Male remating and female fitness in the wolf spider Pardosa astrigera: the role of male mating history

Although the effects of male mating history on female reproductive output and longevity have been studied in insects, few such studies have been carried out in spiders. In a mating system in which females are monandrous while males are polygynous, females may incur the risk by mating with successful males that have experienced consecutive matings and suffer from the possible depletion of sperm and/or associated ejaculates. Here, we examine the effects of male mating history on male courtship and copulation duration, female reproductive fitness, and female adult longevity of the wolf spider, Pardosa astrigera. Results indicated that male mating frequency had little effect on their subsequent copulation success, and of 35 males tested, about half of the males were able to copulate with five virgin females successively at an interval of 24 h. Male mating history had little effect on their courtship duration. However, male mating history significantly affected male copulation duration, female adult longevity, and reproductive output. Males that mated more frequently copulated longer and more likely failed to cause their mates to produce a clutch, although there was no significant difference in the number of eggs laid and the number of eggs hatched regardless of the first clutch or the second one. Multiple mating of male P. astrigera resulted in significant reduction in female adult longevity. Our results indicate that monandrous females mating with multiple-mated males may incur substantial fitness costs.     

Costs of courtship and mating in a sexually cannibalistic orb-web spider: female mating strategies and their consequences for males

The costs of courtship and mating may include increased risks of predation, the transmission of pathogens, and a loss of foraging opportunities. Thus, a female's decision to tolerate a courting male will depend upon how these costs offset the benefits of mating, which will depend on her reproductive and nutritional status. While these costs may be similar for mated and unmated females, the benefits of mating will be less for mated than virgin females. However, the cost of lost foraging opportunities may be higher for females with fewer nutritional reserves necessary for forming eggs. We examined how these costs and benefits influence the courtship and mating behaviour of male and female orb-web spiders, Argiope keyserlingi. In the field, females on webs that also contained a courting male intercepted fewer prey items per hour than females on webs without a male. In the laboratory, the presence of a courting male at the hub also attracted mantid predators to the web, increasing the risk of predation for both male and female. Staged mating experiments in the laboratory revealed that the frequency of female attacks and pre-copulatory cannibalism was greater among mated than virgin females. Feeding history did not affect aggression in virgin females but, among mated females, food-deprived spiders attacked and cannibalized males more frequently than sated females and only the latter ever remated. These differences in female behaviour influenced male mating strategies. Choice experiments demonstrated that males preferred to venture onto the silk threads of virgin rather than those of mated females. Similar patterns of mate selectivity were observed in the field; females with narrow abdomens attracted more males to the webs than females with broad abdomens, and copulations were observed more frequently among females with narrow abdomens. These smaller females are likely to be virgins that have recently molted. Males that preferentially mate with virgin females will not only avoid potentially fatal attacks but also obtain, on average, a higher fertilization success.     

Loss of mating opportunities influences refuge use in the Iberian rock lizard,<Emphasis Type="Italic"> Lacerta monticola</Emphasis>

Because time spent in refuge may be costly if prey lose opportunities to forage, fight, or mate, prey allow predators to approach closer before beginning to flee when opportunity costs are high. Because the same opportunity costs may apply to refuge use as to escape, prey should make similar trade-offs between risk of emerging and cost of remaining in refuge. In the Iberian rock lizard, Lacerta monticola, we studied the effects of sex, reproductive season, speed of predator approach, and potential loss of mating opportunities on time spent in refuge following simulated predatory attacks. Lizards of both sexes adjusted refuge use to the level of risk by spending more time in refuge when approached rapidly than slowly. Females remained in refuge for equal times in the mating and postreproductive seasons, but males emerged sooner during the mating season, suggesting adjustment to a cost of lost opportunity to search for mates during the mating season. When a tethered female was nearby, males emerged from refuge earlier than if no female was present, indicating a trade-off between risk and mating opportunity. Approach speed affected emergence time when females were absent, but not when a female was present. Approach speed did not affect the probability that, after emerging, a male would return to court the female. For males that courted females intensely (bit them) before entering refuge, approach speed did not affect latency to emerge, but males that courted less intensely emerged sooner if approached slowly than rapidly. These findings show that males adjust the length of time spent in refuge to both risk of predation and reproductive cost of refuge use.Communicated by A. Mathis     

Female behaviour mediates male courtship under predation risk in the guppy (Poecilia reticulata)

Previous work has shown that under elevated predation risk, male guppies (Poecilia reticulata) switch from courtship to less conspicuous coercive mating attempts. This behavioural transition is traditionally interpreted as a 'risk-sensitive' response that makes males less conspicuous to predators. However, predation risk leads to behavioural changes (such as schooling and predator inspection) in females that may result in coercive mating attempts being more profitable in high-risk situations. Here, we tested the hypothesis that the switch to coercive mating by male guppies in high-risk situations is mediated by adjustments in female behaviour, rather than directly by the predator. We used replicate models resembling a known guppy predator to simulate predation risk in wild-caught guppies from a high-predation population in Trinidad. Our results revealed that males performed proportionately more coercive mating attempts when presented with a female that had been exposed previously to a model predator compared to when males were paired with non-exposed females. Total mating activity (combined rates of courtship and forced mating attempts) did not differ significantly among the two treatment groups, indicating that overall mating activity is unaffected by predation risk. Importantly, when we subsequently presented both sexes concurrently with a predator model, total mating activity and the proportion of forced mating attempts remained unchanged in the high-risk treatment. Taken together, these results indicate that the transition from courtship to forced mating attempts under elevated predation risk is mediated by changes in female behaviour, which we suggest may favour the use of coercive mating under high predation risk.     

Experience affects mating behavior, but does not impact parental reproductive allocation in a lizard

Investment into reproduction is influenced by multiple factors and varies substantially between males and females. Theory predicts that males should adjust their ejaculate size or quality in response to variation in female experience or phenotypic quality. In addition, sperm investment by males may also be influenced by their own status and experience. Although such adjustments of male ejaculate size can impact reproductive success (via fertilization success), fitness returns from male sperm investment may be influenced (either limited or facilitated) by the level of maternal investment. To test this hypothesis, we conducted an experiment that simultaneously evaluated the effect of paternal and maternal experience (which incorporates mating status, age, body size, and other related variables) on paternal sperm investment and maternal reproductive allocation in the lizard Anolis sagrei. During staged mating trials, experienced males were more likely to copulate with females, but these individuals were less likely to transfer sperm during mating than were naïve individuals. Maternal experience had no impact on these mating behaviors. In contrast to expectations, experience and phenotypic quality (of both sexes) had no impact on male ejaculate size or quality (proportion of live sperm) or on maternal reproductive investment (in terms of egg size and yolk steroids). These findings were intriguing given the mating system and past evidence for differential maternal investment in relation to sire quality in A. sagrei. The results found in this study highlight the complexity of reproductive investment patterns, and we urge caution when applying general conclusions across populations or taxa.     

Predation risk,food deprivation and non-random mating by size in the stream water strider,Aquarius remigis

Summary Non-random mating by size (NRMS) plays a central role in the study of sexual selection and the evolution of mating systems. Theory suggests that NRMS should be influenced by conflicting demands (e.g., predation risk, hunger); few experimental studies, however, have addressed these effects. We used a factorial experiment to examine the influence of predatory green sunfish and food deprivation on NRMS in male and female stream water striders, Aquarius remigis. As predicted by theory, food deprivation decreased the large-male mating advantage. The influence of predation risk, however, went against existing theory; that is, risk increased the large male mating advantage. The degree of large-male mating advantage was negatively related to a measure of the rate of male harassment of females. A behavioral mechanism that can explain these patterns emphasizes the contrasting effects of different competing demands on male harassment rates, female resistance and the role of male size in overcoming female resistance. Females usually resist male mating attempts. Successful mating occurs when males overcome female resistance. If harassment rates (of females by males) are low, larger males have a mating advantage over smaller males perhaps because females resist heavily and thus only larger males can overcome female resistance. If, however, male harassment rates are very high, female resistance might be swamped; mating should then be more random with respect to male size. Food deprivation increases gerrid activity and thus increases harassment rates which should then reduce NRMS. In contrast, risk decreases gerrid activity, thus decreasing harassment rates and increasing NRMS. Females did not show significant NRMS. Females did, however, show a pattern of change in NRMS that is consistent with male choice for larger females. Correspondence to: A. Sih     

Song rate as a signal for nest site quality in blackcaps (Sylvia atricapilla)

In this study we examine male song output as a measure of nest site quality in blackcaps (Sylvia atricapilla). Song rate, breeding success, predation on nests and reaction to playbacks were investigated in individual males. Habitat features determining nest site and song post quality in terms of vegetation cover were compared between successful nests and nests that had suffered predation. We then related song rate of unmated males to habitat factors in territories and nesting sites in order to examine a possible predictor function of blackcap song for habitat quality. Several habitat features are responsible for variation in nesting success. These features also correlate with song rate of unmated males. The study indicates a potential role of song rate in the advertisement of territory quality. Furthermore, the data suggest that females use song rates rather than territory quality in mating decisions. The information females may gain about male quality in relation to territory quality are also discussed.     

Female encounter rate and the calling behavior and mating propensity of male Requena verticalis (Orthoptera: Tettigoniidae)

Mating effort, the energy exerted in finding and persuading a member of the opposite sex to mate, may be influenced by how frequently potential mates are encountered. Specifically, males that frequently encounter females may reduce calling effort and be less eager to mate than males that infrequently encounter females. An experiment was set up to test this hypothesis, using the tettigoniid Requena verticalis. We examined the song structure, calling activity and mating propensity of individual males exposed to one of five different encounter rates with virgin females. Song structure and calling effort were significantly altered by an encounter with a female. After an encounter, males significantly increased chirp rate and decreased variability in interchirp interval. Encounters also stimulated a male to call and to continue to call for up to two hours. The elapsed time since mating affected mating propensity but not calling activity. Mating propensity asymptotically increased to reach a maximum by day 17 since last mating. However, neither the frequency of encounters, nor the number of previous encounters experienced by a male, influenced calling activity or the propensity of a male to mate. The significance of changes in song structure and calling activity following an encounter, and of increasing male mating propensity over time, are discussed. Correspondence to: G.R. Allen     

Role-reversal in katydids: Habitat influences reproductive behaviour (Orthoptera: Tettigoniidae,Metaballus sp.)

Summary Mating behaviour of the katydid Metaballus sp. varies. At sites QS and DB (in 1982) females competed for access to calling males and males chose mates by mating with heavier, more fecund females. At another site (BR) there was no evidence of role-reversal in reproductive behaviour, and males were observed to compete for mates. This species has a large spermatophore, a product of male reproductive glands, that is eaten by the female after mating. Males at the DB site had small reproductive glands. This suggests that some aspect of the QS and DB environments decreases spermatophore production; spermatophores become a limiting resource for females resulting in the reversal in reproductive roles observed at these sites. A field experiment that involved moving individuals from site BR to QS in 1983 determined that mating system was influerced by site (Table 1). At BR, males produced a continuous calling song, a third of the males observed attracted mates, and called for about 30 min before the female arrived; courtship duration was short. Males that were moved from BR to QS encountered a higher density of receptive females as all males attracted females after an average of just 3 min of calling. They changed their behaviour by producing short periodic bursts of song (zipping), and by courting females for long periods of time. The long courtship period may function as as a mate-assessment period for males. The reproductive behaviour of BR males moved to QS differed from that of native QS males only in the length of time spent in copula.     

Bat predation and the evolution of leks in acoustic moths

Theories of lek evolution generally invoke enhanced mating success experienced by males signalling in aggregations. Reduced predation has also been acknowledged as a potential factor driving lek formation, but its role is more ambiguous. Although lekking is a complex behaviour, few empirical studies have investigated the role of both claims. We studied the potential pressures imposed by mating success and predation in an acoustic moth, Achroia grisella, in which males gather in leks and broadcast a calling song attractive to females. We exploited the ability to manipulate the distribution of singing males in laboratory arenas to create different-sized leks and tested female preferences for these aggregations. Because A. grisella are vulnerable to predation by bats while in flight and on the substrate, we also tested the responses of a potential predator, Rhinolophus ferrumequinum, a bat species that feeds on moths, to the experimental leks. We found that the per capita attractiveness of A. grisella males to females rose with increasing lek size. R. ferrumequinum also oriented toward experimental A. grisella leks, but this attraction did not increase at larger leks. Thus, a male’s per capita exposure to predation risk declined as more moths joined the lek. A. grisella males appear to benefit from advertising in larger leks in terms of both increased mate attraction and reduced predation risk. Our results support the idea that multiple factors operating simultaneously may maintain lekking behaviour.     

Experienced males have higher mating success than virgin males despite fitness costs to females

It is generally assumed that mating history has potentially important effects on the mating biology of insects, and differences in mating success of males, in relation to their mating history, have been commonly documented in Lepidoptera. Mating success of male European corn borer Ostrinia nubilalis, in relation to their mating history, and consequent fitness parameters for their female mates, were investigated. Mating experience significantly increased mating success for O. nubilalis males. Mating with experienced males significantly reduced lifetime fecundity and longevity of females. Sperm depletion, transfer of less nutritive substances or more toxic substances may explain the female response to mating with experienced males. Male persistence or some other compensatory mechanism may account for female acceptance of experienced males.     

Male genital length and mating status differentially affect mating behaviour in an earwig

Phenotypic variation in male genitalia may affect copulation behaviour, which can have important fitness consequences for males. Male genitalia commonly possess traits that increase male control over copulation, but in species where females control mating, a poor functional understanding often prevents insight into the processes responsible for such effects. Here, I investigate the effect of male genital length on copulation behaviour in the earwig Euborellia brunneri, where both sexes exhibit extremely elongated genitalia that correspond in shape. This model system is particularly suitable because pairs mate repeatedly and females can limit both the number and duration of copulations. I used both virgin and mated males and females in a double-mating design because longer male genitalia confer benefits in sperm competition. Consistent with a greater predicted male mating effort in mated females, the duration of individual copulations increased, but this traded off against mating frequency as cumulative mating duration remained unchanged. In contrast, male genital length increased both individual and cumulative mating duration, regardless of mating status. This difference suggests that, while males may modify copulation duration in response to mating status, females facultatively adjust mating frequency to prevent mating excessively or express preferences for increased male genital length. Notably, this study demonstrates that male genital phenotypes that are successful in sperm competition also enjoy female-mediated mating benefits.     

Polyandrous mating in treetops: how male competition and female choice interact to determine an unusual carnivore mating system

The diversity of mammalian mating systems is primarily shaped by sex-specific reproductive strategies. In the present study, we explored determinants and consequences of a unique mating system exhibited by fossas (Cryptoprocta ferox), the largest Malagasy carnivore, where females mate polyandrously on traditional mating trees, and males exhibit intrasexual size dimorphism. Males face both contest and scramble competition, and inter-sexual size dimorphism can be pronounced, but its magnitude depends on the male morph. Using a continuous behavioral observation of six estrous females over 4 years, we investigated correlates of male contest competition and female choice based on 316 copulations. Furthermore, we assessed correlates of male scramble competition based on testes size and movement data obtained from GPS tracking. We found that females dominated males regardless of their smaller size and that females actively solicited copulations. Heavy males had highest mating success during the female’s peak mating activity, but were discriminated against afterwards. Female choice and male–male competition thus converged to generate a mating advantage for heavier males. Our results suggest that females actively seek polyandrous matings, presumably for indirect genetic benefits. Since body mass is the major determinant of male mating success and is at the same time dependent on the degree of sociality and associated hunting mode of the respective male morph, a male’s feeding ecology is likely to influence its reproductive tactic. A combination of benefits from female polyandry and the consequences of different subsistence strategies may thus ultimately explain this unusual mating system.     

Song rates of dark-eyed juncos do not increase when females are fertile

Frequency of singing by birds may vary with reproductive stage in ways that reflect variation in the functions of song in intersexual and intrasexual communication. In dark-eyed juncos (Junco hyemalis) high-amplitude song is produced only by males. To investigate the function of this song, we tested whether fertility of females affected singing by their mates or by neighboring males. Using focal observations, song censuses, and radiotracking data, we determined whether song production varied between and among periods when females were fertile and non-fertile. Our findings show that males do not increase song production when their mates are fertile, nor do they increase song production when neighboring females are fertile. These results suggest that male juncos do not signal their intent to defend territories (or mates) more when females are fertile and that they do not use song to advertise to specific potential participants in extra-pair fertilizations. Received: 13 February 1997 / Accepted after revison: 2 May 1997     

Divorce and extrapair mating in female black-capped chickadees (Parus atricapillus): separate strategies with a common target

Patterns of divorce and extrapair mating can provide insights into the targets of female choice in free-living birds. In resident, site-faithful species with continuous partnerships, the better options and the incompatibility hypotheses provide the most likely explanations for divorce. Extrapair mating can be explained by a number of hypotheses often making similar predictions. For example, the good genes and future partnerships hypo- theses predict similar patterns if males with good genes also make the best future partners. By considering both divorce and extrapair mating, it may be possible to distinguish between these comparable hypotheses. We examined natural patterns of divorce and extrapair mating in a long-term study of black-capped chickadees (Parus atricapillus). Out of 144 partnerships over 8 years, we observed 11 divorces and 38 faithful pairs between seasons. Females usually divorced between their first and second breeding seasons for males of higher social rank than their previous partners, had similar reproductive success prior to divorce as females who retained their previous partners, and did not divorce on the basis of previous reproductive success. These results confirm earlier experimental evidence that females divorce for better options. Females who divorced were significantly more likely to have had mixed-paternity broods prior to divorce than females who stayed with their previous partners. There was no evidence that females divorced in favour of previous extrapair partners. These results support the good genes hypothesis for extrapair mating, suggesting that female chickadees use divorce and multiple mating as separate strategies sharing a common target. Received: 4 February 2000 / Revised: 20 July 2000 / Accepted: 4 September 2000     

Male–male competition and alternative male mating tactics influence female behavior and fertility in Japanese medaka (Oryzias latipes)

Intense male–male competition driven by high male density during mating can result in the evolution of alternative mating tactics that increase male fertilization success. The effects of alternative male mating tactics on females can range from increased fertilization and genetic benefits to decreased fertilization and loss of paternal care. However, the influence of male competitive behavior and alternative mating tactics on female behavior and reproductive success has seldom been addressed. In this work, I investigated the occurrence of alternative male mating tactics and their potential influence on female behavior and fertilization success in Japanese medaka (Oryzias latipes). Groups of one, two, or four males competed for access to a female in a repeated-measures experiment. Male density had a significant influence on female reproductive output as a result of a change in competitive mode from contest to scramble competition that coincided with more disruption during mating when more than one male attempted to mate. By contrast, sneaking during mating was beneficial to males, as more than one male sired offspring in most spawnings involving sneaker males. These results suggest that there may be conflict between males and females over mating, such that females are detrimentally affected by the occurrence of alternative mating tactics, whereas males may benefit from sneak mating. The occurrence of conflict between the sexes can be related to ecological factors, such as male density, which cause behavioral change in both males and females.     

Testing the ability of males and females to respond to altered songs in the dueting green lacewing, Chrysoperla plorabunda (Neuroptera: Chrysopidae)

Green lacewings in the carnea group of Chrysoperla engage in species-specific heterosexual duets using low-frequency substrate-borne signals. Within each species, both sexes sing nearly identical songs. Songs are the principal barriers to hybridization between sympatric species in the complex. Here, we investigated the responsiveness of males and females of Chrysoperla plorabunda to synthesized, prerecorded songs that differed from the species mean in the period between repeated volleys of abdominal vibration. We tested 15–16 males and 15–16 females using playbacks of two signals that gradually increased or decreased in volley period, starting at the species mean. We found that (1) duets during courtship are accurate, interactive, and adjustable by each participant; (2) in staged duets, both sexes respond best to song tempos near the mean volley period of their population, but can nonetheless maintain duets with signals of nearly twice, or half, the normal volley period; (3) individuals fine-tune their adjustments to signals of different volley periods by changing their own volley duration and latent period, or less often by inserting extra volleys or skipping every other volley; (4) males are significantly better at matching signals of changing tempo than females; and (5) the range of song responsiveness of C. plorabunda does not overlap the natural range of volley periods found in Chrysoperla adamsi, an acoustically similar sibling species, thus reaffirming strong behavioral isolation. In sum, the precise, almost unbreakable heterosexual duets characteristic of song species of the carnea group result from tight mutual feedback between partners. Effective reproductive isolation between species can be based on song differences alone.     

Tradeoffs between courtship, fighting, and antipredatory behavior by a lizard, Eumeces laticeps

Male fitness in many species depends strongly on social behaviors needed to obtain fertilizations and prevent loss of fertilizations to other males, but courtship, copulation, and fighting may incur increased risk of predation. When demands for reproductive and antipredatory behaviors conflict, fitness may be maximized by accepting some degree of risk to enhance reproductive success. To examine such tradeoffs, I introduced tethered conspecific males or females to adult male broad-headed skinks, Eumeces laticeps, in the field and observed how close they allowed a simulated predator (me) to approach before fleeing, or their latency to approach an introduced female located at different distances from the predator. When conspecific males were introduced, isolated and mate-guarding males initiated agonistic behaviors and permitted closer approach than control males, and mate-guarding males permitted closer approach than isolated males. When females were introduced, both isolated and mate-guarding males courted the introduced females and isolated males permitted closer approach than did mate-guarding males. These results for introduced males and females suggest that increasing risk was accepted when reproductive benefits were greater. Latency for isolated males to approach a conspecific female was greater when the predator was closer to the female, further suggesting sensitivity to predation risk during a reproductive opportunity. Relationships between reproductive and antipredatory behaviors have been studied much less than those between feeding and antipredatory behaviors, but this study indicates that animals balance increased risk of predation with the opportunity to perform several reproductively important behaviors. Received: 5 March 1999 / Received in revised form: 15 July 1999 / Accepted: 25 July 1999     

Costing reproduction: effects of mating opportunity on mating success in male Bicyclus anynana butterflies

It is frequently assumed that males have an almost unlimited reproductive capacity, while access to receptive females is typically limiting. Consequently, sexual selection is expected to favor vigorous courtship behavior in males. If such behavior is associated with non-trivial costs, ample current mating opportunities should be accompanied by a reduction in future mating vigor. To test this hypothesis, three treatments differing in sex ratio were established using the tropical butterfly Bicyclus anynana: 50 males each were housed either with 0, 25, or 50 females. Mating trials involving the competition among males from each treatment for a single virgin female were carried out on days 3, 9, and 15 after allocation to treatments. While there was no difference on day 3, prior mating opportunity clearly reduced mating success on days 9 and 15, being lowest if identical numbers of males and females were housed together. This finding suggests accumulating costs associated with high courtship and/or mating activity. Further, older males were more active and initiated copulation earlier than the younger ones, consistent with the residual reproductive value hypothesis. We found no evidence for a survival or fertility cost of mating.     

Mating system and mating success of the desert spider Agelenopsis aperta

Field studies of the desert spider Agelenopsis aperta revealed a primarily monogamous mating system. However polygyny, polyandry and polygynandry were superimposed upon the primary system, with 9% of the marked males and 11% of the marked females in a field population mating more than once. In the laboratory males commonly mated multiply with fertile offspring resulting, while females were less likely than males to mate multiply. Monogamy under field conditions was enforced by two factors: (1) high travel costs to males, and (2) a significant decline in female receptivity after the first mating. Heavy males were more likely to be accepted by females both in the field, and in female choice experiments conducted in the laboratory. Finally, male weight determined the outcome of male-male agonistic interactions over females. One possible explanation for female choice in this system which lacks male parental investment is that females may be using male size as an indicator of future success of their offspring.