Mate choice plasticity in the field cricket <Emphasis Type="Italic">Teleogryllus oceanicus</Emphasis>: effects of social experience in multiple modalities

Social experience can elicit phenotypically plastic changes in mate choice, but little is known about the degree to which social information from one modality can influence mating decisions based on information from a different modality. I used the field cricket Teleogryllus oceanicus to test whether experience of chemical cues mimicking a high density of sexually mature males causes changes in mate choice based on acoustic signals. T. oceanicus males produce long-range calling songs to attract females for mating, but they also produce waxy, non-volatile hydrocarbons on their cuticle (CHCs) which, when deposited on a substrate, can be detected by females and may provide demographic information. I manipulated female experience of substrate-bound male CHCs and then performed acoustic mate choice trials. When CHCs were present on the substrate during trials, females showed greater motivation to respond to male calling song. This effect diminished with repeated exposure to male songs, demonstrating that the importance of olfactory cues in altering acoustic mate choice decreased with increasing exposure to acoustic signals. However, the temporal nature of CHC experience mattered: previous experience of CHCs did not alter subsequent female choice for male calling song traits. Exposure to male song increased the threshold of mate acceptance over time, and individuals varied considerably in overall levels of responsiveness. Taken together, the results demonstrate that mate choice is dependent on social context mediated by multiple modalities in T. oceanicus, but they do not support the idea that prior experience of social cues in one modality necessarily influences later mating decisions based on other signalling modalities.     

No male agonistic experience effect on pre-copulatory mate choice in female earwigs

Mating with dominant males may confer considerable benefits, but also incur significant costs, hence intrasexual competitiveness is a likely target of mate choice. In addition to established modes of mate assessment, females may use cues or signals associated with agonistic experience effects to assess the relative competiveness of males. Experience effects, where the outcome of a fight increases the likelihood of a similar outcome in subsequent fights, may result from an animal’s altered state after conflict, but can also arise from strategic rival use of information perceived about this altered state. While females may similarly use this information in mate choice decisions, this potential consequence of male–male conflict has largely been neglected. Here, we investigate the effects of experience on subsequent agonistic performance in the earwig Euborellia brunneri by imposing winning or losing experiences on males and rematching them with naïve, size-matched rivals. We reveal a strong loser effect in subsequent fights, with nearly all previous “losers” losing against new rivals. In contrast, we found no equivalent winner effect, with previous “winners” exhibiting no increased likelihood of winning. We subsequently test whether the effects of male agonistic experience extend to pre-copulatory female mate choice. We show that females, when allowed to choose between naïve males and “winners” or “losers”, do not discriminate between males based on their recent agonistic experience. Therefore, while fighting history can play an important role in male–male interactions, females may not attend to this information.     

Age-related female mating decisions are condition dependent in wolf spiders

Female mating behaviors are known to be sensitive to a variety of individual factors both external and internal to a female; however, mating decisions are likely due to a suite of interacting factors. By independently manipulating female and male diet in the wolf spider Rabidosa rabida and testing females across age groups, we demonstrate that, in addition to its independent effect, female nutritional condition interacts with female age to influence female mating behavior. Overall, high-quantity diet (HD) females were more likely to mate than low-quantity diet (LD) females. Within the LD females, older individuals were more likely to mate than younger individuals, while within HD females, mating probabilities were equal across females of different age classes. With respect to mate choice, only female age influenced the likelihood of mating based on male diet. Young females were choosier as they were more likely to mate with HD males than LD males; in contrast, older females were equally likely to copulate with males of each diet treatment. In addition, the likelihood of pre-sexual cannibalism was dependent on both female and male diet. High-quantity diet females were more likely to cannibalize than LD females, and attacks were directed towards LD males most often. We discuss our results in terms of costs versus benefits of female mate choice.     

Responses of female lizards,<Emphasis Type="Italic"> Lacerta monticola</Emphasis>, to males' chemical cues reflect their mating preference for older males

Female mate choice based on visual traits appears to be rare in lizards. Field observations suggest that females of the lizard Lacerta monticola preferred to mate with larger/older males. Although older males are usually green and larger, and younger males brown and smaller, there is some overlap in size and coloration between age classes. Thus, visual cues may not always be reliable indicators of a male's age. We hypothesized that female mate-choice preferences may be based on males' pheromones, which might transmit information about characteristics such as age. In a laboratory experiment, we analyzed the effect of age of males on attractiveness of their scents to females. When we offered scents of two males of different age, females associated preferentially with scents of older males. This suggested that females were able to assess the age of males by chemical signals alone, and that females preferred to be in areas scent-marked by older males. Thus, females may increase their opportunities to mate with males of high quality, or may avoid harassment by sneaking young males. This result agreed with field observations on females mating with old males, and rejection of advances by young males. Our results also suggested that female preference for older males may depend on their own body size. Large females showed a strong preference for older males, whereas smaller females were not so selective. This, together with males' preference for large females, might lead to size-assortative matings. We suggest that the quality and/or quantity of male pheromones could communicate to the female heritable male genetic quality (i.e. age) and thereby serve as the basis of adaptive female choice in lizards.Communicated by W.E. Cooper     

Indirect fitness benefits are not related to male dominance in a killifish

Recent theoretical and empirical studies have shown that male dominance is often at odds with female mate preference and that indirect (genetic) benefits of mate choice may not be related to male dominance. We tested whether female preference corresponded to male dominance and whether mating with dominant males conveyed benefits to offspring fitness in a small freshwater fish, the African annual killifish Nothobranchius korthausae (Cyprinodontiformes), a species without parental care. The experimental design used controlled for the effect of male age, possibility of sperm and egg depletion, and accounted for a potential that females express their preference through maternal effects by manipulation of egg mass during ovulation. By sequentially mating females with males of known dominance, we found that female N. korthausae showed no mate preference in terms of egg numbers deposited with respect to male dominance or body size and no congruent mate preference to specific males was detected. However, males sired offspring with consistently higher hatching success and the effect was repeatable across individual females. Thus, some males provided females with indirect benefits related to additive genetic quality (“good genes”) and expressed via increased hatching rate, but this benefit was not related to male dominance status or body size.     

Rapid onset of mate quality assessment via chemical signals in a woodland salamander (<Emphasis Type="Italic">Plethodon cinereus</Emphasis>)

In the competition to acquire mates, environmental factors can be important in determining the relative quality of an individual. These aspects of quality are often conveyed through signals used for mate assessment by the most energetically invested sex. In red-backed salamanders, Plethodon cinereus, chemical signals communicate a surprising amount of information, much of which is dependent on the current condition of the sender. One such condition-dependent aspect of information conveyed via chemicals is the quality of the sender’s diet, but it is unclear as to whether this information is actively advertised by the sender (i.e., a signal) or simply inferred from food-derived odors (i.e., a cue). The amount of time on different diets required for changes in signaling is also unknown. I examined how quickly gravid female salamanders could detect a difference between the scents of males on high- vs. low-quality diets without fecal cues. The amount of protein present in two known signaling glands (the mental and postcloacal glands) was also measured after experimental feeding. Gravid females were able to infer the differences in male diet quality after only 1 week. Females also responded to the male scents more quickly after differential feeding had begun. High-quality males had significantly more protein present in both signaling glands than low-quality males. This scenario highlights the ongoing interplay between the quality of an individual and its environment, with males actually advertising the status of this relationship as an honest signal for mate assessment.     

Antioxidant status, flight performance and sexual signalling in wild-type parrots

During mate choice, individuals are predicted to assess traits that honestly signal the quality of potential partners. Locomotor capacity may be such a trait, potentially signalling condition and ability to resist oxidative damage. In this study, we experimentally manipulated nutritional status: Male wild-type budgerigars, imported from Australia, were provided with either an enhanced (EQ) or reduced quality (RQ) diet varying in vitamin (particularly retinol and α-tocopherol) and mineral levels. Then, we assessed whether this influenced locomotor capacity, i.e. escape flight performance, and sexual attractiveness in male budgerigars Melopsittacus undulatus. Males in the EQ group showed significantly greater total antioxidant capacity and higher blood plasma concentrations of the dietary antioxidants retinol and α-tocopherol, but not carotenoids, than the RQ group. Over 8 weeks of flight training, males on the EQ diet showed significantly greater improvement on the most strenuous flight test than RQ males. In mate choice trials, females preferred EQ over RQ males. EQ males that were relatively fast in escape flight trials were more strongly preferred in the mate preference arena than their RQ competitors. Interestingly, males with high plasma carotenoid levels flew slower and were less attractive than males with low carotenoid levels. This might indicate that carotenoids are not effective antioxidants in birds. Overall, our results show that dietary-derived antioxidants can influence sexual attractiveness and other fitness-related traits through multiple pathways. Locomotor capacity appears to be an honest signal of male condition in birds.     

Modulation of male aggressiveness through different communication pathways

Communication plays a large role in resource competition, especially for potential mates, and is used by members of the competing sex to assess each other, and simultaneously to evaluate the other sex, which may be advertising its status. To assess the effects of female advertisement on male aggression, males of the decapod Aegla were paired according to body and armament size. Males were left to interact in five different treatments: with receptive females that could use both chemical and visual cues, non-receptive females that could use both types of cues, receptive females that could use only one cue, or no female in the aquarium. Fight duration, time spent in the most aggressive acts, latency period, number of antennal whips/fight duration, and time spent near the female were analyzed. The males had shorter and less intense confrontations when there was a receptive female that could signal with at least one modality. Winning males spent significantly more time near the receptive female only when both chemical and visual cues were present, when compared to the other treatments. The low level of aggression shown by the males may be related to information asymmetry due to the female’s choice: only the preferred male would receive information from the female, or males could compete for other resources that attract females. However, male aggression was modified by the presence of female chemical cues, whereas mate guarding was initiated only when both chemical and visual cues were present. Hence, male aggression can be downregulated by female receptivity.     

Femoral secretions and copulatory plugs convey chemical information about male identity and dominance status in Iberian rock lizards (Lacerta monticola)

Socially dominant males often signal their status to rival males and/or females. We tested the hypotheses that Lacerta monticola femoral gland secretions and copulatory plugs convey chemical information about male identity and dominance status. We estimated male dominance status by staging male–male agonistic encounters in a neutral arena. We then conducted two experiments to compare male tongue-flick behavior toward chemical stimuli consisting of cotton swabs bearing (1) deionized water (control), the lizard’s own femoral secretions, and the femoral secretions of another male and (2) phosphate-buffered saline solution (control), the lizard’s own plug products, and the plug products of another male. Results indicate that males discriminated their own femoral secretions and plugs from those of other males. They also discriminated morphological attributes of other males that were associated with dominance status based on chemical cues arising from femoral secretions and discriminated the dominance status of other males based on chemical cues arising from the plugs. Femoral secretions that convey information about male identity and dominance status may be hypothesized to function in the establishment of L. monticola dominance hierarchies through scent-marking of territories. We suggest that copulatory plugs and femoral secretions may allow males to scent-mark the female body and postulate that this behavior may influence male and female reproductive decisions under selective pressures of sperm competition.     

Male moths undertake both pre- and in-copulation mate choice based on female age and weight

Variation in reproductive potential usually occurs among individuals of both sexes; for example, some individuals may carry more gametes or be able to continue to generate more gametes in their reproductive life than others. Therefore, to maximize their reproductive success, both sexes are expected to show adaptations for mate choice. However, most authors concentrated on how females choose their mates with the belief that females invest more in reproduction than males, and males are generally eager to pair with any female. In this paper, we report our work on a polygamous moth, Ephestia kuehniella Zeller (Lepidoptera: Pyralidae), with special emphasis on male pre- and in-copulation mate choice in relation to the reproductive quality and re-copulation potential of females. We show that in E. kuehniella, younger and heavier females have significantly higher reproductive value than older and lighter ones, and male sperm supply significantly decreases over successive copulations. Males exercise pre-copulation mate choice by selecting females with higher reproductive potential for copulation and in-copulation mate choice by allocating more sperm to females with higher reproductive potential. However, high-quality females are more likely to re-copulate than low-quality ones, and allocation of more sperm to females by males does not increase female reproductive outputs. It is suggested that the allocation of more sperm to high-quality females functions to increase sperm competitiveness against rivals.     

The assessment of female reproductive state during courtship and scramble competition in the fiddler crab, Uca paradussumieri

Male fiddler crabs, Uca paradussumieri, mate underground during a 4- to 7-day period each full and new moon. As soon as the tide recedes, males enter the burrows of females that will ovulate the following day ('pre-ovigerous' females). Males copulate with and guard these females until they ovulate. When interrupted by an intruding male, the first male to reach the female is usually able to defend her and successfully mate with her. In fiddler crabs, females mate multiply and there is last male sperm precedence. Before each semi-lunar mating period, male U. paradussumieri were more likely to court females with whom they would later mate than other nearby females with whom they did not mate. This suggests that males collect information on female reproductive state prior to the females becoming ovigerous. In this species, aggression was common between males that courted the same female. When previously courted females were approached by other males, the initial courter attempted to forcefully disrupt the courtship. This behavior may allow males the exclusive use of information on female reproductive condition. It also suggests a type of scramble competition between males over females. Furthermore, it indicates that males are able to locate receptive females prior to their becoming ovigerous. The shorter guarding period observed in this species, as compared with other fiddler crabs, is caused by females rejecting longer guarding periods. Male ability to assess female reproductive status may therefore be advantageous because it increases male mating success within a scramble type of competitive polygyny.     

Safe betting: males help dull females only when they raise high-quality offspring

In evolutionary biology, whether parents should enhance or reduce parental care according to mate ornamentation is a subject of great debate. However, the evolution of female ornaments can shed light on this question. In theory, female ornamentation should be traded off against fecundity and thus cannot be wholly informative to males without a direct indication of fecundity. Hence, direct cues of offspring quality should affect the relationship between male investment and female ornamentation. Under this hypothesis, we manipulated two direct cues of offspring quality (egg size and color) after first egg laying in the blue-footed booby and registered male incubation patterns. In this species, foot color is a dynamic signal of current condition and in females is traded off with egg size. We found that males spent more time incubating when paired with dull females but only in nests with large eggs. Males also spent less time incubating small dull eggs. Results indicate that egg size, a direct cue of reproductive value, affected the relationship between male effort and female ornamentation. Males may be willing to help females that have invested in offspring at the expense of ornamentation, which suggests compensation when females are in low condition. Another possibility is that males relax their effort when paired with highly ornamented and fecund females because they have high parenting abilities. Our findings suggest that the information conveyed by female ornaments may depend on direct cues of fecundity. Results also highlight that parental decisions are complex, modulated by a combination of information sources.     

Asexuals looking for sex: conflict between species and mate-quality recognition in sailfin mollies (Poecilia latipinna)

When two closely related species are sympatric the process of species recognition (identifying conspecifics) and mate-quality recognition (increased fitness benefits) can yield a conflict when heterospecifics resemble high-quality conspecifics. Conflict in species versus mate-quality recognition may serve as a possible mechanism for the persistence of unisexual, gynogenetic Amazon mollies (Poecilia formosa). Amazon mollies require sperm from closely related species (e.g., sailfin mollies, P. latipinna) to start embryogenesis but inheritance is strictly maternal. When choosing mates, male sailfin mollies from populations sympatric with Amazon mollies may rely on traits indicating species identity rather than those indicating mate quality. Conversely, males from allopatric populations may rely more on traits indicating mate quality. Previous work has found that male sailfin mollies in sympatry exhibit a significantly greater mating preference for female sailfin mollies over Amazon mollies compared to males in allopatry. In addition, male sailfin mollies prefer to associate with and produce more sperm in the presence of larger conspecific females, which are more fecund. We hypothesized that male sailfin mollies experience a conflict in species recognition and mate-quality recognition in the presence of Amazon mollies that are relatively larger than female sailfin mollies. To test this hypothesis, we paired males from sympatric and allopatric populations with a larger Amazon molly and a smaller female sailfin molly. We scored the number of mating attempts that males directed to conspecific and heterospecific females. Males in most sympatric and allopatric populations demonstrate no clear preference for conspecifics. In addition, we found some evidence for a difference in mating preference between allopatric and sympatric populations with males from allopatry showing a greater heterospecific mate preference. These results indicate a conflict between species and mate-quality recognition. In sympatry this conflict may contribute to the persistence of gynogenetic Amazon mollies.     

The lexicon of love: what cues cause size-assortative courtship by male garter snakes?

Cues that females use to select potential mates have attracted substantial research effort, but the criteria for male mate choice remain very poorly known. Red-sided garter snakes ( Thamnophis sirtalis parietalis) court and mate in large aggregations around overwintering dens in southern Manitoba, Canada. Both courtship and mating are size-assortative: small male snakes court small as well as large females, whereas larger males court only large females. This system provides a unique opportunity to assess the cues that males use in selecting mates, and in particular the mechanisms that generate a size-related shift in mate preference. Experiments in which we manipulated body sizes and scents showed that both vision and scent (sex pheromones) were important. Large males directed intense courtship only when the stimulus provided both visual and chemical (skin lipid) evidence of large body size. Small males were much less discriminating in both respects. Thus, size-assortative mating in this system is generated not by larger males excluding their smaller rivals from the largest females (as has been reported in other reptile species), but by a size-related shift in the visual and pheromonal cues that elicit courtship. Males of some species may thus show complex patterns of mate choice, with the cues that stimulate courtship differing even among males within a single population based on traits such as age or body size.     

Post-copulatory aggression toward their mates by males of the rove beetle Leistotrophus versicolor (Coleoptera: Staphylinidae)

Summary Rove beetles (Leistotrophus versicolor) forage and mate at dung and carrion in the riparian forest of northwestern Costa Rica. After copulating, males often launch a post-copulatory attack against their recent partner. We test four hypotheses on the adaptive value of male behavior: (1) The sperm competition hypothesis proposes that the behavior may be the functional equivalent of mate-guarding, (2) the sperm-transfer signal hypothesis states that males bite their mates after copulating to signal that they have successfully passed sperm, (3) the feeding competition hypothesis argues that male aggression toward mates occurs to drive away competitors for fly prey, and (4) the redirected aggression hypothesis is that male attacks after mating occur when threatened males redirect their aggression onto their partners. Only the sperm competition hypothesis withstands testing. As required by this hypothesis, females are usually receptive while at dung, and will mate with more than one male in a morning. In addition, males are more likely to attack a mate when they have fought earlier in the day with other males, an indicator of the presence of rival males and the risk of sperm competition. Contrary to the sperm-transfer signal hypothesis (2), biting of mates does not occur after nearly 40% of all copulations; it seems unlikely that mating males so often fail to transfer sperm. Whether males have fed or not prior to mating has no effect on the probability of post-copulatory attack, a result that contradicts the food competition hypothesis (3). Finally, the occurrence of attacks by males on females in the absence of an immediate threat from a rival argues against the redirected aggression hypothesis (4).     

Cave molly females (Poecilia mexicana, Poeciliidae, Teleostei) like well-fed males

We examined the preference of Atlantic molly females (Poecilia mexicana) to associate with a well-fed or a starved male in simultaneous choice tests. Females from three different populations were tested in three treatments: (1) the females could choose on the basis of multiple cues from the males (visual plus non-visual); (2) only non-visual cues could be perceived in darkness, (3) only visual cues were presented. The three tested populations differ clearly in their ecology: one population occurs in a typical river habitat, the second one in a milky sulfur creek outside a cave, and the third population occurs in a cave habitat (cave molly). In the river-dwelling population, females never showed a preference. In the population from the sulfur creek, females preferred to associate with the well-nourished male when visual cues from the males were available. Only cave molly females exhibited a strong preference for well-nourished males in all treatments. A morphological comparison demonstrated that wild-caught males from river habitats are typically in a good nutritional state. In the sulfur creek, males showed signs of starvation. Cave molly males were in an even worse nutritional state. In the cave population, saturated males probably indicate high fitness, thereby driving the evolution of the preference for good male nutritional state.Communicated by K. Lindström     

Sperm competition and evidence of sperm fertilization patterns in the carrion ball-roller beetle Canthon cyanellus cyanellus LeConte (Scarabaeidae: Scarabaeinae)

Field observation and laboratory experiments were conducted to investigate sperm competition and patterns of sperm fertilization under different experimental conditions in the carrion ball-roller beetle Canthon cyanellus cyanellus. Males in nature can mate with females whose spermathecae contains fertile sperm from other males. Sperm precedence was investigated using a visible genetic marker. The progeny of red (homozygous recessive) virgin females mated once with a red male and afterward, once with a green beetle (homozygous dominant) and vice versa, revealed that regardless of its color, the last male to mate fertilized c.a. 50% of the eggs, suggesting strong sperm competition. Males were able to achieve higher levels of paternity (more than 80%) when mated ad libitum with previously mated females, although large amount of variance in paternity does not exclude the possibility of first male sperm precedence or female cryptic choice. These results suggest that repeated mating and sperm replacement are the mechanisms by which last males achieve sperm precedence.     

Mate preferences by male guppies (Poecilia reticulata) in relation to the risk of sperm competition

We investigated male mate preferences in relation to the perceived risk of sperm competition in the guppy (Poecilia reticulata), a freshwater fish with a promiscuous mating system. Our laboratory experiments revealed that male mate choice behaviour is not influenced by the presence of rival males that are merely in close proximity to a potential mate, as there was no significant difference in the amount of time that males spent with females that were recently either alone or in close proximity to four rival males. Male mate choice behaviour was, however, strongly influenced by the presence of rival males in a second experiment, where those rivals were permitted to copulate with one of the females. In that situation, males spent significantly more time with, and directed significantly more sigmoid courtship displays toward, females that they had recently seen alone compared to females they had seen receiving forced copulations from up to four rival males. Our results therefore indicate that male guppies are sensitive to the risk of sperm competition and alter their mate choice behaviour in an adaptive fashion.Communicated by K. Lindström     

Some constraints on reproduction for male bushcrickets,Requena verticalis (Orthoptera : Tettigoniidae) diet,size and parasite load

Summary Reproduction for male bushcrickets is energetically expensive. Male Requena verticalis invest 70% of their daily energy reserves in calling to attract a female and providing her with a nutritious spermatophore. Males are thereby likely to be constrained in their mating frequency. I investigated constraints on reproduction imposed by body size and the levels of a protozoan gut parasite when males were fed diets that differed in their nutritional value. Males suffered a cost of reproduction in terms of an increasing interval between matings that was independent of diet and parasitic infection. After three successive matings, males decreased the magnitude of investment in courtship feeding when fed a diet poor in protein. Furthermore, these males suffered a reduction in the number of times they were capable of mating relative to males fed a diet rich in protein. Male size constrained mating frequency on both rich and poor diets; small males were able to mate less frequently than large males. There was an interaction between the effects of diet and parasitic infection on male mating frequency. Heavily infected males mated less frequently than uninfected individuals when fed the poor diet. However, males fed the rich diet were able to overcome the constraints imposed by parasitic infection. Reproductive constraints are discussed in relation to the costs of reproduction and their effects on courtship roles.     

Inbreeding avoidance in a poeciliid fish (Heterandria formosa)

Polyandry is hypothesized to give females an opportunity to avoid inbreeding through postcopulatory selection mechanisms if precopulatory inbreeding avoidance is not possible, for example, because of forced matings. Here, we report a postcopulatory, prefertilization, inbreeding avoidance mechanism in the least killifish, Heterandria formosa, a species in which males can force matings. Females had 50% less sperm in their oviducts and ovarian cavities after mating with a sibling compared to a mating with a nonsibling male. Neither sex showed inbreeding avoidance in a dichotomous precopulatory mate choice test or during mating trials. Females in this species invest substantially in each offspring after fertilization (matrotrophy), whereas males invest little more than sperm. Based on theory, females should therefore be more likely than males to avoid inbreeding in this species. We suggest that females do this by reducing the amount of sibling sperm in their reproductive system. However, the possibility that males invested more sperm to nonsiblings could not be ruled out. In a fertilization success experiment, the first male to mate with a female sired all the offspring in most cases, even if it was a sibling. However, large females were more likely to carry offspring of multiple males. Possibly female sperm storage sites were filled by the first male, and only large females had space for the second male's sperm.