Re‐Feeding Food‐Deprived Male Meadow Voles Affects the Sperm Allocation of Their Rival Males

An individual's nutritional status affects the manner in which same‐ and opposite‐sex conspecifics respond to that individual, which may affect their fitness. Male meadow voles, Microtus pennsylvanicus, increase their sperm allocation if they encounter the scent mark of an unfamiliar male that is not nutritionally challenged. If, however, the scent mark comes from a male that has been food deprived for 24 h, stud male voles do not increase their sperm allocation. Food‐deprived males may be viewed as being lower quality and a reduced risk of sperm competition by rival males. We hypothesized that stud males in promiscuous mating systems tailor their sperm allocations depending on whether rival males have been food deprived and then re‐fed. We predicted that newly re‐fed males will be considered a strong risk of sperm competition because of the potentially high fitness and survival costs associated with food deprivation in males, and that they will cause stud males to increase their sperm allocation. Our results, however, showed that the recovery period from 24 h of food deprivation was a relatively slow process. It took between 96 and 336 h of re‐feeding male scent donors that were food deprived for 24 h to induce stud males to increase their sperm allocation to levels comparable to when scent donors were not food deprived. Stud male voles may be conserving the amount of sperm allocated until the male scent donors have recovered from food deprivation and subsequent re‐feeding.     

Seismic Signaling is Crucial for Female Mate Choice in a Multimodal Signaling Wolf Spider

Complex courtship signals can be dissected into distinct components that can either function independently or via interactions with one another. Male Rabidosa rabida wolf spiders use courtship displays that couple a seismic signal with the waving of an ornamented foreleg. While previous studies suggest that female R. rabida exhibit mate choice and that both the seismic and visual modalities are important in mating interactions, it remains unclear how variation in each component influences female mating decisions. To investigate this, we ran two separate experiments in which we manipulated (1) male diets, to induce variation in the seismic courtship signal, and (2) male foreleg color, to artificially induce variation in visual foreleg ornamentation. To determine the influence of variation in each component independently, females were paired with males in environments that allowed the detection of only the manipulated signal component (e.g. seismic signal only and visual signal only). Variability in the seismic signal alone influenced female mate choice, but variability in visual ornamentation alone did not. In a third experiment, we manipulated foreleg color and allowed it to interact with the seismic signal to determine whether inter‐signal interactions influence female mating decisions. When females were able to detect both signal components, variation in visual ornamentation did influence mate choice – females preferred ornamented males. Together, these results suggest that the seismic signal of male R. rabida is integral for female mate choice and that the components of the courtship display interact to influence female mating decisions.     

Discriminating Males and Unpredictable Females: Males Bias Sperm Allocation in Favor of Virgin Females

When both sexes mate with multiple partners, theory predicts that males should adjust their investment in ejaculates in response to the risk and/or intensity of sperm competition. Here, we demonstrate that, in the harlequin beetle riding pseudoscorpion, Cordylochernes scorpioides, males use cues deposited on females by previous males to distinguish between virgin, once‐mated, and multiply‐mated females and adjust sperm allocation accordingly. Sperm number declined in direct proportion to the number of previous males, with virgin females receiving nearly three times more sperm than females exposed to three previous males. Given the lack of first‐male sperm precedence in C. scorpioides, this pattern is not consistent with current sperm competition models and appears best explained by a significant risk of wasting ejaculates on deceitful, mated females. In C. scorpioides, males transfer sperm indirectly to females via a stalked spermatophore deposited on the substrate. Mated females often feign sexual receptivity and cooperate throughout mating, only to reject the sperm packet produced by the male. While indirect sperm transfer facilitates a high level of female deceit and control, females of many species are able to influence the number and fate of sperm transferred during copulation and are likely to conceal their sexual unreceptivity to minimize male retaliation. If males cannot accurately assess female receptivity, increased risk of sperm rejection by mated females could outweigh the risk of sperm competition and favor greater sperm allocation to virgin females.     

Assessment of Rival Males through the Use of Multiple Sensory Cues in the Fruitfly Drosophila pseudoobscura

Environments vary stochastically, and animals need to behave in ways that best fit the conditions in which they find themselves. The social environment is particularly variable, and responding appropriately to it can be vital for an animal’s success. However, cues of social environment are not always reliable, and animals may need to balance accuracy against the risk of failing to respond if local conditions or interfering signals prevent them detecting a cue. Recent work has shown that many male Drosophila fruit flies respond to the presence of rival males, and that these responses increase their success in acquiring mates and fathering offspring. In Drosophila melanogaster males detect rivals using auditory, tactile and olfactory cues. However, males fail to respond to rivals if any two of these senses are not functioning: a single cue is not enough to produce a response. Here we examined cue use in the detection of rival males in a distantly related Drosophila species, D. pseudoobscura, where auditory, olfactory, tactile and visual cues were manipulated to assess the importance of each sensory cue singly and in combination. In contrast to D. melanogaster, male D. pseudoobscura require intact olfactory and tactile cues to respond to rivals. Visual cues were not important for detecting rival D. pseudoobscura, while results on auditory cues appeared puzzling. This difference in cue use in two species in the same genus suggests that cue use is evolutionarily labile, and may evolve in response to ecological or life history differences between species.     

Context‐ and State‐Dependent Male Mate Choice in a Sexually Cannibalistic Spider

High male mating investment may favor selection on male mate choice particularly if females vary in quality. Terminal investment strategies constitute a maximal mating effort and have evolved independently in the absence of paternal investment in several spider taxa including the genus Argiope. To test for male mate preferences in the above context, we used the sexually cannibalistic spider A. bruennichi. We varied male state (mating status and post‐maturation age) as well as the competitive context and quantified male mate choice decisions between females of different states and developmental stages in binary choice tests. We found an overall adaptive preference for the virgin against the mated female regardless of male mating state. Furthermore, we demonstrated that older males paid more attention to female fecundity‐related traits than to mating status. In a second set of experiments, we offered males a choice between a virgin and a subadult female and varied the competitive context which had no effect on male decisions. Curiously, a preference for the virgin adult female was only apparent after exclusion of females that matured <3 d prior to the test. Repeated tests of males supported the hypothesis that males do not distinguish between a freshly matured virgin female and a subadult female. Our results show that male spiders execute mate choice based on information collected from female silk strands and that they integrate their own state into mating decisions.     

Searching Responses of a Hunting Spider to Cues Associated with Lepidopteran Eggs

Cheiracanthium inclusum (Hentz), a hunting spider, feeds on eggs of Helicoverpa zea (Boddie) and other moths. This study investigated whether C. inclusum can use chemical compounds present in H. zea scales and egg residues as kairomones to find these non-motile prey items. In a series of no- choice tests, spiders were presented with a piece of florist paper containing scales alone, scales + egg residues, or untreated controls. Next, spiders were presented with solvent extracts of either scales or eggs. Polar and non-polar solvents were used in the extractions. Contact with scales alone, scales + egg residues, and non-polar solvent extracts of both scales and eggs resulted in retention and/or induction of local searching behavior. Extracts made with polar solvents induced no apparent response, indicating that the chemostimulatory compounds are lipophilic. These results show that C. inclusum responds to kairomones left by ovipositing H. zea and use these chemical cues to detect and locate H. zea eggs.     

Aversive Responses of Female Mice to the Odors of Parasitized Males: Neuromodulatory Mechanisms and Implications for Mate Choice

The detection and avoidance of parasitized conspecifics is proposed to have important consequences for the behavior of animals, especially as related to mate choice. A reduction in pain sensitivity (i.e. analgesia) is a major correlate of exposure to real or potential danger and threatening stimuli, facilitating the expression of various active (e.g. fleeing) and passive (e.g. immobilization) defense responses. The present study examined pain sensitivity (latency of a foot-lifting response to 50 ° C thermal surface) of female mice, Mus musculus, that were exposed to the urine and other odor secretions of male mice subclinically infected with the naturally occurring, enteric, sporozoan parasite, Eimeria vermiformis. A 30-min exposure to the odors of a parasitized male induced an analgesia in the female mice that was found to be mediated by the increased activity of endogenous opioid peptide systems. A brief 1-min exposure to the male odors induced a shorter duration and lower amplitude analgesia of a non-opioid (serotonergic) nature. Maximum analgesic responses were induced by the odors of pre-infective [5 days post-infection (PI)] and infective (day 10 PI) males, with significantly lower responses elicited by the odors of post-infective (day 17 PI) male mice. Exposure to the odors of unparasitized males had no significant effects on the pain sensitivity of female mice. These results indicate that female mice can distinguish between the odors of parasitized and non-parasitized male mice, and find the odors of parasitized males threatening and/or stressful. These odor-induced analgesic responses and their neurohormonal correlates may be part of an adaptive preparatory defense mechanism that facilitates the detection and avoidance of parasitized males by female mice and contributes to female mate choice.     

Cross-fostering Does Not Influence the Mate Preferences and Territorial Behaviour of Males in Lake Victoria Cichlids

The ability to recognise conspecifics in contexts of mate choice and territorial defence may have large effects on an individual's fitness. Understanding the development of assortative behaviour may shed light on how species assortative behaviour evolves and how it may influence reproductive isolation. This is the case not only for female mate preferences, but also for male mate preferences and male territorial behaviour. Here we test with a cross-fostering experiment whether early learning influences male mate preferences and male–male aggression biases in two closely related, sympatrically occurring cichlid species Pundamilia pundamilia and Pundamilia nyererei from Lake Victoria. Males that had been fostered, either by a conspecific female or by a heterospecific female, were tested for their aggression bias, as well as for their mate preferences, in two-way choice tests. Males cross-fostered with conspecific and heterospecific foster mothers selectively directed their aggression towards conspecific intruders. The cross-fostering treatment also did not affect male mate preferences. These results are in striking contrast with the finding that females of these species show a sexual preference for males of the foster species.     

Male-specific Iridescent Coloration in the Pipevine Swallowtail (Battus philenor) is Used in Mate Choice by Females but not Sexual Discrimination by Males

We investigated the potential roles in behavioral interactions of sexually dichromatic iridescent blue coloration found on the dorsal hindwing of male Pipevine Swallowtails (Battus philenor). Behavioral experiments in a large enclosure addressed whether male dorsal hindwing coloration mediated sexual recognition by males, female choice of mates, or both. Models presented to males in the enclosure produced responses that show that males discriminate females from other males using chemical but not visual cues. In contrast, ablation of male dorsal iridescence significantly reduced male mating success with virgin females in the enclosure. The results support the hypothesis that male-specific iridescent coloration in this species is a signal that is important for females in assessing either male species identity or quality in intersexual interactions.     

Female Mate Choice in a Paternal Brooding Blenny: The Process and Benefits of Mating with Males Tending Young Eggs

Female mate preference for males tending young offspring has been demonstrated in many fishes; however, not much is known about the choice process. Using the barred chin blenny Rhabdoblennius ellipes, a fish with male uniparental care, field experiments were conducted to investigate the female preference for males tending young eggs and then whether females choose the males with young eggs by discriminating young eggs from old eggs in the nests. Males tending young eggs (0‐ to 2‐d old) acquired new eggs nine times more frequently than those tending old eggs (3‐ to 5‐d old) regardless of other traits in males and nests. In the two egg‐switching field experiments (old to young and young to old), contrary to our expectation, male mating success was neither enhanced when given young eggs nor inhibited when given old eggs. These results suggested that females choose males with young eggs not by discriminating the developmental stage of eggs in the nests but by using other choice processes. By choosing males with young eggs, females may benefit from the dilution effects of egg predation and filial cannibalism risks and avoid male parental care failure.     

Visual Cues in Mate Recognition by Males of the Damselfly, Coenagrion puella (L.) (Odonata: Coenagrionidae)

Coenagrion puella males search actively for mates and are not aggressive to other males. To study the role of visual cues in male–female discrimination, four types of models were used: (1) bodies of intact insects, (2) models of painted males, (3) models of male–female chimerae, and (4) models of female body parts. Abdomen coloration pattern and presence of wings were the most important cues for sexual recognition by males. Step-by-step elimination of male coloration pattern leads to an increase in the tandem response rate. A female model painted as a male repelled males like the intact male model. The absence of either the head or the thorax slightly decreased the number of tandem responses, but models without both the head and the thorax were not recognized as a mate. Abdomen thickness larger than that of a normal female decreased the attractiveness of the model. Models of the gynochrome female were significantly more attractive than models of the androchrome one. Female models containing male parts were less attractive than models without any structure at this place. Using principal-components analysis, it is shown that models repelling males usually were those containing an intact male abdomen or a female abdomen painted with blue. The results indicate that C. puella males can distinguish males from females visually by morphological structures and coloration pattern.     

Male Competition over Access to Females in a Spider with Last-Male Sperm Precedence

Agonistic behaviour between male cellar spiders (Pholcus phalangioides) was investigated to test whether (1) size difference determines which male achieves access to the female, (2) males are able to monopolize access to the female until egg laying and whether (3) female resource value increases before egg laying because of last‐male sperm precedence. We further investigated whether (4) there is variation in time and energy spent on courtship and copulation depending on the degree of sperm competition, i.e. with or without rival present. In three experimental settings we introduced two males of either different or similar sizes, or a single male to a female. The mating units were constantly video‐observed until the females produced their first egg sac. Experience, ownership and female resource value in terms of body size was controlled. Our results show that larger males achieve almost exclusive access to females. Size symmetrical settings resulted in increased fighting activity and duration but dominance did not influence mating success. If copulations were disturbed by the rival male, copulations were terminated earlier in symmetrical settings compared with asymmetrical settings. In 94.8% of trials only one copulation took place, suggesting that the copulating male successfully monopolized access to the female. Males confronted with a rival copulated longer but courted significantly shorter than lone males. Although the last male to copulate sires 88% of the offspring in P. phalangioides, neither fighting nor courtship activity increased before the female laid a batch of eggs. This suggests that males have no indication of the timing of oviposition.     

Stickleback Males Increase Red Coloration and Courtship Behaviours in the Presence of a Competitive Rival

In species where females preferentially select the most colourful males, males may strategically invest in courtship and nuptial colour according to the presence of rivals. In this experimental study, we tested this in the three‐spined stickleback (Gasterosteus aculeatus) in which mature males exhibit carotenoid‐based red coloration to attract mates and defend their territories against male competitors. We challenged experimental males with either a red‐ornamented dummy male or a non‐ornamented dummy for five min per day in six‐d experimental trial, which was repeated twice during the breeding season. We found that the males presented with a coloured rival exhibited more frequent courtship behaviours (i.e. fanning and gluing) to females than those presented with a non‐coloured intruder during the second experimental trial. At the end of each trial, the experimental males also showed a significantly larger area of red coloration in the presence of a coloured intruder. Our findings suggest that male sticklebacks regulate mating effort according to the presence of competitive rivals by increasing their investment in costly signals when successful mating and territory defence is at risk.     

Females Paired with More Attractive Males Show Reduced Oxidative Damage: Possible Direct Benefits of Mate Choice in Pied Flycatchers

Direct benefits of female mate choice may concern female fertility and fecundity but also physiological status. In birds with biparental care, males may contribute to improve the condition and health of their pair‐mates through help in constructing nests, incubation or incubation feeding and nestling provisioning. They may also reduce harassment of females by non‐pair males. A consequence of these male activities could be expressed in terms of oxidative damage, which may depend on metabolic effort and social stress. Here, we have related male contribution to parental and territorial duties to female oxidative status in the pied flycatcher Ficedula hypoleuca, a species where preferred males present darker dorsal plumage and, in Iberian populations, a large white forehead patch. Darker males were paired with females with high incubation attendance and reduced nestling provisioning rates, which may lead to reduced female exertion. These males owned nest boxes at which there were fewer visits by non‐pair males. Although females paired with dark mates worked less hard, they were able to raise more fledglings. Female oxidative damage measured as malondialdehyde (MDA) level in plasma declined with increasing incubation attendance and male incubation feeding. Moreover, levels of MDA in females declined with both darkness of male dorsal plumage and male forehead patch size when controlling for female forehead patch size and male age. The effect of male plumage darkness was especially strong. Females paired with middle‐aged males (2–3 yr) showed reduced levels of MDA compared with those paired with 1‐yr‐old and more than 3‐yr‐old males. Male age could not explain the effects of male attractiveness. Females paired with attractive males were more successful in reproduction while suffering reduced oxidative damage, possibly mediated by help during incubation and nestling rearing from their pair‐mates. Although correlative, the evidence suggests direct benefits of females paired with more attractive males.     

Sociality, Mate Choice, and Timing of Mating in American Bison (Bison bison): Effects of Large Males

We studied group size, composition, and mating activities in American bison (Bison bison) during rut on the Delta Junction Bison Range in interior Alaska, USA, in 1996 and 1997. Our purpose was to determine the effects of large males (≥5 yr old) on mating and associated activities. Groups with large males were larger than those containing smaller males. Most groups of bison were mixed‐sex (90%), but large males occurred in only one‐half of all groups. Moreover, females in groups with large males were more likely to copulate than those in groups with smaller males, indicating a female preference for large males. Nevertheless, our results are consistent with large males seeking out adult females for mating rather than vice versa. Mating peaked in mid‐August during both the study years and was highly synchronous. Scent marking was coincident with mating, an outcome consistent with a hypothesis of such behavior triggering ovulation. Scent marking by large male bison occurred in both male–male and male–female contexts, but was associated most often with sexual activities. No differences in group size occurred with changes in weather or among vegetation types occupied by bison. Group size of bison, however, was larger with increasing distance from the forest edge, which likely was a response to predation risk in this predator‐rich environment.     

Spermatophore and Sperm Allocation in Males of the Monandrous Butterfly Pararge aegeria: the Female’s Perspective

In insects, spermatophore production represents a non‐trivial cost to a male. Non‐virgin males have been shown to produce small spermatophores at subsequent matings. Particularly in monandrous species, it may be an issue to receive a sufficiently large spermatophore at the first and typically only mating. Females of the monandrous Speckled wood butterfly Pararge aegeria (L.) produce fewer offspring after mating with a non‐virgin male. After mating, females spend all their active time selecting oviposition sites and typically ignore other males. Here, we show that females did not discriminate between a virgin male and a recently mated male in our laboratory experiments. We demonstrate that the number of eupyrene sperm bundles relative to spermatophore mass differed with subsequent male matings. Males transferred a significantly smaller spermatophore after the first copulation, but the spermatophore mass did not decrease further with subsequent matings. However, the number of eupyrene sperm bundles decreased linearly. Therefore, there was proportionally more eupyrene sperm in the male’s second spermatophore compared with the first and the later spermatophores. Such a pattern has been shown in polyandrous species. Hence, it suggests that differences in sperm allocation strategy between polyandrous and monandrous butterflies may be quantitative rather than qualitative. There was also a tendency for females that had mated with a recently mated male to have higher propensity to remate than did females that had mated with a virgin male. We discuss the results relative to the mating system in P. aegeria, including female remating opportunities in the field and male mate‐locating behaviour.     

Paternal Care Decreases Foraging Activity and Body Condition,but Does Not Impose Survival Costs to Caring Males in a Neotropical Arachnid

Exclusive paternal care is the rarest form of parental investment in nature and theory predicts that the maintenance of this behavior depends on the balance between costs and benefits to males. Our goal was to assess costs of paternal care in the harvestman Iporangaia pustulosa, for which the benefits of this behavior in terms of egg survival have already been demonstrated. We evaluated energetic costs and mortality risks associated to paternal egg-guarding in the field. We quantified foraging activity of males and estimated how their body condition is influenced by the duration of the caring period. Additionally, we conducted a one-year capture-mark-recapture study and estimated apparent survival probabilities of caring and non-caring males to assess potential survival costs of paternal care. Our results indicate that caring males forage less frequently than non-caring individuals (males and females) and that their body condition deteriorates over the course of the caring period. Thus, males willing to guard eggs may provide to females a fitness-enhancing gift of cost-free care of their offspring. Caring males, however, did not show lower survival probabilities when compared to both non-caring males and females. Reduction in mortality risks as a result of remaining stationary, combined with the benefits of improving egg survival, may have played an important and previously unsuspected role favoring the evolution of paternal care. Moreover, males exhibiting paternal care could also provide an honest signal of their quality as offspring defenders, and thus female preference for caring males could be responsible for maintaining the trait.     

Ocean Acidification Disrupts Prey Responses to Predator Cues but Not Net Prey Shell Growth in Concholepas concholepas (loco)

Background

Most research on Ocean Acidification (OA) has largely focused on the process of calcification and the physiological trade-offs employed by calcifying organisms to support the building of calcium carbonate structures. However, there is growing evidence that OA can also impact upon other key biological processes such as survival, growth and behaviour. On wave-swept rocky shores the ability of gastropods to self-right after dislodgement, and rapidly return to normal orientation, reduces the risk of predation.

Methodology/Principal Findings

The impacts of OA on this self-righting behaviour and other important parameters such as growth, survival, shell dissolution and shell deposition in Concholepas concholepas (loco) were investigated under contrasting pCO₂ levels. Although no impacts of OA on either growth or net shell calcification were found, the results did show that OA can significantly affect self-righting behaviour during the early ontogeny of this species with significantly faster righting times recorded for individuals of C. concholepas reared under increased average pCO₂ concentrations (± SE) (716±12 and 1036±14 µatm CO₂) compared to those reared at concentrations equivalent to those presently found in the surface ocean (388±8 µatm CO₂). When loco were also exposed to the predatory crab Acanthocyclus hassleri, righting times were again increased by exposure to elevated CO₂, although self-righting times were generally twice as fast as those observed in the absence of the crab.

Conclusions and Significance

These results suggest that self-righting in the early ontogeny of C. concholepas will be positively affected by pCO₂ levels expected by the end of the 21st century and beginning of the next one. However, as the rate of self-righting is an adaptive trait evolved to reduce lethal predatory attacks, our result also suggest that OA may disrupt prey responses to predators in nature.     

Fitness Benefits of Mate Choice for Compatibility in a Socially Monogamous Species

Research on mate choice has primarily focused on preferences for quality indicators, assuming that all individuals show consensus about who is the most attractive. However, in some species, mating preferences seem largely individual-specific, suggesting that they might target genetic or behavioral compatibility. Few studies have quantified the fitness consequences of allowing versus preventing such idiosyncratic mate choice. Here, we report on an experiment that controls for variation in overall partner quality and show that zebra finch (Taeniopygia guttata) pairs that resulted from free mate choice achieved a 37% higher reproductive success than pairs that were forced to mate. Cross-fostering of freshly laid eggs showed that embryo mortality (before hatching) primarily depended on the identity of the genetic parents, whereas offspring mortality during the rearing period depended on foster-parent identity. Therefore, preventing mate choice should lead to an increase in embryo mortality if mate choice targets genetic compatibility (for embryo viability), and to an increase in offspring mortality if mate choice targets behavioral compatibility (for better rearing). We found that pairs from both treatments showed equal rates of embryo mortality, but chosen pairs were better at raising offspring. These results thus support the behavioral, but not the genetic, compatibility hypothesis. Further exploratory analyses reveal several differences in behavior and fitness components between “free-choice” and “forced” pairs.