THE EVOLUTION OF ALTERNATIVE REPRODUCTIVE TACTICS IN MALE CARDIOCONDYLA ANTS

Alternative reproductive tactics are often associated with discontinuous variation in morphology but may evolve independent from each other. Based on life‐history data and a phylogeny we examine how male morphology and reproductive behavior are linked in the evolution of the ant genus Cardiocondyla. Wingless Cardiocondyla males engage in lethal fighting for access to female sexuals, whereas winged males disperse and mate away from the nest. This basic pattern shows considerable variation across species. A phylogeny based on ～3 kbp sequence data shows that male diphenism and lethal fighting are ancestral traits tightly linked in evolution. Winged males were lost convergently in several species groups, apparently in response to the low probability of encountering female sexuals in nests without a resident fighter male. An early dichotomy separates two clades with alternative male morphologies and fighting behavior, but phenotype and fighting strategy are not correlated with the presence of winged males.     

Encounter with heavier females changes courtship and fighting efforts of male field crickets Gryllus bimaculatus (Orthoptera: Gryllidae)

The effects of mating experience on male mating behavior are mediated by four factors: mating cost, such as resource depletion, perception of mating opportunities, self-perception of attractiveness, and female quality. For example, encountering females might increase male expectations of prospective mating opportunities, while copulation increases self-perception of attractiveness in males. To determine the relative importance of these factors, the effect of mating on the two components of reproductive effort (courtship and fighting effort) in Gryllus bimaculatus was examined. Calling activity before and after encountering females was measured, and copulation success was recorded. Subsequently, the intensity and outcome of male–male fighting behavior was recorded. Female encounter increased calling activity irrespective of copulation, thereby indicating that the perception of mating opportunities is important factor for the males. Changes in courtship effort of males were larger and fighting success was lower when they were previously paired with relatively heavier females. These results indicate that male reproductive effort is also affected by quality of previous mating partners.

     

Hunting and Nesting Phenology Influence Gobbling of Wild Turkeys

Wild turkeys (Meleagris gallopavo) use a polygynous mating system whereby males engage in multiple courtship behaviors, including vocalizations (gobbling) to attract females and compete with other males for breeding opportunities. Males must balance the risk of courtship behaviors with the reproductive potential of each courtship behavior. Male turkeys are primarily hunted during the reproductive period, so the associated risk of courtship behaviors is increased. Many state agencies attempt to set hunting season frameworks that maximize hunter satisfaction by allowing hunting when gobbling activity is greatest and most females are theoretically incubating nests, but the relationship between gobbling activity and nesting phenology is unclear. We used autonomous recording units and global positioning system transmitters to monitor gobbling activity by male turkeys and reproductive behaviors of female turkeys in the Piedmont region of Georgia, USA. We used 13,177 gobbles, behavioral data from 82 females during the reproductive season, and daily estimates of harvest of males by hunters to examine relationships between daily gobbling activity, cumulative removal of males, and reproductive behaviors (laying, incubating) of females during 2017–2018. We observed a weak negative relationship between daily gobbling activity and gobbling activity the following day. As the reproductive season progressed, gobbling activity decreased. As the proportion of females engaged in laying or incubating behaviors increased, expected daily gobbling activity increased. Conversely, we observed that hunting and removal of males had a negative effect on daily gobbling activity, and this effect was disproportionately greater than the positive effect of female reproductive behaviors. Our findings suggest that hunting and removal of males are important determinants of gobbling activity, and that corresponding reductions in gobbling activity may have mediating effects on the mating system of wild turkeys. © 2019 The Wildlife Society.     

Impaired sperm quality,delayed mating but no costs for offspring fitness in crickets winning a fight

The outcome of male–male contest competition is known to affect male mating success and is believed to confer fitness benefits to females through preference for dominant males. However, by mating with contest winners, females can incur significant costs spanning from decreased fecundity to negative effects on offspring. Hence, identifying costs and benefits of male dominance on female fitness is crucial to unravel the potential for a conflict of interests between the sexes. Here, we investigated males' pre‐ and post‐copulatory reproductive investment and its effect on female fitness after a single contest a using the field cricket Gryllus bimaculatus. We allowed males to fight and immediately measured their mating behaviour, sperm quality and offspring viability. We found that males experiencing a fight, independently of the outcome, delayed matings, but their courtship effort was not affected. However, winners produced sperm of lower quality (viability) compared to losers and to males that did not experience fighting. Results suggest a trade‐off in resource allocation between pre‐ and post‐mating episodes of sexual selection. Despite lower ejaculate quality, we found no fitness costs (fecundity and viability of offspring) for females mated to winners. Overall, our findings highlight the importance of considering fighting ability when assessing male reproductive success, as winners may be impaired in their competitiveness at a post‐mating level.     

Causes of a male-biased operational sex ratio in the fiddler crab<Emphasis Type="Italic"> Uca crenulata</Emphasis>

This research investigates the causes of a male-biased operational sex ratio in a population of the California fiddler crab, Uca crenulata. Mensurative studies revealed there were almost twice as many adult males as females, mating occurred across half of the days within the breeding season, and females had much longer individual reproductive cycles than males. Therefore, many more males than females were available for mating on each breeding day. Perhaps as a consequence, males spent a large proportion of their time fighting with neighbors and rapidly waving their large claws when females passed by. Electronic Publication     

Changes in Fecal Estrogen Levels and Sexual Behavior in Captive Sichuan Snub‐Nosed Monkeys (Rhinopithecus roxellana) Following a Male Replacement

Rhinopithecus roxellana are generally seasonal breeders, although copulation can occur throughout the year. Previous studies suggest that estradiol modulates female sexual behavior during the mating season. However, the effects of social context on estrogen levels and behavior have not been fully explored. We studied the relationship between sexual behaviors and fecal estrogens in a group of captive R. roxellana during a period of social instability. We collected behavioral data for six months and collected fecal samples at 2–3‐day intervals for four months spanning the mating and nonmating seasons, and analyzed fecal estrogen levels via RIA. Females showed clear cyclic solicitation and copulation peaks in the mating season, which corresponded with sharp peaks in fecal estrogens. During the nonmating season, solicitation rates, copulation rates, and fecal estrogens were generally low. However, one nonpregnant female displayed a sharp peak in solicitations, copulations, and estrogens during the nonmating season 10–14 days after a male replacement. Our results provide preliminary evidence that social and behavioral changes affect estrogen levels in R. roxellana. Zoo Biol. 32:342–346, 2013. © 2012 Wiley Periodicals, Inc.     

Characterization of social behavior in the spiny mouse,Acomys cahirinus

While there are many species that are commonly used for the study of mammalian social behavior, there remains a need for lab-suitable organisms that are appropriate for examining sociality specifically in non-reproductive contexts (i.e., social behavior not in the context of mating or parenting). The spiny mouse, Acomys cahirinus, is a cooperatively breeding rodent that lives in large groups and is a species that holds great potential for studying a wide range of social behaviors in reproductive and non-reproductive contexts. Here, we characterize the basic social behaviors in male and female A. cahirinus to obtain a foundation for future study. We tested adult A. cahirinus in social approach, social preference, social interaction, social recognition, and group size preference paradigms. Regardless of sex, novelty, or familiarity, we found that both males and females rapidly approach conspecifics demonstrating high social boldness. Additionally, both sexes are significantly more prosocial than aggressive when freely interacting with conspecifics. However, we observed effects of sex on social preferences, such that males exhibit a preference to affiliate with same-sex conspecifics, whereas females exhibit a preference for affiliating with opposite-sex conspecifics. We discuss how this preference may relate to the cooperative breeding system of spiny mice. Lastly, both sexes show a robust preference for affiliating with large over small groups, indicating they may be an ideal species for the study of mammalian gregariousness. These data lay a basic foundation for future studies that seek to assess complex group dynamics and the mechanisms underlying reproductive and non-reproductive social behaviors in a highly social mammal.     

Relationships among male sexually selected traits in the bean bug,Riptortus pedestris (Heteroptera: Alydidae)

Current concepts of sexual selection suggest that male reproductive success is determined by multiple sexual traits. As expression and production of multiple sexual traits are frequently associated with each other, positive or negative correlations among multiple sexual traits ensue. These relationships among traits associated with male reproductive success may be crucial in the evolution of male reproductive strategies. Here, we investigate phenotypic relationships among sexually selected traits in the armed bean bug Riptortus pedestris. In this insect, males with a larger body and weapon are more likely to win male–male competitions, and males with a larger weapon or higher courtship rate are more attractive to females. There was a significant positive correlation between body size and weapon size, whereas the courtship rate had significant negative correlations with body size and weapon size. Our results suggested that there was a phenotypic trade‐off between courtship rate and male morphology. In this insect, smaller males may make more effort in courtship behavior as an alternative mating tactic.     

Aggression in adult male primates: A comparison of confined Japanese macaques and free-ranging olive baboons

The frequencies and types of adult male aggressive behavior of confined Japanese macaques (Macaca fuscata)and free-ranging olive baboons (Papio anubis)were compared. The baboons, which do not have a mating season, were more aggressive to conspecific males than were the macaques during their nonmating season. The baboons also solicited aid during aggressive encounters more frequently than the macaques. However, during their mating season, the macaques were more aggressive to conspecific females than were the baboons. The macaques were also involved in more triadic sequences of aggression, and the frequency of occurrence of these patterns supported Chase’s theory of dominance hierarchy formation and maintenance. The differences in aggressive behavior appeared to be related to the seasonal reproductive cycle of the macaques.     

韭菜迟眼蕈蚊成虫行为活动昼夜节律

【目的】本研究旨在明确韭菜迟眼蕈蚊Bradysia cellarum成虫行为活动的昼夜节律,为其室内饲养、行为学研究和研发精准、高效的监测与防控新技术提供依据。【方法】室内系统观察记录了韭菜迟眼蕈蚊成虫羽化盛期求偶、交配、产卵、移动与静息等行为的昼夜节律,分析了其特定行为的时间分配特征。【结果】韭菜迟眼蕈蚊成虫的运动行为和静息行为白天和黑夜均有发生,夜间的繁殖行为(求偶、交配和产卵)频次高于白天的,各时间点的移动、静息和繁殖行为频次占比间均具有显著性差异。韭菜迟眼蕈蚊雌雄成虫的求偶节律相同,一天中求偶有两次高峰,分别出现在3:00和7:00。韭菜迟眼蕈蚊雌雄成虫的交配节律与雌成虫的产卵节律相似,雄成虫移动行为频次占比的峰值时刻均出现在3:00时,雌成虫移动行为频次占比的峰值时刻则出现在19:00时。雌雄成虫静息行为的频次占比均无明显昼夜节律高峰。在日时间分配上,雌、雄成虫以静息行为的时间占比最多,分别为89.53%和89.90%;求偶行为的时间占比最少,分别为0.56%和0.89%。雌雄成虫各行为的日时间占比差异不显著。【结论】韭菜迟眼蕈蚊成虫的求偶、交配、产卵、移动和静息行为具有明显...     

Male age affects female mating preference but not fitness in the monandrous moth Dendrolimus punctatus Walker (Lepidoptera: Lasiocampidae)

It is widely accepted that male age can influence female mating preference and subsequent fitness consequences in many polyandrous species, yet this is seldom investigated in monandrous species. In the present study, we use the monandrous pine moth Dendrolimus punctatus to examine the effects of male age on female mating preference and future reproductive potential. In multiple male trials, when permitted free mating from an aggregation consisting of virgin males aged 0 (young), 2 (middle-aged) and 4 (old) days, virgin females preferentially mate with young and middle-aged males, although mating latency and mating duration are independent of male age. In single male trials, when virgin females are randomly assigned single virgin males of known age, a negative correlation is found between mating success and male age in this species. However, we find that male age also has no effect on mating latency and mating duration. Further fitness analysis reveals that females do not receive benefits in terms of oviposition period, total egg production, average daily egg production, percentage of egg hatching, longevity, expected reproduction and relative expected reproduction from mating with young and middle-aged males compared with mating with old males. The results of the present study are the first demonstrate that females mated preferentially with younger males but gain no apparent fitness benefits in a monandrous moth species.     

Adult Male Scent-Marking in Lemur catta and Eulemur fulvus rufus

Scent-marking and olfactory communication are used extensively by prosimians and can provide spatial and temporal records of group movement and behavior. We compare rates of male scent-marking in relation to reproductive seasons, male dominance rank, and habitat use in two related prosimians: Lemur catta and Eulemur fulvus rufus. We collected scent-marking data on adult male Lemur catta at Beza-Mahafaly Reserve (dry forest), and on Eulemur fulvus rufus at Ranomafana National Park (rainforest), Madagascar. In Lemur, rates of overall scent-marking differed significantly by reproductive season, with higher rates occurring in mating and lactation/migration periods, whereas in Eulemur, reproductive season did not appear to affect scent-marking rates. Dominance rank of male Lemur catta did not affect rates of scent-marking. Among male Eulemur fulvus, dominance relations were not apparent; however, 2 of the 5 focal males scent-marked somewhat more frequently during the mating season and also experienced greater mating success. In Lemur catta, higher rates of scent-marking in the mating season may relate to indirect reproductive competition during a period of high aggression, while such mating competition was not as marked in Eulemur fulvus. Furthermore, higher rates of marking in resident male Lemur catta during male migration may correlate with vigilance toward immigrating males. Greater overall scent-marking rates in ring-tailed lemurs may relate to extensive intergroup home range overlap and no area of exclusive use, whereas the red-fronted lemur groups tended to forage in areas of their home range where little-to-no intergroup overlap occurred.     

The Haremic Mating System and Mate Choice in the Wide-Eyed Flounder, Bothus podas

The social structure and reproductive behaviour of the wide-eyed flounder, Bothus podas, was studied in the coastal waters around the Azorean Islands. Both sexes are territorial throughout the year. Adult males defend large territories, which include several smaller female territories. Intraspecific agonistic behaviour was frequent and differed between sexes: males were more aggressive towards other males, while females were only aggressive towards each other and juveniles. During the reproductive season and only at dawn, territorial males court and mate successively with females in their territories, and females seem to show mating fidelity to their dominant male. Such territoriality and mating patterns indicate a haremic social system in the wide-eyed flounder. In order to identify potential factors influencing female mate choice acting on this haremic system, we examined male mating success and some of its potential correlates. We found no evidence for female preference for any of the males' physical or territory characteristics. However, courtship effort was strongly correlated with the total number of attempted and successful spawnings, indicating that females seem to mate preferentially with males that court them more vigorously. Thus, our data suggest that courtship plays an important role in determining male mating success in the wide-eyed flounder and, that it may possibly serve as an honest indicator of male `quality' for female choice.     

Males with short horns spent more time mating in the Japanese horned beetle Allomyrina dichotoma

We examined the relationships between male body and horn sizes and mating duration in the Japanese horned beetle, Allomyrina dichotoma. Smaller males possessing shorter horns spent more time for copulation with a female and mounting the female without copulation. The results of multiple regression analyses indicate that the horn length is a determining factor for the time spent by the males during mating. A previous study has documented that the horn length of male A. dichotoma primarily determined the outcomes of aggressive male–male interactions; hence, predicts access to females. Therefore, instead of fighting for females, males possessing short horns may maximize their fertilization success by mating longer with the few females they have access to.     

Social isolation increases male aggression toward females in the field cricket Gryllus bimaculatus

Social isolation has often been reported to facilitate male aggressiveness in various animal species. If social isolation also escalates male aggressive behavior towards females, the mating success of the aggressive males will be low. This study evaluated the effect of social isolation on mating behavior in the field cricket Gryllus bimaculatus, which has traditionally been considered to be an asocial species. The results showed that social isolation from same-sex individuals enhanced male aggressiveness to females, and the mating success of aggressive males was reduced under the experimental conditions. More aggressive males exhibited a longer latency to court than less aggressive males. These results suggest that because male aggressiveness causes a delay in courtship, aggressive males may have reduced mating success. This demonstrated that social relationships are a critical factor affecting male mating success, even if the species is normally considered solitary.     

Day–night fluctuations in floral scent and their effects on reproductive success in <Emphasis Type="Italic">Lilium auratum</Emphasis>

We examined the contribution of diurnal and nocturnal pollination to male and female reproductive success in Lilium auratum. Plants were bagged for either 12 h during the day or at night to allow either only nocturnal or only diurnal visitors to forage throughout the flowering period. We found that there was no significant difference in the seed:ovule ratio among diurnally pollinated, nocturnally pollinated, or control flowers. However, in terms of male reproductive success, it was more advantageous for the plants to be pollinated both diurnally and nocturnally: the numbers of pollen grains remaining in diurnally pollinated or nocturnally pollinated flowers were significantly greater than those in control flowers. The total amount of floral volatiles of L. auratum was significantly higher at night than during the day. The constituents of floral scent of all time series examined were mostly monoterpenoids, many of which serve as attractants for nocturnal hawkmoths. Such nocturnally biased floral scent emission of L. auratum might achieve male reproductive success by attracting nocturnal visitors, which may suggest that the relative contribution of floral scent in this species is biased towards male reproductive success.     

First Evidence of Inclusive Sexual Selection in the Ant Cataglyphis cursor: Worker Aggressions Differentially Affect Male Access to Virgin Queens

Sexual selection typically shapes the traits of individuals capable of reproduction. However, in social species, it may also affect the behaviour of non‐reproductive individuals if this increases their inclusive fitness. We examined the potential role of workers during mating in the polyandrous ant Cataglyphis cursor. In this species, mating takes place on the ground near the nest entrance, and workers chase away foreign males trying to mate with young queens (gynes). We investigated the effects of worker aggression on male mating attempts by placing three or four males in the presence of one foreign gyne and 20 of her nestmate workers and contrasted this with a second group of 3–4 males in the presence of another gyne without nestmate workers (n = 8 paired replicates). Workers strongly attacked males (more than 20 aggressions per minute), although this did not affect the total number of mating attempts by the 3–4 males or the skew in the number of mating attempts among these males. However, there was an interaction between worker aggression and relative male weight, as relatively heavy males attempted to mate more often than lighter males in the absence of worker whereas the reverse occurred in the presence of workers. Our study is the first to experimentally test the potential role of non‐reproductive individuals in sexual selection. It shows that workers could potentially change male mating success by disfavouring heavier males.     

Plumage color as a status signal in male–male interaction in the red-flanked bushrobin, Tarsiger cyanurus

Recent studies have suggested that structural-based coloration is an honest signal of male genetic and/or conditional quality in sexual selection. However, whether structural coloration functions in intrasexual competition is unknown. We examined whether plumage color functions as a status signal during intrasexual interactions in the red-flanked bushrobin Tarsiger cyanurus; adult males have many blue plumes as structural coloration whereas yearling males and females are olive brown with few blue plumages. Blue males did not always dominate olive-brown males. The number of interactions did not differ with the colors of the two birds involved. The interactions of a blue male and an olive-brown male were less aggressive than those of two blue or of two olive-brown males. In this study, we found that structural plumage coloration may serve as a signal of aggressive intent and lower the escalation level of an aggressive interaction in a manner consistent with hypotheses regarding the evolution of delayed plumage maturation.     

The coevolution of juvenile play–fighting and adult competition

Although play–fighting is widespread among juvenile mammals, its adaptive significance remains unclear. It has been proposed that play is beneficial for developing skills to improve success in adult contests (motor‐training hypothesis), but the links between juvenile play–fighting and adult aggression are complex and not well understood. In this theoretical study, we investigate the coevolution between juvenile play–fighting and adult aggression using evolutionary computer simulations. We consider a simple life history with two sequential stages: a juvenile phase in which individuals play–fight with other juveniles to develop their fighting skills; and an adult phase in which individuals engage in potentially aggressive contests over access to resources and ultimately mating opportunities, leading to reproductive success. The simulations track genetic evolution in key traits affecting adult contests, such as the level of aggression, as well as juvenile investment in play–fighting, capturing the coevolutionary feedbacks between juvenile and adult decisions. We find that coevolution leads to one of two outcomes: a high‐play, high‐aggression situation with highly aggressive adult contests preceded by a prolonged period of juvenile play–fighting to improve fighting ability, or a low‐play, low‐aggression situation in which adult contests are resolved without fighting and there is minimal investment in play–fighting before individuals mature. Which of these outcomes is favoured depends on the mortality costs and on the type of societal structure: societies with strong reproductive skew, favouring monopolization of resources, show high levels of adult aggression and high investment in juvenile play–fighting, whereas societies with low reproductive skew have both low adult aggression and low levels of play–fighting. A review of empirical evidence, particularly in the primate genus Macaca, highlights some limitations of our model and suggests that other, complementary functional explanations are needed to account for the full range of competitive and cooperative forms of play–fighting. Our study illustrates the power of evolutionary simulations to shed light on the long‐standing puzzle of animal play.