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.     

Do Male Veiled Chameleons,Chamaeleo calyptratus,Adjust their Courtship Displays in Response to Female Reproductive Status?

Variation in male courtship behavior may be due to inherent differences among males or may arise from males adjusting their courtship displays according to female responsiveness. Female veiled chameleons, Chamaeleo calyptratus , exhibit two distinctive suites of body coloration and behavior patterns that vary according to receptive and non-receptive stages of their reproductive cycle. We presented male chameleons with both receptive and non-receptive females, and recorded differences in their mating frequency, courtship intensity and courtship behavior patterns. As expected, males were more likely to court and attempt mating with receptive females. Although fewer males courted non-receptive females, their courtship displays were significantly longer than those directed towards receptive females. Males also adjusted the contents of their displays according to female reproductive condition. Certain behavior patterns were unique to courtship displays directed towards each class of females. Males exhibited the behavior pattern `head roll' only when paired with receptive females, and `chin rub' was displayed only during courtship of non-receptive females. We hypothesize that these differences in male courtship frequency, intensity and content reflect differences in female reproductive value. Although males may benefit from mating with both receptive and non-receptive females, the costs associated with courtship may depend on female responsiveness. Thus, males adjust their courtship tactics accordingly.     

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.     

Reproductive Signals of Female Lizards: Pattern of Trait Expression and Male Response

Relative to the volume of studies concerning the function and evolution of male‐biased sexually dimorphic traits, instances of female‐biased sexual dimorphisms remain largely unstudied, especially in species with conventional sex roles. I investigated the signal function of a female‐specific ornamental trait using the striped plateau lizard (Sceloporus virgatus, Phrynosomatidae) as a model system. During the reproductive season, female S. virgatus develop orange color on their throats that is absent in conspecific males. I established the relationship between color expression and female reproductive state, and determined male response to female color. I show that dynamic changes occurring within the color patch can potentially identify each stage of the female reproductive cycle, largely because of a lag in patch growth relative to color intensification. Sexual receptivity is associated with intense patches rapidly growing in size; ovulation occurs near peak color expression; and the unreceptive period is associated with large patches fading in intensity. Because females express orange color during both the receptive and unreceptive periods, the pattern of color expression is consistent with the courtship‐stimulation and courtship‐rejection hypotheses of signal function. Males may preferentially associate with females that have more highly developed color patches during the courtship season, and/or ignore such females when they are unreceptive. An examination of male behavior towards unfamiliar females indicates that female color has a role in courtship stimulation but has little, if any, role in courtship rejection. During the pre‐mating season, males maintained significantly closer affiliation with, and tended to perform more social behavior towards females with more intense color. During the post‐mating season, female color had no apparent effect on male behavior. The evolution and current function of female ornaments may vary among taxonomically‐related species as a result of differences in ecology, social system, and life‐history.     

Complex Courtship Behavior in the Striped Ground Cricket, Allonemobius socius (Orthoptera: Gryllidae): Does Social Environment Affect Male and Female Behavior?

Complex courtship in the striped ground cricket, Allonemobius socius, involves a series of behaviors alternating between the sexes. We examined if complex courtship allows either or both genders to evaluate their mate and how mating behavior changes in different social environments. While complex courtship may allow discrimination by both sexes, here only females exhibited a preference. Males did not alter their courtship behavior or change spermatophore size for different size females. In contrast, females initiated copulation more quickly with bigger males possessing bigger spermatophores. In a different social environment (additional male, female, or both), males were less likely to omit courtship songs and female discrimination of mates changed. The distinct differences in male and female behavior suggest that subtle changes in social environment can have important consequences in structuring courtship and mating behavior.     

Male dwarf chameleons assess risk of courting large, aggressive females

Conflict between the sexes has traditionally been studied in terms of costs of mating to females and female resistance. However, courting can also be costly to males, especially when females are larger and aggressively resist copulation attempts. We examined male display intensity towards females in the Cape dwarf chameleon, Bradypodion pumilum, in which females are larger than males and very aggressive. We assessed whether aggressive female rejection imposes potential costs on males and whether males vary their display behaviour with intensity of female rejection, female size or relative size differences. Males persisted in courtship after initial female rejection in 84% of trials, and were bitten in 28% of trials. Attempted mounts were positively associated with males being bitten. Males reduced courtship with increased intensity of female rejection. Male courtship behaviour also varied with female size: males were more likely to court and approach smaller females, consistent with the hypothesis that larger females can inflict more damage. These results suggest that, in addition to assessing female willingness to mate, male dwarf chameleons may use courtship displays to assess potential costs of persistence, including costs associated with aggressive female rejection, weighed against potential reproductive pay-offs associated with forced copulation.     

Spiders,microbes and sex: Bacterial exposure on copulatory organs alters mating behaviour in funnel‐web spiders

Environmental microbes have the potential to be involved in nearly all behavioural processes. For example, mating systems where males use intromittent organs to transfer sperm to females represent a means by which environmental microbes collected by males can breach entry into females' body cavities during mating. However, the degree to which the acquisition of environmental microbes onto important sex structures alters courtship behaviours remains unknown. Here, we collected bacteria from the copulatory organs of Agelenopsis pennsylvanica funnel‐weaving spiders in situ to test whether exposure to bacteria on copulatory organs can alter hosts' courtship behaviour, reproductive success and survival. We used a standardized assay to repeatedly measure each spider's aggressiveness, a behavioural component of both male courtship and female sexual receptivity. Then, we experimentally altered the bacteria present on male and female spiders' copulatory organs with an application of either (a) a mixture of bacteria collected from conspecifics to increase bacterial presence, (b) an antibiotic to reduce bacterial presence or (c) a procedural control. Each spider was paired with a size‐matched spider of the opposite sex whose copulatory organs were unaltered, and we measured the latency until the onset and the duration of courtship. Spiders were then isolated, and we measured each individual's time until death and female fecundity over the next 40 days. We found that female exposure to bacteria had multiple effects on mating dynamics. Males took over four times longer to begin courting females that had been exposed to bacteria compared to unexposed and antibiotic‐treated females. Only when courting these bacteria‐exposed females, males began courtship sooner when females were more aggressive. Lastly, females whose mate had been exposed to bacteria experienced reduced survival. These data suggest that bacteria present on animals' copulatory organs can alter courtship behaviours, female survivorship, and may potentially play a role in mating dynamics.     

Courtship behaviour of the African Maize Stem Borer: Busseola fusca (Fuller) (Lepidoptera: Noctuidae) under laboratory conditions

Busseola fusca (Fuller) (Lepidoptera: Noctuidae) is the most important African stem borer damaging maize and sorghum. Chemical mediators play an essential role in all life cycle of this moth, especially for mating recognition and host plant choice. The female sex pheromone, courtship and mating behaviours act on the reproductive isolation within insect populations. B. fusca courtship behaviour was studied to decipher each step that could account as a process for reproductive isolation. B. fusca males and females presented a very simple and fast courtship behaviour, without any particular events or male pheromone emission.     

Seasonal sex role changes in the blenniid Petroscirtes breviceps, a nest brooder with paternal care

The plasticity of the sex roles in the blenniid fish Petroscirtes breviceps , a nest brooder with exclusive paternal care, was studied throughout an 8 month breeding season. Males performed most courtships early and late in the breeding season, whereas females performed most in the middle of the season. These results indicated that the sex of individuals initiating courtship changed seasonally, with courtship role reversal in the middle of the season. Intrasexual aggression in both sexes occurred much more frequently in mid-season than in the early and late seasons. Males frequently fought when available nest sites were limited, regardless of the presence of females, suggesting that males competed for nests in order to qualify to mate (resource competition). In contrast, courting females fought only in mid-season, when females' relative success in entering nests decreased, indicating that females competed for limited mating opportunities (mating competition). The reversed courtship roles and female mating competition in mid-season suggested that the sex roles in P. breviceps changed seasonally from the conventional roles to reversed roles and back again during one breeding season. This study provides the first empirical evidence of multiple changes in the sex roles of animals within a breeding season.     

Alternative reproductive tactics and courtship in the common goby

In the marine common goby Pomatoschistus microps , males either build nests and attract females to spawn or act as sneakers, forcing themselves into other males' nests during mating, to fertilize some of the eggs in the clutch. The reproductive tactics of small males were studied in the presence of surplus nest material and an excess of gravid females, while a larger, nestholding male spawned with a female. Of 24 small males, 13 only attempted to sneak, seven only courted females and/or spawned in their own nest, and four performed both behaviours. The small males who attracted females successfully were significantly larger than the ones who failed to do so. Females had been courting the small male more often when he subsequently spawned in his own nest, than when he did not get a female. The small males did not court the females very frequently, and their courtship rate did not depend on whether or not they attracted females. Instead, female courtship behaviour may be a cue for a male either to spawn as a nest holder or become a sneaker. The study pointed out the importance of female behaviour for male reproductive tactics, which could be more important in fish than is known currently.     

An acute stressor alters steroid hormone levels and activity but not sexual behavior in male and female Ocoee salamanders (Desmognathus ocoee)

Stressors that are chronic have clear suppressive effects on reproductive behaviors in both males and females. Stressors that are acute have effects on reproductive behavior that are less clear. We measured the effects of an acute bout of handling in laboratory-housed male and female Ocoee salamanders (Desmognathus ocoee), a species with a prolonged mating season. Handling resulted in decreased locomotory activity and elevated plasma corticosterone, a hallmark of the vertebrate stress response. Handling also decreased plasma testosterone in males and elevated plasma estradiol in females. Despite the handling-induced changes in hormone levels, handling had minimal impact on courtship and mating. Other species in which reproduction is insensitive to acute stressors may live in extreme environments with limited reproductive opportunities, whereas Ocoee salamanders live in a relatively temperate environment with multiple reproductive opportunities. Together, these data indicate that an allostatic response to a stressor can alter locomotory activity and elevate corticosterone without suppressing nonessential behaviors like courtship and mating in a species in which reproductive opportunities can occur over a period of multiple months. The lack of reproductive suppression in Ocoee salamanders might be due to the low energetic cost of courtship and mating in this species combined with potentially elevated energetic stores, highlighting the importance of considering energy budgets when making predictions about behavioral effects of acute stressors.     

Mating system and the evolution of sex-specific mortality rates in two nymphalid butterflies

Life-history theory predicts that organisms should invest resources into intrinsic components of lifespan only to the degree that it pays off in terms of reproductive success. The benefit of a long life may differ between the sexes and different mating systems may therefore select for different sex-specific mortality rates. In insects with polyandrous mating systems, females mate throughout their lives and male reproductive success is likely to increase monotonously with lifespan. In monandrous systems, where the mating season is less protracted because receptive females are available only at the beginning of the flight season, male mating success should be less dependent on a long lifespan. Here, we show, in a laboratory experiment without predation, that the duration of the mating season is longer in the polyandrous comma butterfly, Polygonia c-album, than in the monandrous peacock butterfly, Inachis io, and that, in line with predictions, male lifespan is shorter than female lifespan in I. io, whereas male and female lifespans are similar in P. c-album.     

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

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

Effect of multiple mating on the reproductive performance of the rice leaffolder moth,Cnaphalocrocis medinalis (Lepidoptera: Crambidae)

To clarify whether multiple mating of females and males affects the reproductive performance of the rice leaffolder moth, Cnaphalocrocis medinalis (Guenée), we examined the effect of the number of matings (once, twice, or three times) for females (female treatment) and males (male treatment) on the incidence of moth mating, number of eggs laid, egg hatchability, and adult longevity. We also compared the effect of multiple mating imposed on males or females separately with the effect of that imposed on both sexes simultaneously (both sexes treatment). The incidence of mating of females and males that mated three times (3-mated females and males) was significantly lower than for females and males that mated twice or once (2-mated or 1-mated females and males). The incidence of mating of 1-mated moths (both sexes) was significantly higher than for 2-mated or 3-mated moths (both sexes). Two-mated or 3-mated females laid significantly more eggs with significantly higher hatchability than 1-mated females. Females that mated with 1-mated males (second male mating) or 2-mated males (third male mating) laid significantly fewer eggs than those that mated with virgin males (first male mating). Females laid significantly more eggs after the second and third matings for moths of both sexes than after the first mating for moths of both sexes. The mechanisms of improvement and decline of female reproductive performance when multiple mating was imposed on males or females are also discussed in relation to the reproductive biology of C. medinalis.     

Seasonal Variation in Mate Choice of Photinus ignitus Fireflies

Mate choice by either sex may vary with changes in the associated costs and benefits, determined by factors such as the availability of potential mates and variation in mate quality. We examined seasonal variation in operational sex ratio, courtship behavior, spermatophore mass, egg count, and the relationship between morphological traits and mating success in Photinus ignitus fireflies to determine if mate choice in either sex varied with the availability and relative reproductive investment of fertilizable females and sexually active males. Successfully mating males had larger lanterns than unsuccessful males when the operational sex ratio was male‐biased. In addition, female responsiveness to male signals increased as the number of courting males decreased, and male spermatophore mass decreased with body size across the mating season. Successfully mating females had larger body mass than unsuccessful females. Female body mass predicted egg count and female rejection by males increased as the season progressed and female size decreased. These results suggest that both male and female P. ignitus exhibit mate choice, and that such choice is influenced by seasonal variation in the abundance and quality of potential mates.     

Mating behaviour and reproductive isolation of three species of reptile tick

The mating behaviour of three species of reptile tick, Aponomma hydrosauri, Amblyomma albolimbatum and Amb. limbatum is similar and involves a female sex pheromone which activates males to detach from their hosts and search. After contacting females, a stereotyped six-phase courtship sequence occurs. There are qualitative differences between the species in courtship behaviour at phase 3 (reversal of position by dorsally mounted males) and phase 6 (the copulation position). On-host observations of non-conspecific mating show that females of the three species have species-specific activation pheromones, which is contrary to reports in other species of tick. Such specificity should result in reproductive isolation of the three species; however, under certain circumstances it may not prevent non-conspecific contacts between the sexes. Off-host observations of courtship behaviour show that once males of the three species contact non-conspecific females, they attempt courtship and are persistent with their courtship. Rarely did non-conspecific courtships proceed beyond phase 4 of the courtship sequence, as non-conspecific females did not lift their bodies to allow males venter contact. Differences between the species in leg orientation in the copulation position, together with body size differences, are responsible for a complete barrier to successful non-conspecific copulation. The observations illustrate the role that behavioural mechanisms play in reproductively isolating these three species of tick.     

Size-biased Mating in Both Sexes of the Black-horned Tree Cricket, <Emphasis Type="Italic">Oecanthus nigricornis</Emphasis> Walker (Orthoptera: Gryllidae: Oecanthinae)

Previous studies on tree crickets have demonstrated female choice of males based on size and courtship feeding but less is known about sexual selection under conditions of direct mating competition. I studied courtship, aggression and mating of the black-horned tree cricket Oecanthus nigricornis (Walker) to test size-related sexual selection under conditions of direct sexual competition. Results show that larger individuals of both sexes mated more frequently than their smaller counterparts, and this was due to the ability of large individuals to out compete rivals. Large males achieved the advantage by aggressively reducing courtship by small males, whereas large females responded to male courtship more quickly but with little aggression. Although there was no evidence here for mate choice, there were advantages for having larger mates; fecundity increased with female size and spermatophores (which females consume after mating) increased with male size. Size of the specialized metanotal courtship gift, however, was not related to male size.     

Love Is Blind: Indiscriminate Female Mating Responses to Male Courtship Pheromones in Newts (Salamandridae)

Internal fertilization without copulation or prolonged physical contact is a rare reproductive mode among vertebrates. In many newts (Salamandridae), the male deposits a spermatophore on the substrate in the water, which the female subsequently takes up with her cloaca. Because such an insemination requires intense coordination of both sexes, male newts have evolved a courtship display, essentially consisting of sending pheromones under water by tail-fanning towards their potential partner. Behavioral experiments until now mostly focused on an attractant function, i.e. showing that olfactory cues are able to bring both sexes together. However, since males start their display only after an initial contact phase, courtship pheromones are expected to have an alternative function. Here we developed a series of intraspecific and interspecific two-female experiments with alpine newt (Ichthyosaura alpestris) and palmate newt (Lissotriton helveticus) females, comparing behavior in male courtship water and control water. We show that male olfactory cues emitted during tail-fanning are pheromones that can induce all typical features of natural female mating behavior. Interestingly, females exposed to male pheromones of their own species show indiscriminate mating responses to conspecific and heterospecific females, indicating that visual cues are subordinate to olfactory cues during courtship.     

Females of the Cellar Spider Discriminate Against Previous Mates

Mate choice for novel partners should evolve when remating with males of varying genetic quality provides females with fitness‐enhancing benefits. We investigated sequential mate choice for same or novel mating partners in females of the cellar spider Pholcus phalangioides (Pholcidae) to understand what drives female remating in this system. Pholcus phalangioides females are moderately polyandrous and show reluctance to remating, but double‐mated females benefit from a higher oviposition probability compared to single‐mated females. We exposed mated females to either their former (same male) or a novel mating partner and assessed mating success together with courtship and copulatory behaviours in both sexes. We found clear evidence for mate discrimination: females experienced three‐fold higher remating probabilities with novel males, being more often aggressive towards former males and accepting novel males faster in the second than in the first mating trial. The preference for novel males suggests that remating is driven by benefits derived from multiple partners. The low remating rates and the strong last male sperm precedence in this system suggest that mating with novel partners that represent alternative genotypes may be a means for selecting against a former mate of lower quality.