Payoffs of the two alternative nesting tactics in the African dung beetle, Scarabaeus catenatus

1. Scarabaeus catenatus is a ball-rolling scarab in the subfamily Scarabaeinae. This species, however, makes use of two tactics for nest building: rolling and tunnelling. The tunnelling tactic differs substantially from the rolling tactic in that (1) it always involves repeated movements to and from the dung source and the nest, whereas rolling does not, and (2) it involves a shorter distance between the two sites.
2. Brood-nest founders were usually males and less often females, with about 25% adopting the rolling tactic and 75% adopting the tunnelling tactic. During nest building, the founder paired off with a scarab of the opposite sex, and they co-operated in the work. The female made one to four brood balls from the dung in the nest, each of which contained one egg.
3. Each scarab seemed to be able to employ both tactics. The tactic employed was independent of an individual's status, e.g. body size and timing of nest founding.
4. The rolling tactic offered only male founders a greater nest-defence success than the tunnelling tactic due to a lower intrusion into the rolled nest and a higher intensity of male–male fighting. The tunnelling tactic offered both male and female founders a larger number of brood balls than the rolling tactic because it enabled scarabs to take a larger amount of dung into the nest.
5. The reproductive success for the two tactics was estimated from the product of nest-defence success and the number of brood balls. As a result, the two tactics had equal fitness payoffs for males, but unequal payoffs for females.
6. The results suggest that male alternation of tactics is controlled by a mixed strategy. Female alternation, however, cannot be explained by mixed strategy, alternative strategies or conditional strategy.     

The High Price of Success: Costs of Mating Effort in Male Primates

While males are generally the low investing sex when it comes to offspring care, males of many species experience intense and persistent mating effort. Mating effort incurs a variety of costs which are expected to have non-negligible effects on fitness, as well as how reproductive tactics are selected and investment in mating activity is moderated over time. This special issue features contributions investigating the costs of male mating effort across primate species. Here, we place these exciting new works in context, addressing the specific types of mating effort expected for male primates and the significance of these costs for our understanding of primate life histories and socioecology.     

The Foraging Costs of Mating Effort in Male Chimpanzees (Pan troglodytes schweinfurthii)

Costs of mating effort can affect the reproductive strategies and lifetime fitness of male primates, but interspecific and interindividual variation in the magnitude and distribution of costs is poorly understood. Male costs have primarily been recognized in seasonally breeding species that experience concentrated periods of mating competition. Here, we examine foraging costs associated with male mating effort in chimpanzees (Pan troglodytes schweinfurthii), a polygynandrous species, in which mating opportunities occur intermittently throughout the year. To quantify male feeding, aggression, and mating, we conducted focal follows on 12 males in a wild community (Kanyawara, Kibale National Park, Uganda) for 11 mo. Males fed less on days when high-value mating opportunities (estrous parous females) were available than on days without any mating opportunities. Reductions in feeding time were related to increased rates of aggression and copulation, indicating that the proximate cause of changes in male foraging was mating effort. Surprisingly, however, there was no relationship between dominance rank and the extent to which feeding time was reduced. High costs of mating effort may reduce the degree of reproductive skew and limit the use of possessive tactics in chimpanzees. We suggest that male bonding in chimpanzees may be favored not only for its benefits but because intragroup competition is so costly. Our results complement the available data on mammals, and primates in particular, by showing that mating effort can have measurable foraging costs even in species, in which breeding is aseasonal and only moderately skewed.     

Variation in testosterone levels and male reproductive effort: Insight from a polygynous human population

Recent evidence suggests that, in humans, variations in testosterone (T) levels between men reflect their differential allocation in mating versus parenting efforts. However, most studies have been conducted in urbanized, monogamous populations, making generalizations from them questionable. This study addresses the question of whether indicators of male reproductive effort are associated with variations in salivary T levels in a polygynous population of agriculturists in rural Senegal. We first show that pair-bonding and/or transition to fatherhood is associated with T profiles: married fathers (N = 53) have lower morning and afternoon T levels than unmarried non-fathers (N = 28). Second, among fathers, individual differences in parenting effort, as well as variations in mating effort, predict morning T levels. Indeed, men highly investing in parental care show lower morning T levels. Moreover, among men under 50, polygynous men show higher morning T levels than monogamous men. Taken together with previous results in monogamous settings, these findings suggest that the endocrine regulation of reproductive effort is probably a general feature of human populations.     

Interspecific variation in spawning time and male mating tactics of the parrotfishes on a fringing coral reef at Iriomote Island,Okinawa

Spawning time and male mating tactics of parrotfishes (family Scaridae) were investigated on a fringing coral reef at Iriomote Island, Okinawa. Spawning was observed in 14 species, and more frequently in more abundant species such as Chlorurus sordidus, Scarus rivulatus and Chlorurus bowersi. At the reef-edge spawning site, C. bowersi spawned at high tide, C. sordidus spawned both at high tide and in the early morning, whereas Calotomus carolinus and most of the Scarus species such as S. rivulatus spawned only in the early morning, mostly 0630–0830 h. Spawning only in the early morning irrespective of tide phase and moon age has seldom been reported from the scarid species of other localities. It is suggested that spawning in the early morning would be adaptive in species such as S. rivulatus, which migrated considerable distances (ca. 500 m) to the inshore feeding sites, in order to minimize feeding losses due to migration. For male mating tactics, pair spawning by territorial TP (terminal phase) males occurred in all 14 species, and streaking and group spawning by nonterritorial small IP (initial phase) males were seen more frequently in more abundant species. Moreover, group spawning by nonterritorial TP males, which were larger than the IP males but smaller than the territorial TP males, frequently occurred in S. rivulatus. Such mating tactics of TP males have not been reported from Scaridae.     

Patterns of Niche Partitioning and Alternative Reproductive Strategies in an East African Dung Beetle Assemblage

Sympatric species of coprophagus dung beetles rely on essentially the same resource for provisioning broods, which raises the question of how local species diversity is maintained. Interspecific competition may be mitigated to some extent by large-scale spatial (e.g., habitat type) and temporal (e.g., seasonal) variation in activity. Niche partitioning also occurs at the scale of individual dung pads. We examined the extent to which inter- and intraspecific variation in adult morphology and behavior contribute to spatial and temporal partitioning above and below the dung pad in the dung beetle community of Kibale Forest, Uganda. Excavations in the vicinity of dung pads revealed significant heterogeneity among species and guilds in the lateral distance between tunnels and dung, and in tunnel depth. Patterns of succession and diel activity demonstrated strong temporal dissociations among guilds. In one species, Onthophagus multicornis, we found a bimodal distribution of male horn sizes and a tunneling pattern consistent with alternative reproductive tactics.     

A cost of maternal care in the dung beetle Onthophagus taurus?

Parental care theory assumes that investment in current offspring will trade against future investment. A number of field studies on birds have used clutch size manipulations to demonstrate a survival cost to chick rearing. However, such studies do not account for costs accrued during earlier stages of reproduction because not all aspects of reproductive effort are manipulated by varying the number of nestlings. In this study, we investigate the effect of reproductive effort on female survival in the dung beetle, Onthophagus taurus. By experimentally manipulating mating status and dung availability, we demonstrate that virgin females survive longer than mated females and that the survival of mated females was negatively associated with the number of brood masses produced. Using a novel manipulation of the mating system, we separated the effects of egg production and maternal care on female survival. Previously, we have shown that females provisioning with the assistance of a major male provide relatively less care than unassisted females. However, paternal assistance did not alter the number of brood masses produced and hence the amount of reproductive effort that was allocated to egg production. Therefore, our finding that female survival was increased when receiving paternal assistance provides, to our knowledge, the first definitive evidence that maternal care reduces female lifespan. These results are of major importance to theoretical models on the evolution of parental care.     

Alternative mating tactics in the yellow dung fly: resolving mechanisms of small-male advantage off pasture

Recent work suggests that the yellow dung fly mating system may include alternative patroller–competitor mating tactics in which large males compete for gravid females on dung, whereas small, non-competitive males search for females at foraging sites. Small males obtain most matings off pasture, yet the behavioural mechanism(s) giving rise to this pattern are unknown. We investigated the male and female behaviours that determine mating success in this environment by conducting field mating experiments and found small males to benefit from several attributes specific to the off-pasture mating environment. First, small males from foraging sites exhibited higher mating propensity, indicating that large males away from dung may be depleted of energy and/or sperm. Second, small males were more discriminating, being significantly less likely to attempt with non-gravid females, which are absent on dung but common off pasture. Third, non-gravid females were generally more likely to actively struggle and reject mating attempts; however, such behaviours occurred disproportionately more often with large males. Female Scathophaga stercoraria thus appear to preferentially mate with small males when off pasture. These findings challenge assumptions about male–female interactions in systems with alternative mating tactics and reveal hidden processes that may influence selection patterns in the field.     

Predation risk and alternative mating tactics in male Trinidadian guppies (Poecilia reticulata)

In the guppy (Poecilia reticulata), males have two alternative mating tactics. Individual males may either display to a receptive female prior to attempting to copulate with her or attempt to quickly sneakcopulate with a female without first displaying to her or without a prior receptive response from her. In this study, I experimentally investigated the effects of simulated local increases in the risk of predation (in the form of a cichlid fish predator model in situ) on the mating tactics used by free-ranging male guppies in two typical macrohabitats (riffle and pool) of a Trinidadian river. Focal male guppies displayed to females significantly less often on average, and conversely attempted sneak copulations more often, in the presence of the predator model than in its absence; this pattern was similar for both habitats. These fish therefore performed a lower proportion of sigmoid displays and increased their sneaky mating attempts when the apparent risk of predation had increased locally. This predator-mediated response is consistent with a trade-off between mating success and risk of mortality due to predation. The results are the first to confirm risk-sensitive mating behaviour in free-ranging male guppies within a population, and demonstrate the potential importance of predators in influencing the relative use of alternative mating tactics in this species on a microgeographical scale in the wild.     

Stepparental Behavior as Mating Effort in Birds and Other Animals

In many animal species, widowed or divorced parents may remate before young of the prior union are independent. In such circumstances, stepparents may kill their predecessors' offspring, may tolerate them without providing care, or may invest in them more or less as genetic parents do. Rohwer proposed that all three of these responses may be understood as mating tactics, adapted to different social and ecological circumstances. We discuss the selection pressures that would favor each of these alternatives and review relevant evidence on nonhuman stepparenting, especially in birds. Stepparental tolerance and (partial or full) care, which are the predominant human responses, are common in nonhuman animals too, and in many cases there is evidence supporting their interpretation as stepparental mating effort adaptations. In general, however, this interpretation is not as well established for tolerance and care as it is for stepparental infanticide. Because tolerance and care are not distinct modes of behavior peculiar to stepparents, the hypothesis that they are nonadaptive by-products of parental psychology often remains tenable. We discuss the kinds of evidence needed to choose between by-product and stepparental adaptation hypotheses.     

The function of nuptial feeding in insects: a review of empirical studies

Nuptial feeding encompasses any form of nutrient transfer from the male to the female during or directly after courtship and/or copulation. In insects, nuptial gifts may take the form of food captured or collected by the male, parts, or even the whole of the male's body, or glandular products of the male such as salivary secretions, external glandular secretions, the spermatophore and substances in the ejaculate. Over the past decade, there has been considerable debate over the current function of nuptial feeding in insects. This debate has centred on the issue of whether nuptial gifts function as paternal investment (i.e. function to increase the fitness and/or number of the gift-giving male's own offspring) or as mating effort (i.e. function to attract females, facilitate coupling, and/or to maximize ejaculate transfer), although the two hypotheses are not mutually exclusive. In the present article, evidence for the potential of nuptial gifts to function as either paternal investment, mating effort, or both is reviewed for each form of nuptial feeding in each insect taxon for which sufficient data are available. Empirical evidence suggests that many diverse forms of nuptial feeding in different insect taxa function, at least in part, as mating effort. For example, nuptial prey and salivary masses in the Mecoptera, regurgitated food in Drosophila (Diptera), hind-wing feeding in Cyphoderris (Orthoptera) and the secretion of the male's cephalic gland in Neopyrochroa (Coleoptera) and Zorotypus (Zoraptera) appear to function to entice females to copulate and/or to facilitate coupling. Nuptial prey and salivary masses in the Mecoptera also appear to function to maximize ejaculate transfer (which is also a form of mating effort), as do nuptial prey in Empis (Diptera), external glandular secretions in Oecanthus and Allonemobius (Orthoptera) and the spermatophylax in gryllids and tettigoniids (Orthoptera). Large spermatophores in, for example, the Lepidoptera and Coleoptera, also appear to be maintained by selection on the male to maximize ejaculate transfer and thereby counter the effects of sperm competition. In contrast to the large amount of evidence in support of the mating effort hypothesis, there is a relative lack of good evidence to support the paternal investment hypothesis. Certain studies have demonstrated an increase in the weight and/or number of eggs laid as a result of the receipt of larger gifts, or a greater number of gifts, in tettigoniids, gryllids, acridids, mantids, bruchid beetles, drosophilids and lepidopterans. However, virtually all of these studies (with the possible exception of studies of the spermatophylax in tettigoniids) have failed to control adequately for hormonal substances in the ejaculate that are known to affect female reproductive output. Furthermore, in at least four tettigoniids (but not in the case of two species), three lepidopterans, a drosophilid and probably also bruchid beetles and bittacids, evidence suggests that the male has a low probability of fertilising the eggs that stand to benefit from his nuptial gift nutrients. Therefore, the hypothesis that paternal investment might account for the function of nuptial gifts in general is not supported.     

Dung beetles as secondary seed dispersers in a temperate grassland

The two-phase dispersal event in which dung beetles move seeds after endozoochory is often assumed to be advantageous for plant regeneration. Because seeds are expected to end up in favourable and safe germination sites, it is considered as an example of directed dispersal. However, literature so far is restricted to tropical rain forest ecosystems, while data for temperate regions are lacking. In this study, the effect of dung beetles on seedling establishment of endozoochorically dispersed seeds is evaluated for a temperate grassland ecosystem. We performed a field experiment in which cages excluded dung beetles from horse and cattle dung samples with mixed-in grass seeds. Seed germination from these samples was significantly higher than that from samples which were accessible to dung beetles. This indicates that the effect of dung beetles on short-term seedling establishment was negative, which contrasts with the patterns found for large-seeded species used in tropical studies. This is most likely attributed to the lack of roller species and the larger depth at which tunneling Geotrupes species bury seeds.     

Seasonal changes in reproductive and physical condition, sexual dimorphism, and male mating tactics in the jewelled blenny Salarias fasciatus

To investigate seasonal changes in reproductive and physical condition, sexual dimorphism, and mating tactics in the jewelled blenny Salarias fasciatus, monthly collections were conducted on the fringing reef in northern Okinawa, Japan. Monthly variation in the female gonadosomatic index suggested that the breeding season of this species is from April to June. The physical condition and hepatosomatic index of both sexes considerably deteriorated during the course of the breeding season. Such declines may be primarily due to egg production in females and brood care in males. Males’ elongated anal spines were longer than females’, but no other apparent sexual dimorphisms were detected, indicating that S. fasciatus exhibits a low degree of sexual dimorphism. Examinations of testes size and the testicular gland area during the estimated breeding season revealed that the gonadosomatic index values of smaller males did not differ from those of larger males. This result may differ from other blenny species that exhibit alternative male reproductive tactics. However, patterns of male testicular glands of S. fasciatus were similar to those of other blennies with alternative tactics, i.e., smaller males had poorly developed testicular glands compared with larger males.     

Divergent life history traits in two closely related species of Simocephalus (Cladocera: Daphnidae) inhabiting lentic environments in Venezuela

We describe life history tactics under laboratory conditions of two species of cladocerans of the genus Simocephalus. The populations live in two habitats with different characteristics. S. acutirostratus was isolated from a small temporary pool without fish. S. latirostris was found in the marginal vegetation of a reservoir with fish. Their life history was monitored for differences in traits such as clutch size, neonate size, age distribution, reproductive effort and adult survival. Our results show that S. acutirostratus (the larger-sized species) grows until it reaches the optimal foraging size and then begins to reproduce, while S. latirostris (the smaller-sized species) starts breeding before reaching the optimal foraging size, allocating energy mainly to reproduction. We explore the possibility that divergences in life history may arise as a response to environmental stress such as that produced by fishes.     

Repeated matings offset costs of reproduction in female crickets

Summary Courtship food gifts can be a significant source of nutrition to females and costly for males to produce; hence, costs of reproduction should be reduced for multiple-mating females and increased for multiplemating males in a gift-giving species. We tested this hypothesis by experimentally manipulating mating opportunities of males and females of two cricket species,Gryllodes sigillatus, a gift-giving species andGryllus veletis, a non-gift-giving species. Females of both species consume the externally attached spermatophore after mating, but inG. sigillatus, the sperm-containing ampulla is accompanied by a large gelatinous spermatophylax. In both species, survival of mated females given limited access to males was reduced relative to virgin females, thus suggesting a cost of reproduction to females. However, females given unlimited mating opportunities lived as long as virgins and also produced significantly more offspring than limited-access females. These results suggest that benefits of repeated matings, particularly those arising through spermatophore consumption, offset costs of reproduction in females. Lack of a treatment by species interaction suggests that females of both species derive nutritional benefits through spermatophore consumption, and that any additional advantage to the consumption of the spermatophylax inG. sigillatus is offset by more frequent mating byG. veletis females. In contrast to females, varying mating opportunities had no effect on male survival, suggesting that mating effort is not very costly to males. Male survival increased linearly with body mass but only when males were food-deprived, suggesting that larger males possess greater initial energy reserves to sustain their longevity when food-stressed.     

INBREEDING AND ADVERTISEMENT CALLING IN THE CRICKET TELEOGRYLLUS COMMODUS: LABORATORY AND FIELD EXPERIMENTS

If sexually selected traits reveal a male's heterozygosity or condition to females, then such traits should exhibit declines with inbreeding. We tested this by examining the effect of inbreeding on advertisement calling in male crickets Teleogryllus commodus. We investigated the effect of one generation of full‐sibling mating on calling effort and fine‐scale call structure. Inbreeding reduced calling effort but had no effect on call structure. We then compared the attractiveness of inbred and outbred calls in the field by monitoring how many wild females were attracted to each call type. From the field data, we conducted a selection analysis to identify the major axes of linear and nonlinear multivariate sexual selection on call structure. A comparison of multivariate attractiveness of inbred and outbred calls along each major axis of selection revealed no difference in attractiveness. Our results suggest that inbred male calls have a fine‐scale structure that is no less attractive to females than that of outbred calls. However, because inbred males call less often, and female T. commodus prefer males with a higher calling effort, inbred males will suffer reductions in mating success. Females who base mate choice on call rate are therefore using a signal correlated with male heterozygosity and/or condition.     

Marriage Markets and Male Mating Effort: Violence and Crime Are Elevated Where Men Are Rare

The negative social outcomes in populations with male-biased sex ratios are a growing concern. In general, the expectation is of heightened violence as a result of excess men engaging in antisocial behavior and crime, thereby threatening societal stability. While intuitive, these claims are largely unsupported in the literature. Using mating market theory as our guide, we examine indicators of male mating effort, including (1) violent competition between men (homicide, aggravated assault) and (2) indicators of uncommitted sexual behavior (rape, sex offenses, and prostitution). Our unit of analysis is U.S. county-level data. We find that counties with more men have lower rates of crime and violent behavior. Our findings challenge conventional claims of male excess leading to elevated levels of violence. Instead, in support of mating market predictions, we find that criminal and violent behavior related to male mating effort is least common in male-biased sex ratios. We discuss the implications of our findings for public policy regarding incarceration and criminal behavior.     

Costs of Mate Guarding and Opportunistic Mating Among Wild Male Japanese Macaques

A trade-off relationship between mating and feeding effort is important when considering reproductive strategies of long-lived species. I compared the influence of male sexual activities, female mate-choice behaviors and the daily activity budget on male mating success among males in a group of wild Japanese macaques (Macaca fuscata yakui) on Yakushima Island. The 1st-ranking male, which had immigrated into the troop at this rank, more frequently approached peri-ovulatory females, spent more time grooming peri-ovulatory females and in mounting series and spent less time feeding than subordinate males did. The 1st-ranking male attained the highest mating success as a result of his high expenditure of time and energy in sexual behaviors directed toward peri-ovulatory females. Mating success of subordinate males did not relate to the amount of sexual effort, but instead to the frequency of female approaches, female rush toward males and the number of peri-ovulatory females within the group. The pattern of intermale competition shifted from nearly contest competition to scramble competition as the number of peri-ovulatory females in the group increased. Feeding time of subordinate males did not vary between the days when they copulated and the days when they did not. The findings demonstrate that mate guarding in the 1st-ranking male is a high-cost mating tactic, while opportunistic mating in subordinate males is a low-cost mating tactic. The differences in male mating tactics are probably related to male life history and to the formation of groups with a high socionomic sex ratio.