Inbreeding and local mate competition in the ant Cardiocondyla batesii

The ant species Cardiocondyla batesii is unique in that, in contrast to all other ant species, both sexes are flightless. Female sexuals and wingless, ergatoid males mate in the nest in autumn and young queens disperse on foot to found their own colonies in spring. The close genetic relatedness between queens and their mates (r_qm=0.76±SE 0.12) and the high inbreeding coefficient (F=0.55; 95%CI 0.45–0.65) suggest that 83% of all matings are between brothers and sisters. As expected from local mate competition theory, sex ratios were extremely female biased, with more than 85% of all sexuals produced being young queens. Despite the common occurrence of inbreeding, we could not detect any adult diploid males. Though the probability of not detecting multiple mating was relatively high, at least one-third of all queens in our sample had mated more than once. Multiple mating to some extent counteracts the effects of inbreeding on worker relatedness (r_ww=0.68±SE 0.05) and would also be beneficial through decreasing diploid male load, if sex was determined by a single locus complementary system.Communicated by L. Sundström     

Alternative male mating tactics in a cichlid, Pelvicachromis pulcher: a comparison of reproductive effort and success

Pelvicachromis pulcher is a small African cichlid which breeds in holes. Males may either reproduce monogamously (pair males), polygynously (harem males), or be tolerated as helpers in a harem territory (satellite males). These helpers share in defence of the territory against conspecifics, heterospecific competitors and predators. There are two male colour morphs that are fixed for life and are apparently genetically determined. These differ in their potential mating strategy. Red morph males may become harem owners, while yellow morph males may become satellite males, and males of both morphs may alternatively pair up monogamously. We compared the reproductive effort and success of these three male reproductive strategies. Effort was measured as attack rates, time expenditure and the risk of being injured or killed when attacking competitors or predators of three sympatric fish species. Reproductive success was measured by observing how many eggs were fertilized by each male when this was possible, and by using genetic markers. The number of fry surviving to independence of parental care was used as a criterion of success. The reproductive success of harem males was 3.3 times higher than that of pair males and 7 times higher than that of the average satellite male. Dominant satellite males, however, were as successful as monogamous pair males, using the measure of fertilized eggs. To our knowledge, this has not been found previously in any fish species. Both harem and pair males had lower parental defence costs per sired offspring, however, than males using the alternative satellite tactic. Defence effort was significantly related to the risk of injury. Received: 17 January 1996 / Accepted after revision: 9 June 1997     

Male reproductive success in free-ranging feral horses

In the social organization of feral horses, adult males compete to monopolize groups or bands of females, sometimes called harems. Alternative male strategies are to remain alone or with other bachelors or, less commonly, to accept subordinate status within a harem. The hypothesis that dominant harem stallion status confers a reproductive advantage was tested in free-ranging feral horses. The presence of foals in harems headed by vasectomized (VSX) versus intact stallions was used to assess the ability of these stallions to control reproduction in their harems. Of harems headed by VSX stallions, 17 and 33% contained foals during years 2 and 3 post-treatment, respectively. In contrast, 86 and 80% of harems headed by non-VSX stallions contained foals in those years. Acquisition of pregnant mares appeared more likely than sneak copulations by bachelor stallions to account for foals in harems with a single stallion. However, most foals were born into harems that included a subordinate stallion, an occurrence that was undoubtedly exacerbated by the extended breeding season resulting from the sterility of the harem stallion. Thus, in comparing alternative reproductive tactics, bachelors appeared less successful than subordinate stallions within a harem. However, the highest reproductive success was achieved by the harem stallion, further demonstrating that alternative tactics are not equally profitable. Received: 13 July 1999 / Received in revised form: 24 July 1999 / Accepted: 24 July 1999     

Male-male interference competition decreases spawning rate in the European bitterling (<Emphasis Type="Italic">Rhodeus sericeus</Emphasis>)

We investigated the consequences of male-male interference competition associated with alternative male mating tactics in a freshwater fish, the European bitterling (Rhodeus sericeus). Male bitterling defend territories around living mussels and attract females to lay their eggs in the gill cavities of mussels. We experimentally manipulated spawning-site abundance and male density at two spatial scales. We showed that the total number of eggs spawned by females was constrained by the number of mussels available for oviposition. The effect was mediated by behavioral interactions among competing males because of variation in the Operational Sex Ratio (OSR) in close proximity to a mussel and not by a direct limitation in mussel capacity to accommodate the eggs. Both total and local male densities affected spawning behavior, and interacted in their effect on female spawning rate. Territorial male aggression caused courtship interruptions that prolonged the time until successful spawning and increased with male density. However, territoriality broke down at the highest male density, with a consequent stabilizing effect on spawning rate.Communicated by J. Krause     

Multiple paternity in the cooperatively breeding fish <Emphasis Type="Italic">Neolamprologus pulcher</Emphasis>

In cooperative breeders, mature males may compete for fertilizations. In this study, we measured the degree of multiple paternity in a natural population of a cooperatively breeding fish. Neolamprologus pulcher (Perciformes: Cichlidae) is a highly social cichlid endemic to Lake Tanganyika. We used highly variable microsatellite loci to survey 12 groups with an average number of 10.6 brood care helpers per group and a total of 43 offspring (mean 3.6 per brood). In 11 of 12 groups, all young were assigned to the dominant female. The dominant male sired all offspring in three groups, part of the offspring in four groups, and in five groups, he had no paternity at all. In total, 44.2% of young were not fathered by the current male territory owner. Multiple paternity was found in 5 of 12 broods (41.7 %), with 8 of 35 young (22.9 %) being sired by males other than the respective territory owners. This is an exceptionally high rate of extra-pair paternity among cooperatively breeding vertebrates. Neither helpers present in these territories during collection nor neighbouring males were unequivocally assigned to have sired these extra-pair young. However, behavioural observations suggest that male helpers may have produced these young before being expelled from the territory in response to this reproductive parasitism. We discuss these results in the light of reproductive skew theory, cooperative breeding in vertebrates and alternative reproductive tactics in fish.     

The adaptive significance of male polymorphism in the acarid mite Caloglyphus berlesei

Summary Two forms of males occur in the acarid mite, Caloglyphus berlesei. One of them uses its thickened third pair of legs to kill other males; the other, without modified legs, does not attack other males. Previous studies have shown that the form of the male is environmentally determined: fighter males develop only at low population densities and their development can be suppressed by substances emanating from dense colonies. In this study, the duration of development, longevity and virility of the two male morphs were measured. The only significant difference was that fighters had a shorter development time between the two last moults. In another experiment, two groups of colonies were maintained under the same conditions but had different numbers of individuals: small colonies contained 2 fighters and 2 non-fighters, whereas large colonies contained 30 males of each type. The relative reproductive success of both morphs was estimated from numbers of matings. In small colonies, the estimated reproductive success (ERS) was significantly higher for fighters than for non-fighters, mainly because in over 50% of these colonies a single fighter male managed to kill all the rival males and monopolize the females. The opposite was true in large colonies, in which non-fighters achieved significantly higher ERS, mainly because the fighters were killed in fights more often than the non-fighters. This implies that the ratio of costs to benefits obtained by adopting the fighter-male strategy increases with the number of rival males, which may explain conditional male development in C. berlesei.     

Genetic analysis of parentage within experimental populations of a male dimorphic beetle,<Emphasis Type="Italic"> Onthophagus taurus</Emphasis>, using amplified fragment length polymorphism

We used the multilocus DNA fingerprinting technique, amplified fragment length polymorphism, to examine parentage of 902 offspring from eight experimental populations of the dung beetle, Onthophagus taurus. The males of this species exhibit a suit of morphological and behavioural traits that characterise alternative mating tactics. Hornless minor males sneak copulations with females that are guarded by horned major males. Our aims were to provide a prospective assessment of the potential role of frequency dependence in the maintenance of alternative mating tactics, to assess the levels of polyandry, and to determine the patterns of sperm usage by multiply mated females. The average proportion of offspring sired by major and minor males did not co-vary with the relative frequency of each morph present in experimental populations. However, there was some indication that the effective mating frequency (number of females producing offspring sired by a given male) of major and minor males may exhibit frequency dependence. Mating success of both male types declined with increasing numbers of major males. Paternity was positively associated with effective mating frequency. Females produced offspring sired from between one and eight males and, on average, paternity was distributed equally amongst a females mates, regardless of the number of males mated. Differences in fertilisation success among males were not associated with alternative male phenotypes. Neither did a males fertilisation success depend on his genetic dissimilarity with the female. These results are discussed in the context of the evolution of alternative mating strategies, and mechanisms of postcopulatory sexual selection.Communicated by N. Wedell     

Effectiveness of territorial polygyny and alternative mating strategies in northern fur seals, <Emphasis Type="Italic">Callorhinus ursinus</Emphasis>

We conducted a 6-year longitudinal behavioral and genetic investigation of a highly polygynous pinniped, the northern fur seal (Callorhinus ursinus), to determine the contribution of terrestrial polygyny to male fertilization success and to assess the occurrence of alternative mating strategies. Genetic samples from 37 adult males, 50 adult females, and 85 pups were collected and genotyped using five polymorphic microsatellite loci. Pup paternity was assigned using Cervus 2.0 at 99% confidence level. Paternity of 83 pups (98%) was assigned to terrestrial males who held territories or stayed temporarily in the study area during the breeding season when fertilization occurred. For 56 pups of which attendance records of their mothers were available, paternity of 45 pups (80%) was assigned to the associate males in whose territory their mothers stayed during the perioestrus period. In addition to defending breeding territories, territorial males have often been observed attempting to forcibly abduct adult females from adjacent territories (female stealing): We observed a total of 95 such cases, in which the stealers had significantly fewer females than the territorial males from whose territories they stole females. Our results indicate that terrestrial resource-defense polygyny is the major mating system in this species and that nonassociated paternity occurs mostly as a result of alternative mating strategies of less successful males. Male northern fur seals thus appear to adopt conditional alternative strategies that depend on their current social status to maximize their life-time reproductive success.     

Sex,food and conflicts: nutrition dependent nuptial feeding and pre-mating struggles in scorpionflies

Female and male reproductive interests often differ. In species in which matings are accompanied by a transfer of resources valuable for both participants, such as nuptial prey gifts, conflicts may readily occur. Scorpionflies may use alternative mating tactics. One is to offer a prey item (dead arthropod) to females in exchange for mating. This prey gift tactic includes a conflict because a male must decide on whether to offer the gift rather than to fight the female and consume the gift. The outcome may depend on the nutritional status of both males and females. Males may be more willing to give if they themselves are satiated and the condition of the females may influence the payoff from the males’ investment. Similarly, females may be more willing to accept food gifts if they are in poor nutritional condition. In this study of the scorpionfly Panorpa cognata, I experimentally manipulated the feeding history of both males and females. I observed the outcome of the direct interactions that followed when males that were holding prey were approached by females. I found that well-fed males offered the food gift sooner than males in poor nutritional condition that fed extensively on the food item before offering. Female condition had no significant influence on whether prey items were offered by males or accepted by females. I also found that well-fed males rarely searched for prey to pursue the prey gift tactic in courtship. Thus, the prey tactic does not seem to be the males’ first option.     

Sperm swimming speed and energetics vary with sperm competition risk in bluegill (<Emphasis Type="Italic">Lepomis macrochirus</Emphasis>)

Under sperm competition, a males fertilization success depends largely on the ejaculate characteristics of competing males. Theoretical models predict that, in external fertilizers, increased risk of sperm competition should result in selection for increased sperm swimming speed. To test this prediction, we studied the behavior of sperm from parental and sneaker male bluegill (Lepomis macrochirus), a fish species characterized by high levels of cuckoldry due to alternative reproductive tactics of males (parentals and cuckolders). Because cuckolders (sneakers and satellites) always spawn in the presence of a parental male, but the reverse is not true, cuckolders experience the greater risk of sperm competition. We show here that the spermatozoa of sneakers have faster initial swimming speeds but shorter periods of motility than the sperm of parental males. Moreover, we show that sperm swimming speeds shortly after activation (when most fertilization occurs) are correlated with starting ATP levels in spermatozoa, suggesting that sperm competition has selected for higher energetic capacity in the sperm of sneakers. Thus, the higher energetic capacity and initial swimming speed of sneaker sperm may explain why, despite having fewer sperm per ejaculate than parentals, sneakers fertilize more eggs than parental males when they compete to fertilize a clutch of eggs.Communicated by L.W. Simmons     

A dynamic model of mating behavior in digger wasps: the energetics of male-male competition mimic size-dependent thermal constraints

I develop a state-based dynamic model of behavior to demonstrate that size-dependent differences in temperature tolerances are not necessary to account for the activity of small male digger wasps late in the day. In the model, males defend or patrol the nesting area, wait near nests, or feed away from the nesting area depending on time of day, energy reserves and size rank. I assume a large male competitive advantage, so mating opportunities decrease with size rank for territorial or patrolling males and are rare for all waiting males; the costs of patrolling or defense are higher than the costs of waiting. If energy reserves of all males are initially small, all males alternate feeding and territorial or patrolling behavior. If energy reserves are initially large, large males patrol or maintain territories until they risk starvation and leave the area to feed. At this time, smaller males that have conserved their resources by waiting and feeding may defend territories or patrol. I simulate the behavior of three populations representing two species of Microbembex by assuming large initial energy reserves for populations in which males were territorial and small initial reserves for populations in which males patrolled, and then convert the predicted time of activity to temperature using local regressions from field studies. Temporal patterns in the activity of large and small males were similar to those actually observed, and relationships between size and temperature predicted by the model corresponded to most observations and were sometimes positive. Thus, the delayed activity of smaller males does not correspond to activity at higher temperatures and is probably not attributable to size-dependent thermal tolerances, but may represent a temporal displacement of mating activity due to intra-sexual competition and mediated by energetics. The model makes testable predictions on the timing of feeding and depletion of energy reserves in relation to size and initial energy state, and suggests how differences among species may influence the temporal and spatial organization of male mating behavior. Received: 27 February 1997 / Accepted after revision: 26 July 1997     

Male size and mating tactic influence proximity to females during sperm competition in bluegill sunfish

Many fishes are characterized by intense sperm competition between males that use alternative mating tactics. In externally fertilizing fishes, males’ proximity to females during spawning can be an important determinant of fertilization success. Here, we assess how mating tactic, body length, speed during streak spawns, and periphery cover affect males’ proximity to females during sperm competition in the externally fertilizing bluegill (Lepomis macrochirus). Bluegill are characterized by three mating tactics referred to as parental, sneaker, and satellite. Parentals are territorial and construct nests, while sneakers use a streaking behavior, and satellites use female mimicry to steal fertilizations from parentals. We show that a small body length is important for sneakers but not for satellites to obtain a close position to the female during spawning. Specifically, smaller sneakers obtain a closer position to females than larger sneakers in part by positioning themselves closer on the periphery of a parental’s nest before streaking but show no difference in the speed at which they streak. The amount of peripheral vegetation around a parental’s nest did not appear to affect proximity of sneakers to females, and there was no relationship between the amount of peripheral vegetation and the frequency of intrusions by either sneakers or satellites. Finally, parentals were farther from the female when a sneaker or satellite intruded than when they spawned alone with the female.     

Patterns of paternity skew in Formica ants

Reproductive skew among cooperatively breeding animals has recently attracted considerable interest. In social insects reproductive skew has been studied in females but not in males. However, cooperative breeding of males occurs when two males mate with the same queen and father offspring. Here we present the first analysis of comparative data on paternity skew in ants. We show that, across seven species of Formica ants, the average skew in paternity among worker offspring of doubly mated queens is negatively correlated with the population-wide frequency of multiple (mostly double) mating. We also demonstrate that this trend is relatively robust in additional analyses taking phylogenetic relationships between species into account. The observed trend is opposite to the one normally found in non-social insects with second-male precedence through sperm displacement, but agrees with predictions based on queen-male conflict over sperm allocation as a consequence of facultative, worker controlled, sex allocation – an interpretation which assumes first-male precedence. However, alternative (but not mutually exclusive) explanations are possible and further studies will be needed to discriminate between these alternatives. Received: 16 May 1997 / Accepted after revision: 26 September 1997     

Plumage characteristics, reproductive investment and assortative mating in tree swallows Tachycineta bicolor

Elaborate ornamental plumage has been associated with various measures of individual quality in many species of birds. Male plumage characteristics, which have been relatively well studied, have been shown to reflect past reproductive investment, as well as the potential for reproductive investment in the current breeding attempt. In contrast, the signalling functions of female traits remain largely unexplored. In this study, we investigated the relationship between plumage attributes of breeding adult tree swallows and past reproductive investment, current reproductive investment and social mate pairing strategy. Both males and older females possess metallic green to metallic blue iridescent plumage on their dorsal surface, making this a suitable species for this type of investigation. We did not find any effects of past reproductive investment and success on the plumage attributes of returning breeders. In contrast, female plumage hue covaried with fledging success, and female plumage brightness was positively associated with mean clutch egg mass. In addition, we found that social pairs mated assortatively with respect to plumage brightness. We argue that since plumage characteristics vary with age in both male and female tree swallows, plumage attributes in this species are indicative of breeding experience and may be honest signals of quality. Positive assortative pairing could be the result of mutual mate choice or intra-sexual competition for nest sites by both males and females.     

Male mating effort in a polygynous ungulate

Survival and mating success are key fitness components of mammalian males. Because energy is limited, life history theory predicts trade-offs between current and future reproduction. While many studies have examined fitness trade-off in females, we know little about trade-offs faced by males. In polygynous species, male mating success is largely dependent upon intrasexual competition. Consequently, males have greater uncertainty over the benefits of a given allocation than over its costs, and the correlation between mating effort and success is likely much weaker in males than in females. We analyzed 14 years of data on the mating effort and survival of marked bighorn rams to investigate fitness trade-offs. Dominant rams defended single estrous ewes (“tending”) while subordinates attempted to copulate after separating the tending pair (“coursing”). We estimated the participation in tending and coursing for each ram and the effort in searching for breeding opportunities by each ram each year. We compared these three behavioral indices of male mating effort to demographic parameters, individual characteristics, and both yearly and long-term survival. Mating effort during the rut was unrelated to ram overwinter survival, but longevity was positively correlated with mating effort between 2 and 5 years of age. Persistent variation among rams is likely to explain this pattern, suggesting that in natural populations a few high quality males enjoy both high mating success and high survival.An erratum to this article can be found at     

Male reproductive success: paternity contribution to queens and workers in Formica ants

The relative number of workers and female sexuals fathered by two males mated with a queen were directly assessed using microsatellite and allozyme markers in field colonies of the ants Formica exsecta and F. truncorum. In both species one of the two males consistently fathered more offspring than the other. There was, however, no evidence that one male might be particularly successful in fathering a disproportionally high proportion of female sexuals relative to the proportion of workers. Moreover, in F. exsecta, the proportions of worker pupae and worker adults fathered by each male did not differ significantly between cohorts. The most likely explanation for this pattern is that females store different amounts of sperm from the two males they mated with. Received: 10 January 1997 / Accepted after revision: 22 March 1997     

Pair and extra-pair mating success relative to male quality in red-winged blackbirds

We tested whether the reproductive success of male red-winged blackbirds (Agelaius phoeniceus) varied with male secondary sexual traits or with haematozoa prevalence, and whether these patterns were consistent with females preferring genetically superior males. We also determined whether the traits that correlated with male success on their own territories were also correlated with male success at siring young on other males territories. Our analysis included data from a 6-year study involving 617 nestlings for which paternity was determined by DNA profiling. Larger males sired more young on their territories, principally because they obtained larger harems. The success of larger males at acquiring more mates did not appear to be a consequence of larger males holding larger or better-quality territories. Older and longer-lived males sired more young by extra-pair fertilizations. Larger males sired the most offspring overall (on territory + off territory). Variation in epaulet size and color, responses to male and female models, nest defence and parasitism was not correlated with male success either on or off their territories or overall. Male success in a given year was significantly correlated with success the previous year, as expected if females were selecting genetically superior males. The male that was by far the most successful individual in this study was highly consistent from year to year. Because male body size is positively correlated with survival in this population (although not within the sample of males included in this study), female preference for larger males may have reflected a general preference for males with superior Survival ability. We propose that the direct advantage realized by older males in extra-pair matings might indicate that experience is important, such that experienced males are better at creating or exploiting the opportunities for extra-pair mating. This hypothesis is consistent with a pairwise analysis of cuckoldry that showed that cuckolders were most often older than the males they cuckolded.     

Mate choice games, context-dependent good genes, and genetic cycles in the side-blotched lizard, Uta stansburiana

According to mate choice models, a female should prefer males with traits that are reliable indicators of genetic quality which the sire can pass on to their progeny. However, good genes may depend on the social environment, and female choice for good genes should be context dependent. The side-blotched lizard, Uta stansburiana, exhibits genetically based throat colors (orange, blue, or yellow) that could be used as a sexually selected signal since they reliably predict the genetic quality of mates. The frequencies of male and female morphs cycle between years, and both male and female morphs have an advantage when rare; thus genetic quality will depend on morph frequency. A female should choose a sire that maximizes the reproductive success of both male and female progeny. We examine a game theoretical model that predicts female mate choice as a function of morph frequency and population density. The model predicts the following flexible mate choice rule: both female morphs should prefer rare males in ’boom years’ of the female cycle (e.g., ’rarest-of-N rule’), but prefer orange males in ’crash years’ of the female cycle (’orange-male rule’). Cues from the current social environment should be used by females to choose a mate that maximizes the future reproductive success of progeny, given the social environment of the next generation. We predict that the cue is the density of aggressive orange females. In the side-blotched lizard, cycling mate choice games and context-dependent mate choice are predicted to maintain genetic variation in the presence of choice for good genes. Received: 8 March 2000 / Revised: 26 August 2000 / Accepted: 4 September 2000     

Experimental assessment of ecological and phenotypic factors affecting male mating success and polyandry in northern watersnakes, Nerodia sipedon

To resolve conflicting field observations regarding the action of sexual selection, we used breeding experiments and paternity analysis of the 927 resulting offspring to assess how male size, condition, tail length, genetic similarity to the female, and variation in operational sex ratio (OSR) affected male reproductive success and the incidence of polyandry in northern watersnakes (Nerodia sipedon). Only size affected male mating success. Large males were more successful, but only when male size varied substantially and competition among males was intense (i.e., male-biased OSR). The conditional nature of the size advantage may explain why studies of free-living watersnakes have produced inconsistent results regarding the relationship between male size and mating success. Size differences between males did not affect the proportion of offspring each male sired within multiply sired litters. We found positive size-assortative mating, but only when the OSR was female biased, suggesting that smaller males had improved access to females when competition among males was reduced, but that competition with larger males still restricted mating opportunities of small males to less preferred, smaller females. Most litters (58%) were multiply sired and larger females were more likely to produce multiply sired litters, similar to free-living watersnakes. There was no association between the incidence of multiple paternity and OSR, however, suggesting that polyandry is not simply a function of opportunity, with females passively waiting for males to court them.     

Dominance, access to colonies, and queues for mating opportunities by male boat-tailed grackles

Species characterized by female-defense polygyny have extreme variance in male mating success. Many studies have considered alternative male strategies for access to females, but few have considered age-specific strategies. Male boat-tailed grackles (Quiscalus major) compete for access to colonies of females and form linear dominance hierarchies. I observed two groups of males that competed for females at seven colonies. Dominance rank was significantly correlated with mass, but not after controlling for age. In contrast, the correlation between dominance rank and age remained significant after controlling for mass. Older males dominated younger males and dominance relationships were very stable. Thus, dominance hierarchies represent queues for mating opportunities. A male's rank in the hierarchy determined how closely he approached a colony. Furthermore, males of all ranks prevented lower-ranked individuals from approaching the colonies. Dominance rank thus determined access to nesting females. One top-ranking male's loss of mass over the course of the breeding season presumably reflected the energetic cost of defending females, but he maintained his position in the hierarchy despite the small loss of mass. One alpha male held a colony for at least 4 years, and the ages of males from two queues indicated that males wait 6 or more years before becoming an alpha male. Therefore, most males die before acquiring a colony of females. Spatial structure such as that documented here could obscure recognition of queues and explain why they have not been documented in more species. Received: 13 May 1996 / Accepted after revision: 26 April 1997