Sexual size dimorphism in fallow deer (Dama dama): do larger, heavier males gain greater mating success?

Sexual size dimorphism may evolve as a result of both natural and sexual selection. In polygynous mammals, the main factor resulting in the evolution of large body size in males is the advantage conferred during competition for mates. In this study, we examined whether sexual selection acts on body size in mature fallow bucks (Dama dama) by examining how the following traits are inter-related: age, body (skeletal) size, body mass, prerut dominance rank, rut dominance rank and mating success. This is the first study to examine how all these factors are together related to the mating success of a large sexually dimorphic and polygynous mammal. We found that male mating success was directly related to body size, but not to body mass. However body mass was related to prerut dominance rank which was in turn strongly related to rut dominance rank, and thus there was an indirect relationship between mating success and body mass. Rut dominance rank was the variable most strongly related to mating success. Mating success among mature males was unrelated to age. We conclude that larger mature fallow bucks have advantages over other males when competing for matings, and sexual selection therefore continues to act on sexual size dimorphism in this species. Heavier fallow bucks also have advantages, but these are mediated through the dominance ranks attained by males before the rut.     

Lifetime mating success, sexual selection and life history of fallow bucks (Dama dama)

We used data from a long-term study (15 years) of fallow deer to report for the first time the lifetime mating success, overall variance in lifetime mating success, and age-specific mortality levels of males. Fallow bucks that gain matings have higher social dominance rank, higher rates of fighting, and invest more in vocal display during the breeding season than unsuccessful males. Therefore, we examined if mating was associated with trade-offs in terms of survival, lifespan, and mating potential. We found that the variance in lifetime mating success was very high: 34 (10.7%) males mated, and of those, the 10 most successful males gained 73% of all matings (n=934). Mortality rates were generally high and only 22.3% (71/318) of males reached social maturity, i.e., 4 years. The oldest male was 13 years old. We found that fallow bucks that mated were not more likely to die during the following year, did not suffer from a reduction in lifespan, and did not incur lower mating success later in life as a result of mating during the early years of social maturity. Our results show that mating males at age 5 years (and possibly 9 years) may be more likely to survive than non-mating males. Additionally, the number of matings gained by males during the first years of social maturity was positively correlated with lifespan. We suggest that mating males are of higher quality than non-mating males because they are not more likely to incur trade-offs as a result of their increased reproductive efforts. Received: 9 November 1999 / Revised: 30 April 2000 / Accepted: 27 May 2000     

Fallow deer polyandry is related to fertilization insurance

Polyandry is widespread, but its adaptive significance is not fully understood. The hypotheses used to explain its persistence have rarely been tested in the wild and particularly for large, long-lived mammals. We investigated polyandry in fallow deer, using female mating and reproduction data gathered over 10 years. Females of this species produce a single offspring (monotocous) and can live to 23 years old. Overall, polyandry was evident in 12 % of females and the long-term, consistent proportion of polyandrous females observed, suggests that monandry and polyandry represent alternative mating strategies. Females were more likely to be polyandrous when their first mate had previously achieved high numbers of matings during the rut or was relatively old. However, polyandry was not related to the following factors: female age, the stage of the rut, the dominance ranks of mates, or the number of daily matings achieved by males. Polyandrous and monandrous multiple-mating females were not more likely than single-mating females to be observed with an offspring during the following year, and there were no significant differences in offspring size between these females. These results provide support for a fertility insurance hypothesis, with females remating if fertilization from the first mating was uncertain due to possible sperm depletion. The potential for different female mating strategies among large, polygynous mammals has generally been overlooked. Our findings highlight the complexity of female reproductive strategies and the possible trade-offs between fertilization success, preferences for high-quality males, and potential costs of polyandry, particularly for monotocous species.     

Fertilisation success in the fly Dryomyza anilis (Dryomyzidae): effects of male size and the mating situation

Male fertilisation success in relation to male size and the mating situation (ordinary pair formation with a single, nonvirgin female vs. take overs) was examined in the fly Dryomyza anilis. In ordinary matings, large males achieved higher fertilisation success than small ones when they were the second to mate with the female. Take overs differ from ordinary pair formation in that the second male experiences intensified sperm competition. This is because in take overs the female is not able to discharge any of the sperm inseminated by the first male as she usually does before a new mating. Compared with ordinary matings, take overs reduced the fertilisation success of the second male by 8–10%, whereas that of the first male was 7–14% higher in take overs. Even though the intruder was always larger than the paired male his superior fertilisation success did not compensate for the effect of the sperm already present in the female. In D. anilis, males can increase their fertilisation success by tapping the female's external genitalia with their claspers or having several copulation bouts per mating. Thus, in a take over, the intruder could respond to the intensified sperm competition by performing more tapping sequences per copulation bout or more copulation bouts per mating. In matings observed in the wild, males performed more tapping sequences after a take over than after pair formation with a single female, although the difference was not significant. The results show that there are differences in fertilisation success between males of different size. In addition, different mating situations can result in considerable variation in the fertilisation success of an individual male. Higher fertilisation success for the first male after a take over may be significant, in particular, for the reproductive success of small males, which frequently lose their females to large males.     

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.     

Extra-pair paternity in willow ptarmigan broods: measuring costs of polygyny to males

Willow ptarmigan are one of only three monogamous grouse species in North America. However, in some populations between 5 and 20% of individuals pair polygynously. It has been suggested that monogamy may be maintained by the high cost of polygyny to males. We have used DNA fingerprinting to assess the actual reproductive success of both monogamous and polygynous adults. We determined whether or not the putative parents were the biological parents of the chicks from 38 broods. Of these clutches 30 were from monogamous matings, and 8 were from bigamous matings. Of the 207 chicks from monogamous matings 96% were within-pair offspring, compared to 67% of the 49 chicks from bigamous matings. All extra-pair offspring chicks resulted from extra-pair fertilizations (EPFs), and there were no instances of intraspecific nest parasitism. Mate guarding by monogamous males seems to be a highly effective method for maintaining genetic monogamy, as the only cases in which EPFs occurred were when the resident female left the territory for a few days or when a second female visited the territory. Our results support the notion that certainty of parentage may be one factor constraining willow ptarmigan males to be monogamous.     

Lifetime mating success in a natural population of the damselfly,Enallagma hageni (Walsh) (Odonata: Coenagrionidae)

Summary Variance in lifetime mating success was measured for individuals of a population of Enallagma hageni, a non-territorial damselfly in northern Michigan. E. hageni is an explosive breeder with scramble competition for mates. Highly skewed operational sex ratios resulted in intense male-male competition which took the form of interference with tandem pairs. 41% of the males failed to mate in their lifetime as opposed to only 3.6% mating failure in females. The effect on mating success of size, age, longevity, and time spent at the breeding site were investigated. Intermediate sized males obtained the most matings, and male lifetime mating success was highly correlated with longevity.     

Parental investment, potential reproductive rates, and mating system in the strawberry dart-poison frog, Dendrobates pumilio

We studied the effect of relative parental investment on potential reproductive rates (PRRs) to explain sex differences in selectivity and competition in the dart-poison frog Dendrobates pumilio. We recorded the reproductive behavior of this species in a Costa Rican lowland rainforest for almost 6 months. Females spent more time on parental care than males, and `time out' estimates suggest that PRRs of males are much higher than than those of females, rendering females the limiting sex in the mating process. Males defended territories that provide suitable calling sites, space for courtship and oviposition, and prevent interference by competitors. Male mating success was highly variable, from 0 to 12 matings, and was significantly correlated with calling activity and average perch height, but was independent of body size and weight. Estimates of opportunity for sexual selection and variation in male mating success are given. The mating system is polygamous: males and females mated several times with different mates. Females were more selective than males and may sample males between matings. The discrepancy in PRRs between the sexes due to differences in parental investment and the prolonged breeding season is sufficient to explain the observed mating pattern i.e., selective females, high variance in male mating success, and the considerable opportunity for sexual selection. Received: 9 June 1998 / Received in revised form: 27 March 1999 / Accepted: 3 April 1999     

Alternative mating strategies in the water strider Gerris elongatus (Heteroptera,Gerridae)

Summary Males of the water strider Gerris elongatus established territories which included copulation and oviposition sites (small pieces of fallen bamboo). Males were aggressive and competition for territory and females was observed frequently. Male midlegs were more developed than female midlegs and were used as weapons. Reproductive behaviour changed as the breeding season advanced. Early in the season immature females were attracted by male surface wave courtship signals, then copulated white floating on the water surface without ovipositing (type 1). In midseason, males established territories, produced calling signals and attracted females which copulated and oviposited there with male postcopulatory guarding (type 2). In late season, many females oviposited without postcopulatory guarding on pondweed mats near fallen bamboo. Non-territorial males waiter for the arrival of these females and copulated without courtship, but mating success was low (type 3). These alternative mating strategies appeared to depend on differences in male size. Larger males were superior to smaller males in many ways (establishing territory, fighting, mating etc.). The largest males defended territories and had higher mating success than small non-territorial males. Medium sized males used all three strategies according to the number of empty territories and seasonal femald distribution.     

Territory establishment in lekking marine iguanas, Amblyrhynchus cristatus: support for the hotshot mechanism

The territory establishment of male marine iguanas and their subsequent mating success were analysed to identify spatial spillover (hotshot) and temporal spillover effects on lek formation. Males started to establish small display territories 2 months ahead of the mating season. Males did not establish territories in temporal synchrony and did not settle at sites where the probability of encountering females was highest. However, males arriving later preferentially established their territories in the neighbourhood of already established territories independently of the density of female-sized iguanas in these territories. Although settling in close proximity, there were no fights between those males. The number of fights between territorial males increased towards, and peaked during, the mating season. Fights did not result in the transfer of space, indicating that space per se was no resource. Instead, fights were directed towards central (hotshot) males. These central males had higher mating success than marginal males. Female density during the time of territory establishment did not predict the mating success of males, because females changed their spatial preferences between early establishment and mating periods. Similarly, the areas where males achieved the highest numbers of copulations changed during 4 years of our study. Thus, there was no evidence for temporal spillover between subsequent seasons. However, most male-male interactions served to distract successful males and may lead to spatial spillover of females into territories of unsuccessful males. In marine iguanas, territorial establishment appears largely governed by hotshot processes.     

Mating strategies of a nocturnal,desert rodent (Dipodomys spectabilis)

Summary The mating system of a nocturnal, desert rodent, the banner-tailed kangaroo rat (Dipodomys spectabilis) was studied through direct observation, live-trapping, and radiotelemetry over a 13-month period from August 1986 to August 1987. Mating behavior varied from exclusive matings between male and female neighbors to competitive mate searching and direct male competition. In summer matings and early in a November to May breeding season, males located receptive females and mated exclusively with them without disturbance from other males. As the operational sex ratio changed in favor of males, multiple males converged on an estrous female's territory and competed for access to her. However, an older, experienced male usually monopolized the matings of the same one to two close female neighbors for the entire breeding period, and females mated with the same male neighbor over several estrous cycles. Monopolization of females by neighbor males was facilitated by female relaxation of individual territorial defense. Dominant males spent considerable time in the territories of the females they monopolized before and during mating. This relaxation in territorial defense was seen in dyadic encounters in which females tolerated all males but allowed significantly more contact by neighbor than stranger males. Neighbor recognition, therefore, seems important in coordinating the mating interactions of this solitary rodent.     

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.     

Determinants of paternity success in the spider Pholcus phalangioides (Pholcidae: Araneae): the role of male and female mating behaviour

In double mating experiments, we examined whether and to what extent various male and female behavioural traits influence the course of mating and fertilization success in the cellar spider. In males, we focussed on pre-copulatory behaviour and on the rhythmic twisting movements that the male performs with his pedipalps during copulation. In females, we investigated remating decisions and the effect of female termination of copulation. Second males fertilized a high proportion of the eggs (P ₂: median 89%) despite much shorter second matings, with high variation in relative paternity success. The number of pedipalp movements (PPMs) of either male was a better predictor of paternity than copulation duration. Our results suggest that in second matings, PPMs help to remove sperm from previous males, whereas in first matings a high number of PPMs enhances fertilization success, either due to numerical sperm competition or cryptic female choice. Furthermore, we found a negative male age effect on paternity in second matings, implying that age-related deterioration of spermatozoa may promote variation in fertilization success. Female receptivity decreased significantly in second matings; only 70% of the females remated. Females that accepted a second copulation were found to terminate these much earlier and with higher probability than first matings. This suggests that the intensity of conflict between the sexes is higher in second matings. Increased intensity of sexual conflict may be responsible for stronger selection on male traits, as pre-copulatory behaviour and age only affected male copulatory performance and paternity in second matings. Electronic Publication     

Male–female socio-spatial relationships and reproduction in wild chimpanzees

Recent research on primates and other taxa has shown that the relationships individuals form with members of the same sex affect their reproductive success. Evidence showing that intersexual relationships also influence reproduction, however, is more equivocal. Here, we show that male chimpanzees living in an exceptionally large community display long-term tendencies to associate with particular females. These association patterns are likely to arise because individuals of both sexes selectively range in specific areas of the communal territory, with males inheriting the ranging patterns of their mothers. These differentiated male–female socio-spatial relationships involved males of widely varying ranks, and their effect on reproduction is as strong as that of male dominance rank, which in turn is as strong a predictor of reproductive success at Ngogo as in other smaller chimpanzee communities. These results show that male–female socio-spatial relationships can play a large role in chimpanzee male reproductive strategies, although they probably neither weaken nor strengthen the relationship between male dominance rank and reproductive success. Our results linking male–female socio-spatial relationships to reproduction in chimpanzees suggest that the gap between the social and mating systems of humans and their closest living relatives may not be as large as previously thought.     

Correlates of copulatory success in a fallow deer lek

Summary We studied the behavior and copulatory success of fallow deer (Cervus dama) males at a lek. We recorded 471 copulations in 3 years. Most males did not copulate. The top three males accounted for between 60 and 90% of copulations each year. Lek attendance time was the major determinant of male copulatory success, but territory location also affected the number of copulations achieved. Copulatory success was correlated with the number of females in a male's territory and possibly with dominance status, but not with fighting success or fighting rate. All males that defended lek territories were 5 years of age or older. Copulatory success may improve with age. Body condition appears to be an important determinant of male copulatory success, because only males in superior condition could defend a lek territory for up to 2 weeks. Males do not feed while defending lek territories. Foraging ability during the year probably determines condition at the onset of the rut. Females appear to choose mates at least partially on the basis of location, preferring males located near traditional routes. Females may ultimately select mates in the best body condition.     

Mating success on white-bearded manakin (Manacus manacus) leks: male characteristics and relatedness

Morphological, territorial, and behavioural characteristics were measured on white-bearded manakin (Manacus manacus) males from two leks in two consecutive years. These data were combined with data on marker-inferred relatedness to study possible co-variation with mating success. In one year, male size and male condition were correlated with mating success. In both years, males holding courts nearer the lek centre gained more matings. No observed male display behaviour appeared to be an independently important factor in explaining variance in male mating success. Successful males made more aggressive displays than non-successful males and more displays were between close relatives. Number of aggressive displays increased as the distance between male courts decreased. Mating success in the white-bearded manakin is most likely mediated by a combination of morphological and behavioural characteristics, influencing both male-male competition and female choice. Females could potentially use centrality on the lek as an indicator of male characteristics. However, levels of relatedness may influence spatial arrangement of males on a lek thereby affecting male-male interactions and ultimately influencing patterns of mating success.     

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     

Positive association between social and extra-pair mating in a polygynous songbird, the dickcissel (Spiza americana)

In polygynous species, males appear to gain additional offspring by pairing with multiple females simultaneously. However, this may not be true if some females copulate outside of the social pair bond. Polygynous males could experience lower paternity because of trade-offs among gaining multiple social mates, guarding fertility with these mates, and pursuing extra-pair matings. Alternatively, polygynous males could simultaneously gain extra social mates and have high paternity, either because of female preferences or because of male competitive attributes. We tested four predictions stemming from these hypotheses in a facultatively polygynous songbird, the dickcissel (Spiza americana). Unlike most previous studies, we found that males with higher social mating success (harem size) also tended to have higher within-pair paternity and that the number of extra-pair young a male sired increased significantly with his social mating success. Females that paired with mated males were not more likely to produce extra-pair young. In contrast, extra-pair paternity was significantly lower in the nests of females whose nesting activity overlapped that of another female on the same territory. This pattern of mating was robust to differences in breeding density. Indeed, breeding density had no effect on either extra-pair mating or on the association between polygyny and paternity. Finally, nest survival increased with harem size. This result, combined with the positive association between polygyny and paternity, contributed to significantly higher realized reproductive success by polygynous male dickcissels.     

Polyandry, multiple mating, and female fitness in a water strider Aquarius paludum

Contrary to Bateman’s principle, polyandry appears to be a common female mating strategy. Several hypotheses have been proposed to explain the evolution of polyandry. It is assumed that females gain either material or genetic benefits from multiple matings, or that they are coerced into mating by males. In water striders, mating is generally assumed to be costly to females, and they are thought to mate for reasons of convenience, adjusting their resistance to mating according to male harassment. Here, we tested the effect of number of matings (with the same male) and number of partners on female fitness in a water strider Aquarius paludum. In the first experiment, we regulated the time females spent with a male and found that females’ egg production increased with multiple matings up to a point. The result supports the existence of an optimal female mating frequency. In the second experiment, we tested how polyandry affects the number of eggs laid and egg hatching success. We conducted three different trials: females mated four times with either a single male, two different males, or with four different males. Females that mated with four different males laid the lowest number of eggs and had the lowest egg hatching success, suggesting that polyandry reduces females’ egg production and egg hatching success in A. paludum. We conclude that A. paludum females probably gain material benefits from mating but no genetic benefits were found in this study.     

Alpha-male paternity in elephant seals

The aim of this study was to assess paternity of males that dominated mating in harems at northern (Mirounga angustirostris) and southern (M. leonina) elephant seal rookeries using DNA fingerprinting and microsatellite DNA analysis. Southern alpha males had greater reproductive success than most northern alphas at similar-sized harems. Comparison of the relatedness between pups within harems also suggested that fewer males achieved matings in the southern elephant seal population. This was consistent with behavioral observations that suggest greater competition for mates in northern elephant seal harems. Reproductive success was consistent with estimates of mating success in some cases, but lower than expected for some northern elephant seal alpha males. A lower reproductive success than predicted from mating behavior may arise from a variety of factors including sperm competition, male sperm depletion from frequent mating, or reduced fertility. The alternatives are discussed in the context of environmental and historical factors. Received: 17 August 1998 / Received in revised form: 28 March 1999 / Accepted: 30 May 1999