Passing the buck: resource defence,lek breeding and mate choice in fallow deer

Summary Lek breeding systems, where males defend small, clustered mating territories, are thought to occur where the distribution of females is heavily clumped but males are unable to defend resources used by females. In this paper, we describe a breeding system in fallow deer where males are able to defend resources used by females but the most successful bucks instead defend small territories on a traditional mating ground; where the lek is sited in an area not heavily used by females at other times of year and is visited primarily by females in or close to oestrus; and where mating success on the lek is related to territory position and to male phenotype but not to the resources available on different lek territories. Comparisons with other ungulates suggest that lek breeding species fall into two groups: those where leks are regularly visited by herds of females many of which are not in oestrus and those, like fallow deer, where leks are visited primarily by oestrous females. In the latter species, it is unlikely that females visit the lek for ecological reasons.     

Male-biased maternal expenditure and associated costs in fallow deer

This study tested whether fallow deer mothers, Dama dama, bias their investment towards sons and, thus, whether sons are more costly to produce than daughters. Young (2 years) and old (≥3 years) hinds were analysed separately. Old hinds who raised sons accumulated less body mass than those who raised daughters, during the period between late gestation and the end of lactation. This difference in body mass persisted to the following spring. Mothers who had raised sons gave birth later and their offspring's pre-winter mass was lower the following year than for mothers who had raised daughters. These results indicate higher expenditure for hinds who raise sons and support theories of male-biased maternal investment. However, young mothers with sons and those with daughters did not differ in reproductive performance the following year. One reason might be that young mothers are close to the maximum level of maternal expenditure, since they are still growing, and cannot invest any extra resources in sons. Received: 28 August 1997 / Accepted after revision: 5 April 1998     

Effectiveness of multiple release sites in reintroduction of Persian fallow deer

Releasing animals in more than one location may increase or decrease the probability of success of a reintroduction project, yet the question of how many release sites to use has received little attention. We used empirical data from the reintroduction program of the Persian fallow deer (Dama mesopotamica) (Galilee region in northern Israel) in an individual-based spatially explicit simulation model to assess the effects of releasing deer from multiple sites. We examined whether multiple release sites increase reintroduction success, and if so, whether the optimal number of sites for a given scenario can be determined and whether the outcome differs if animals are released alternately (i.e., the location of the release alternates yearly between sites) or consecutively (i.e., one release site is used for several years, then another is used, and so forth). We selected 8 potential release sites in addition to the original site and simulated the release of 180 individuals at a rate of 10 individuals per year in different combinations of the original site and 1-4 additional sites. In our model, releasing animals into the wild at multiple sites produced higher population growth and greater spatial expansion than releasing animals at only one site and a consecutive-release approach was superior to an alternate-release approach. We suggest that through the use of simulation modeling that is based on empirical data from previous releases, managers can make better-informed decisions regarding the use of multiple release sites and greatly improve the probability of reintroduction success.     

Components of variance in male lifetime copulatory and reproductive success in a seed bug

Summary Males and females of the seed bug, Neacoryphus bicrucis Say, were individually numbered in the field in southeastern Georgia (USA) and census taken daily for 6 weeks. Variation in male mating efficiency (ME = no. copulations/no. sightings) exceeded that in females and was significantly greater than that generated by a null model. Lifetime copulatory success, estimated as the product of ME and longevity, ranged from 0 to 41, with ME accounting for over 88% of the variation. Lifetime reproductive success (LRS), estimated as the product of ME, longevity, and clutch size, ranged from 0 to 898. Among males copulating at least once, ME accounted for over 40% of the variation in LRS, while longevity alone or in combination with clutch size accounted for 21% and 46%, respectively, of the variation in LRS. Failure of some males (26%) to copulate contributed 38% of the total variation in LRS. Thus, among males surviving to adulthood, sexual selection pressure arising from variation in ME is approximately as strong a force for phenotypic change as is natural selection pressure arising from variation in longevity.     

Correlates of extra-pair fertilization success in hooded warblers

We examined correlates of extra-pair fertilization (EPF) success in the hooded warbler (Wilsonia citrina), a species where females pursue extra-pair matings. The good genes hypothesis predicts that females choose extra-pair mates with morphological or behavioral traits that reflect differences in male genetic quality. EPFs were common, as 35.3% (42/119) of broods contained extra-pair young (EPY) and 26.7% (95/356) of nestlings were the result of EPFs. There was a strong skew in male EPF success which increased the variance in annual male mating success 2–3 fold. However, male morphology did not predict EPF success, as extra-pair males were not older or larger than the males they cuckolded. Likewise, there were no significant correlations between the proportion of extra-pair young in a brood and male size or age. The good genes hypothesis predicts that high-quality males will be consistently preferred as genetic mates, but the number of young sired by a male with his social mate was not consistent from one year to the next. There was a significant negative correlation between female age and proportion of EPY produced, which could result if older females obtain higher-quality social mates. We found no strong evidence that females choose extra-pair mates for good genes, but females may use behavioral rather than morphological cues to assess relative male quality. Received: 31 May 1996 / Accepted after revision: 17 October 1996     

Ecological sexual segregation in fallow deer (Dama dama): a multispatial and multitemporal approach

We questioned the different interpretations of ecological sexual segregation from a novel perspective, i.e., by carrying out diverse temporal and spatial scale analyses within a long-term study (1984–2003). Thus we combined spatial (small/large) and temporal (small/large) scale analyses to identify the factors generating sexual segregation in fallow deer in San Rossore, Italy. The study site was divided into an eastern sector characterized by human disturbance (DS) and a western undisturbed sector (US). According to census data, human presence increased in DS from 1984, and while females gradually abandoned it, males remained—thus supporting the predation risk hypothesis (large spatial and temporal scale)—and actually increased their presence in DS, where they seemingly benefited from a lower female density. This supported the indirect competition hypothesis. The analysis of data on a large temporal and small spatial scale confirmed that intersexual competition, in particular for grass, was higher in a crowded pasture in US. Observations by means of radio-telemetry of 23 adult females and 25 adult males (1997–2001, reduced temporal and large spatial scale) showed that large scale segregation was relevant during the day and disappeared at night, when disturbance was absent and also the females reached DS. This also supported the predation risk hypothesis. Moreover, sexes showed different habitat choices inside DS at night, thus supporting the forage selection hypothesis (small spatial and temporal scale). In conclusion, failure to address the whole set of combinations of spatial and temporal scale analyses would have led to monocausal explanations of ecological sexual segregation.     

Mating strategies and mating success of fallow (Dama dama) bucks in a non-lekking population

The rutting behaviour of bucks in an enclosed population was investigated between 1988 and 1990. A substantial proportion of the matings were observed. After preliminary observations in the 1987 rut we categorised bucks into one of four rutting strategies based mainly on their degree of territoriality. We investigate the effects of age, dominance and mating strategy on mating success. Territories were aggregated in an area of oak woods and mating success was highly skewed. Bucks of between 5 and 7 years old achieved the majority (over 90%) of observed matings. Mating success was highly correlated with dominance but only weakly related to fighting success. The possession of a territory was crucial to achieving high reproductive success, with a 38-fold difference between the most and least successful strategies. Bucks pursuing the different strategies also differed in the time they commenced groaning, timing of matings, mating interference and the locations where they achieved their matings. Although high-ranking males devoted considerable effort to obtaining and defending a territory only 36% of each buck's matings were achieved on his territory and males tended to abandon these sites when the tendency of females to visit them decreased.     

Social cohesion and foraging decrease with group size in fallow deer (Dama dama)

We studied the impact of group size on foraging behaviour and level of movement synchronisation among female herdmates of a fallow deer population in Central Italy. Both proportion of foraging events and movement synchronisation decreased with increasing group size. The proportion of foraging events was higher for animals on the edge of the group than for deer in the centre of the group; hence, there appears to be a trade-off between protection against predators and foraging interference, both of which decrease from the centre to the periphery of the group. This is the first time this type of behaviour has been recorded for wild ungulates. As expected, we also found that the movement of peripheral animals was less synchronised than that of central animals. Consequently, peripheral animals may lose contact with their herdmates and split off the group. We conclude that social inequalities may lead to conflicting requirements among group members and instability of large groups. Movement synchronisation (as a function of group size) appears to interact with habitat openness to produce variations of group size (which appear to be adaptive for individuals) as an emergent property of these aggregations.     

Correlates of male mating success in the ruff Philomachus pugnax,a lekking shorebird

Summary Male ruffs (Philomachus pugnax), a lekking shorebird, can generally be divided into two morphs according to behavior and plumage coloration. Predominantly dark-colored, independent male ruffs defend small territories on a lek, whereas lighter colored satellite males are non-territorial and less site-faithful to a particular lek. The traits associated with the mating success of independent male ruffs were studied at two leks during two seasons on the island of Öland, Sweden in May and June of 1985 and 1987. Using multivariate analyses, three characteristics were found to be significantly related to mating success: high frequency of visits by satellites to an independent male's residence, consistency of lek attendance, and low rates of aggressive behavior. In contrast, mating success was unrelated to the degree of darkness of the independent male nuptial plumage, territory location on the lek, or rate of courtship displays. The use of multivariate analyses helped to resolve conflicting conclusions from previous studies which employed simple statistical analyses, or none at all. Experimental manipulations are suggested which will help to further determine the influence of female mate choice in this lekking system.     

Behavioural interference between ungulate species: roe are not on velvet with fallow deer

Interference is expected to occur at feeding areas between species with a similar diet, but few studies have tested this idea for wild ungulates. We analysed interactions between fallow deer, European roe deer and wild boar, in three sites, in a Mediterranean area. We expected that interference should be greater between deer than between them and wild boar. We documented the negative effects of behavioural interference by fallow on foraging behaviour of roe deer, under field conditions. Deer species built up 90% interference interactions, with fallow always dominant on roe, also through direct aggression. Although roe deer decreased feeding and increased vigilance levels in proximity (<50 m) of either fallow deer or wild boar, they were displaced significantly more often by the former than by the latter. Fallow deer were neither displaced nor alarmed by roe and rarely by wild boar. No deer species displaced wild boar. Interference was significantly greater on solitary roe deer, especially females, in spring and roe left the feeding ground most often in the smallest site (13 ha). Roe deer avoided areas where the local density of fallow deer was the highest. During our 4-year-study, roe deer density decreased whereas fallow deer numbers increased. Behavioural interference may explain how fallow deer outcompete roe deer through spatial exclusion from feeding sites and avoidance of areas with high densities of the former. Fallow deer evolved in semi-arid, relatively poor habitats of Asia Minor: interspecific defence of crucial resources could have developed as a beneficial tactic for its survival.     

Characteristics that influence male reproductive success on a lek of Lethrinops c.f. parvidens (Teleostei: Cichlidae)

Sexual selection has long been proposed as a mechanism leading to the diverse cichlid (Teleostei: Cichlidae) fauna of Lake Malawi, Africa. Many of the shallow-water, sand-dwelling, bower-building cichlid species are particularly well suited for studies of sexual selection because they participate in leks. Since females in lekking systems appear to acquire only genetic material from their mates, it has been suggested that leks are ideal systems to study female mate choice. The objectives of the investigation were to examine Lethrinops c.f. parvidens male bower characteristics (i.e., bower size and location) as well as other male characteristics (i.e., length, gular color, and duration on the lek) for their influence on male mating success as measured by the number of visits, circles, and eggs laid by females. These measures are nested in that a visit by a female may or may not lead to circling, and circling by a female may or may not lead to egg-laying. We found increased bower height and higher numbers of conspecific neighbors (analogous to shallow-water, near-shore bower positions) to be positively, significantly associated with the number of visits by females. The only significant correlate with the number of circles was visits, and similarly circles was the only significant correlate with the number of eggs laid. The R ² value for the egg-laying regression was quite low (19.8%) compared with visits (54.3%) and circling (78.9%), suggesting that females may be using additional cues, that we failed to measure, when in close proximity to males or simply that a small proportion of the females were ready to spawn. Both indirect selection and direct selection pressure due to egg predation may have influenced female choice on the lek. Received: 10 April 1999 / Received in revised form: 26 July 1999 / Accepted: 18 September 1999     

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     

Extra-pair copulations in a lek: the secondary mating system of monogamous razorbills

Summary I studied the mating strategies of razorbills Alca torda, a monogamous colonial seabird, during 1987–1989 in Wales, U.K. The outstanding feature of the study population's mating system was the performance of most copulations in mating arenas outside the colony. Females visited the arenas and sometimes accepted extra-pair copulations (EPCs), and males aggressively competed for EPCs. During the fertilizable period of most females, the arenas resembled leks because they contained no resources except mates, and males contributed only sperm to extra-pair females. Typical of leks, the operational sex ratio was strongly male-biased, and male mating success was highly skewed. To examine the potential for sexual selection to operate in this system, I correlated male EPC success with variables that could be associated with male-male competition and/or female choice. The frequencies of three behaviors (interference of copulation attempts, fights won against other males, and EPC attempts), accounted for 62% of the variance in EPC success and were strongly intercorrelated, with interference explaining most of the variance in a multivariate test. EPC success was not correlated with body size, age, paired status or either of two estimates of the time spent in the arena. The hypothesis that male EPC success was determined by active female choice was tested by examining the relative success (EPCs per attempt) of males. Relative success was not correlated with any of the three behaviors associated with absolute success, or with age, body size or attendance, suggesting that male-male competition, mainly in the form of copulation interference, is the principal correlate of EPC success. These findings in a monogamous species illustrate that EPC is a secondary mating system of razorbills in which sexual selection operates. The existence of this system outside the colony raises the possibility that lekking for EPCs may also occur within the nesting territories of other monogamous, colonial species but is hidden by competition for nests and breeding partners.     

Correlates of male mating success in two species of Anthidium bees (Hymenoptera : Megachilidae)

Summary Field observations were made on the mating behavior of two congeneric species of solitary bees, Anthidium porterae in an arid grassland and A. palliventre in a coastal sand dune habitat. Males of both species exhibited resource defence polygyny and defended hostplants to gain access to females foraging for nectar and pollen. The mating frequencies of marked and measured resident (territorial) males were monitored during periods of continuous observation, following which measurements of territory size and floral resources were obtained. Mating success of A. palliventre males was strongly influenced by territory characteristics: Males that defended small areas with a few rich hostplant patches mated more often than males that held larger territories containing many hostplant patches of low floral density. Large males generally held high-quality sites and thus had a mating advantage over smaller individuals. In A. porterae, on the other hand, male mating success was unrelated to any measure of territory quality. Copulation frequency and male size were positively correlated, however, apparently due to the increased ability of large males to seize and hold females for mating. The two species also differed in the incidence of non-territorial, sneaky males. While absent in A. palliventre, sneaky males accounted for 12% of all mating observed in A. porterae. Males of A. porterae that displayed sneaky tactics mated, on average, as often as resident males. Offprint requests to: E.M. Villalobos     

Male copulatory guarding enhances female foraging in a water strider

Summary In the laboratory, females of Gerris remigis foraging singly after being separated from a copulating male averaged 16 times longer to repel male copulatory attempts compared to repelling time when copulating (60 trials). In<10 min after separation, all females either became immobile at the edge (in 34% of all trials), or recopulated and either continued to forage (61%) or swam to the edge and became immobile (5%). Copulating females captured 85% of imitation prey, but single females, harassed by males, captured only 32% of prey.During a census of 2 small streams, in the pools with swimming single males, >99% of all swimming females were copulating, <1% being single. Ninety seven percent of all single females were immobile at the edge, whereas only 11% of all pairs were at the edge. In pools without swimming single males, 32% of all single females were swimming, compared to <1% in pools with swimming males. Introductions of a male into 4 pools with a female swimming singly in each resulted in the females either becoming immobile at the edge or copulating. The 2 streams had a : sex ratio of 1.6:1, and 79% of all females were copulating during the census. Foraging, copulation and copulatory attempts continued at a reduced level during the night.Thus in streams where male G. remigis are attempting to copulate, females can forage effectively only by carrying a copulating male who apparently repels copulatory attempts by other males.     

Correlates of male reproductive success in the browncheek blenny,Acanthemblemaria crockeri (Blennioidea: Chaenopsidae)

Summary Acanthemblemaria crockeri exhibits a resource defense polygyny mating system with male parental care. Females preferred to mate with the larger of two males in laboratory experiments, and male size was positively correlated with the number of eggs defended by males at two sites in the Gulf of California, Baja California, Mexico. Females appeared to avoid mating with males defending heavily-fouled shelters. The role of other factors including the intensity of male courtship coloration and displays in determining male reproductive success was studied in Bahia San Carlos, Sonora, Mexico by providing glass vials as shelters. This provided a non-destructive technique for counting of eggs defended by males and allowed repeated assessment of the reproductive success of individual males. Males varied greatly in their duration of residency of shelters and in their color score and intensity of courtship displays. The number of eggs received by males was positively correlated with their duration of residency in shelters, and females appeared to avoid mating with new residents. Mated males deserted shelters less frequently and were more likely to receive future matings than were unmated males. The mean color score of males was unrelated to their reproductive success, while the intensity of male courtship displays was negatively correlated with the number of eggs received. This may have resulted because female mate choice is based on multiple criteria, including some that more accurately reflect the quality of parental care afforded by males.     

Neighbours, strangers and male-male aggression as a determinant of lek size

Interactions between males on leks may play an influential role in lek formation and the regulation of lek size. In this paper I present the results of a playback experiment that simulated de novo settlement at sites adjacent to currently existing display territories of the ochre-bellied flycatcher, Mionectes oleagineus. In the study population, males displayed both solitarily and at small leks. A large proportion of males held no display territory at all. A stranger's song was played to both solitary and lekking males from 10 m outside their territorial boundaries. In separate playbacks, lekking males were also played neighbour's song. Both lekking and solitary territorial males reacted to the playback by decreasing their song rate, approaching the playback speaker and, on occasion, attacking the model. Solitarily displaying males responded more aggressively to playback of stranger's song than did lek males. Lek males were able to distinguish between their neighbour's and a stranger's song and did so irrespective of whether it was played from the neighbour's territory or from outside the lek. In addition to distinguishing between neighbours and strangers, lek males modified their responses to these different playbacks depending on where the playback originated. These results suggest that male-male interactions can be influential in structuring leks. In M. oleagineus, interactions between males are aggressive and act to limit rather than augment lek size. Received: 6 March 1996 / Accepted after revision: 9 December 1996     

Correlates of male mating success in great bustard leks: the effects of age, weight, and display effort

We examined how mating success varied in relation to age, weight, body size, and display behavior among great bustard Otis tarda males. The estimated mating success was strongly skewed, with 45% of adult males being involved in copulation attempts and only 9.7% actually seen copulating successfully. Unlike most birds, body size continued increasing in great bustards several years after reaching sexual maturity. Age, weight, and display effort were all significant and independent predictors of male mating success. The higher display effort involved performing longer full-display bouts. Older males could detach from the male flock earlier in the season as well as on each day and spend longer seasonal and daily periods displaying as solitary birds, which contributed to increase their mating success. In contrast, males weighing more did not invest more in display, which suggests that they could be recognized as dominants by other males and selected by females through assessment of their plumage sexual traits. In contrast to most other bird species, the system described for great bustards resembles that found in some lek-mating ungulates, where social rank is a complex trait determined by both age and mass, and as in these mammals, it suggests that sexual selection continues to favor a high male weight in this extremely sexually dimorphic species.     

Field playback of male display attracts females in lek breeding sage grouse

Summary Recent correlational studies of lekking sage grouse suggest that male vocal display attracts females. To test this hypothesis further, the natural displays of a territorial male were supplemented with the tape-recorded display of another, reproductively-successful, individual. Significantly, more females approached the speaker's location on days when the recording was played, and also on non-playback days immediately following a playback, than on other non-playback days (Fig. 1). Analysis of male displays indicated that females were responding to the playback itself rather than to changes in male behavior. The after-response following a playback suggests that some females present during a playback remembered its location and approached on a subsequent lek visit. The results provide necessary support for the epigamic function of vocal display, and suggest ways in which female responses to male display may influence lek structure.