Mating system evolution in response to search costs in the speckled wood butterfly, Pararge aegeria

A general and intuitive prediction from models of mate preference is that when the cost of searching for mates increases, individuals should become less choosy. Here, we test this prediction by comparing the mating propensity of females in two populations of the butterfly Pararge aegeria. The populations originated from southern Sweden and Madeira and due to different adult emergence patterns throughout the year, the average density of males per female is likely to be lower on Madeira. Therefore, we expected that the cost of searching should be greater on Madeira and, consequently, that the Madeiran females should be less choosy. In line with predictions, the Madeiran females mated significantly sooner after the first interaction with males than did females from southern Sweden. This difference may reflect a weaker preference for territorial males over non-territorial patrollers in the Madeiran population, because of the greater costs of searching. The Madeiran females also showed a shorter time lag between mating and the start of oviposition. We discuss this unexpected result and propose that the same mechanism could also explain this population difference, i.e. different costs of searching for suitable host plants. Both search processes are fundamental for female reproductive success and we find it plausible that they can be generalised into the same theory of optimal search behaviour. Received: 14 May 1998 / Accepted after revision: 13 December 1998     

Determinants of male mating success in the red bishop (Euplectes orix)

We studied sexual selection in the red bishop, Euplectes orix, a colonial, polygynous weaverbird widely distributed over sub-Saharan Africa. Male reproductive success measured in terms of the number of nests accepted by females and the number of eggs and nestlings in all the nests on a male's territory varied considerably. The standardized variance (variance/mean²) in male reproductive success ranged from 0.505 to 1.737 in different years, indicating a high potential for sexual selection in this species. An analysis of genetic parentage for 432 nestlings by non-radioactive, multilocus DNA fingerprinting confirmed that male reproductive success (number of young sired on the territory) in this species can be reliably estimated by the measures introduced above. In all 4 study years there was a strong positive correlation between male mating success and the total number of nests that males built in their territories. The number of nests built can be partitioned into the number of weeks a male held a territory and his nest-building performance. Both factors exert a significant positive effect on male mating success and in combination explained between 53.3 and 86.3% of the variation in male reproductive success. Male morphological characters were found to be of no importance. Males that established a territory in the following season built more nests and held their territories for longer than males that did not establish a territory in the following season, suggesting that these measures might be indicators of male condition and quality. Male nest-building performance (number of nests built per week) seems to be unrelated to male condition or quality. Received: 8 January 1999 / Received in revised form: 7 June 1999 / Accepted: 13 June 1999     

Female choice in the opportunistic mating of wild chimpanzees (Pan troglodytes schweinfurthii) at Mahale

For female chimpanzees (Pan troglodytes schweinfurthii) in the Mahale Mountains National Park, Tanzania, the most common mating pattern is opportunistic. In such opportunistic matings, females copulated promiscuously but not randomly. This study describes female mate choice during 1-year observation of six females who exhibited regular genital-swelling cycles. During the study period, 169 opportunistic matings and four restrictive matings were recorded over the course of 51 days. As female estrus progressed, mating frequency and the number of adult male mating partners increased, although the number of potential mating partners did not change. Criteria of female choice examined were the direction and consent/rejection of courtship, proximity maintenance, and female grooming. Adult-male courtships were successful more often than those of adolescent males. During the earlier phase of estrus, females copulated rather promiscuously with many males. But during the later phase of estrus when the likelihood of conception is expected to be highest, they copulated repeatedly with high-ranking adult males. There was a positive correlation between female grooming frequency and mating frequency when the likelihood of conception was greatest. Female chimpanzees are thought to choose high-ranking males as fathers of their offspring. Moreover, female chimpanzees may adopt one or both of two mating strategies, i.e., a many-male strategy and a best-male strategy. Received: 23 November 1998 / Received in revised form: 12 April 1999 / Accepted: 26 April 1999     

Imperfect female choice and male mating skew on leks of different sizes

We present a model of error-prone female choice on leks, and investigate the effects of different degrees of error on the distribution of mating success among males present at leks of different sizes. At higher levels of error, the best male is predicted to gain a smaller share of matings, while low-ranking males gain a larger share. Males who are of high rank but not the most desirable on the lek do best at intermediate levels of error, since the top-ranked male does not then claim all the matings, but assessment is still sufficiently accurate for females to discriminate between high-ranking and low-ranking competitors. The effects of error are shown to be more pronounced on larger leks, due to smaller expected differences in mating value between males of adjacent ranks. This interaction between lek size and error suggests that observed negative relationships between lek size and mating skew need not be attributed solely to intrasexual competition, as previously suggested, but could also be a result of imperfect choice. Received: 20 February 1998 / Accepted after revision: 25 October 1998     

A framework for allocating conservation resources among multiple threats and actions

Land managers decide how to allocate resources among multiple threats that can be addressed through multiple possible actions. Additionally, these actions vary in feasibility, effectiveness, and cost. We sought to provide a way to optimize resource allocation to address multiple threats when multiple management options are available, including mutually exclusive options. Formulating the decision as a combinatorial optimization problem, our framework takes as inputs the expected impact and cost of each threat for each action (including do nothing) and for each overall budget identifies the optimal action to take for each threat. We compared the optimal solution to an easy to calculate greedy algorithm approximation and a variety of plausible ranking schemes. We applied the framework to management of multiple introduced plant species in Australian alpine areas. We developed a model of invasion to predict the expected impact in 50 years for each species-action combination that accounted for each species’ current invasion state (absent, localized, widespread); arrival probability; spread rate; impact, if present, of each species; and management effectiveness of each species-action combination. We found that the recommended action for a threat changed with budget; there was no single optimal management action for each species; and considering more than one candidate action can substantially increase the management plan's overall efficiency. The approximate solution (solution ranked by marginal cost-effectiveness) performed well when the budget matched the cost of the prioritized actions, indicating that this approach would be effective if the budget was set as part of the prioritization process. The ranking schemes varied in performance, and achieving a close to optimal solution was not guaranteed. Global sensitivity analysis revealed a threat's expected impact and, to a lesser extent, management effectiveness were the most influential parameters, emphasizing the need to focus research and monitoring efforts on their quantification.     

Mate sampling behaviour of black grouse females (Tetrao tetrix)

We studied female mate sampling behaviour in lekking black grouse (Tetrao tetrix). Females mainly visited males occupying territories in the centre of the lek with relatively large territories. They were also more likely to visit males that had high attendance. The same factors were also correlated with male mating success. A multiple regression model including these factors explained more of the variance in female visits per male (53%) than in mating success (33%). The pattern of female sampling conformed with a pool comparison (best-of-n) tactic. Such a tactic is expected if the costs of sampling are low. Females of high body mass visited more males than lighter females, however, which indicates that females may vary in their search tactics and suggests that there may be search costs. The existence of costs is further suggested by the fact that if the mate from a previous year was still present, females always mated with the same male in the following year. Though search costs were not measured directly, our findings suggest that some costs are negligible (e.g. energetic exhaustion or predation) whereas others (timing of mating) may be more important.     

The influence of potential reproductive rate and variation in mate quality on male and female choosiness in the sand goby, Pomatoschistus minutus

Recent models of choosiness in mate choice have identified two particularly important factors: the potential reproductive rate (PRR) of the choosing sex relative to that of the chosen sex, and the variation in quality of potential mates. This experimental study tested how these factors affected choosiness in male and female sand gobies, Pomatoschistus minutus. We manipulated relative PRR by means of water temperature, and mate quality by means of body length. The choosing male or female was offered a choice between two mates with either a small or a large difference in body length representing a small or a large variation in mate quality. Choosiness was measured as (1) preference for the larger mate, and (2) as whether or not spawning occurred with the smaller mate, while the larger mate was visible but screened off. We found that females preferred large males, and that their level of choosiness was affected by variation in male quality, but not by their own relative PRR. Males, on the other hand, seemed unselective in all treatments and were in general more likely than females to spawn with their provided partner. This suggests that in the sand goby, variation in male mate quality has a greater influence than relative PRR on facultative changes in female choosiness. However, a general difference in PRR between males and females may be one important factor explaining the observed sex difference in choosiness. Received: 17 April 2000 / Revised: 24 June 2000 / Accepted: 17 July 2000     

Variance in female quality, operational sex ratio and male mate choice in a bushcricket

Male bushcrickets, Kawanaphila nartee, exercise mate choice when nutrients are limited. Male mate choice is associated with a female-biased operational sex ratio (OSR) that arises from an increased relative paternal investment under nutrient limitation. However, increased male choosiness could be attributable to the fact that females vary more in fecundity, and consequently in mate quality, when nutrient limited. Our objective was to experimentally partition the influences of OSR (male or female bias) and variance in mate quality (high or low) and to assess their relative influence on the intensity of mate choice by male bushcrickets. Female quality was manipulated by controlled feeding regimes that directly affected female fecundity. We found that males and females engaged in sexual interactions sooner under a male-biased than a female-biased OSR. Males were more likely to reject females on their first encounter when variance in female quality was high. However, the effect of quality variance on the total number of rejections during a 4-h observation period was dependent on the perceived OSR. A male's prior experience of variance in female quality did not influence male choosiness. Our observed rates of mate rejection conformed well with those predicted from recent theoretical models of sexual differences in choosiness. In conclusion, our results show that the opportunity for selection via male mate choice is influenced by an interaction between OSR and the variance in mate quality that arises within nutrient-limited populations of females. Received: 5 January 1998 / Accepted after revision: 25 October 1998     

Are nest characters of importance when choosing a male in the fifteen-spined stickleback (Spinachia spinachia)?

Male fifteen-spined sticklebacks (Spinachia spinachia) build their nests by wrapping epiphytic algae around macroalgae and securing them with secretional threads. In this study, I show that female fifteen-spined stickleback prefer males which have built their nests above the surrounding vegetation. High-located nests are safer, because they are less likely to attract egg-predators such as shore crabs (Carcinus maenas). Thus, I found that shore crabs more rapidly find eggs in nests built close to the bottom than in nests well above the it. Moreover, male-male competition could be an additional explanation as to why males build nests high up. Thus, my field results suggest that the closer the males were to a neighbour, the higher they built. Larger males were also found to have larger territories but male size did not correlate with the height of the nest above the surrounding vegetation. Females showed no preference for nest size, regardless of the presence of egg-predators. However, they tended to choose nests that had no egg-predators in the vicinity over nests with egg-predators close by. Female choosiness for certain nest characters may provide one explanation for the evolution of male care in this species. Received: 29 December / 1999 Revised: 18 April 2000 / Accepted: 28 May 2000     

Signature whistle variations in a bottlenosed dolphin,Tursiops truncatus

To examine whether context-specific information is superimposed upon the individual cues present in the whistling of the bottlenosed dolphin, Tursiops truncatus, parameter variations within the two most frequently emitted whistle types of a captive individual were investigated in three different behavioural contexts. The study concentrated on comparing signal features of spontaneously occurring vocalizations in two possible phases following the performance of a trained discrimination task and those occurring during isolation. Phases of the discrimination task differed according to whether the animal showed correct (reward given) or incorrect performance (no reward). Signature whistles were most common in isolation, but also represented just over half of the whistles following a choice task. Of 14 signature whistle frequency and time parameters measured 9 differed significantly between isolation and at least one of the phases following a choice task (Table 1). Three parameters also varied according to whether performance was correct or incorrect. In contrast, only one out of four parameters (start frequency) measured from the second most frequent whistle type varied significantly between contexts (isolation vs. phase following correct choice). The results indicate that not only identity but also context-related information is available in the whistles of a bottlenosed dolphin.