Sexual differences in reproductive effort: time-activity budgets of monogamous killdeer,Charadrius vociferus

By measuring the time allocated to various reproductive and somatic activities it was possible to look at differences in the strategies of the sexes for killdeer, a monogamous species. Killdeer time-activity budgets supported the general hypothesis that males and females maximize fitness in different ways. Males allocated more time than females to mating activities: mate attraction and territorial defence. However, the parental roles of the sexes were not those predicted for a monogamous species; males allocated more time than females to parental activities throughout a nesting attempt. Males appeared to be less time-constrained, and better able to perform parental activities. Females allocated more time to foraging than did males. It is suggested that the amount of time females allocated to parental activities is limited by the high energy demands of egg production. Further, high nest-failure rates may select for females maintaining good condition for re-nesting. The costs of desertion for killdeer were considerable. Adult removal experiments supported the hypothesis that two parents were necessary for successful reproduction. No individuals managed to raise offspring alone successfully. Lone males, however, were able to hatch chicks while females were not. The need for continuous parental attentiveness may results in selection for biparental care for killdeer.     

Activity budgets and activity rhythms in red ruffed lemurs (Varecia rubra) on the Masoala Peninsula, Madagascar: seasonality and reproductive energetics

The activity budgets and daily activity rhythms of Varecia rubra were examined over an annual cycle according to season and reproductive stage. Given the relatively high reproductive costs and patchy food resources of this species, I predicted that V. rubra would 1) travel less and feed more during seasonal resource scarcity in an attempt to maintain energy balance, and 2) show sex differences in activity budgets due to differing reproductive investment. Contrary to the first prediction, V. rubra does not increase feeding time during seasonal food scarcity; rather, females feed for a consistent amount of time in every season, whereas males feed most during the resource-rich, hot dry season. The results are consistent with other predictions: V. rubra travels less in the resource-scarce cold rainy season, and there are some pronounced sex differences, with females feeding more and resting less than males in every season and in every reproductive stage except gestation. However, there are also some provocative similarities between the sexes when activity budgets are examined by reproductive stage. During gestation, female and male activity budgets do not differ and appear geared toward energy accumulation: both sexes feed and rest extensively and travel least during this stage. During lactation, activity budgets are geared toward high energy expenditure: both sexes travel most and in equal measure, and rest least, although it remains the case that females feed more and rest less than males. These similarities between female and male activity budgets appear related to cooperative infant care. The high energetic costs of reproduction in V. rubra females may require that they allot more time to feeding year round, and that their overall activity budget be more directly responsive to seasonal climate change, seasonal food distribution, and reproductive schedules.     

Orangutan activity budgets: Monthly variations and the effects of body size,parturition, and sociality

The activity budgets of individual orangutans were investigated at the Kutai Reserve, Indonesia. Activity profiles within and between individuals were compared to examine monthly variations in feeding patterns, potential energetic constraints imposed by large body size and parturition, and the costs of sociality. Animals showed monthly changes in travelling, feeding, and resting patterns. Monthly increases in travelling and feeding were associated with marked reductions in the time spent resting. Inter-individual variations in activity budgets did not exist among animals of the same age-sex class. Activity patterns differed, however, as a function of age and sex. Adult females and subadult males travelled and fed significantly longer than an adult male. Parturition had predictable effects on activity; one female reduced her feeding and travelling immediately following parturition. Adult male orangutan sociality appears to be limited by travel costs. Associations with females forced a male orangutan to travel significantly more compared with periods in which he was solitary. The male did not lose an appreciable amount of time feeding when accompanying a female.     

Effect of multiple mating on the reproductive performance of the rice leaffolder moth,Cnaphalocrocis medinalis (Lepidoptera: Crambidae)

To clarify whether multiple mating of females and males affects the reproductive performance of the rice leaffolder moth, Cnaphalocrocis medinalis (Guenée), we examined the effect of the number of matings (once, twice, or three times) for females (female treatment) and males (male treatment) on the incidence of moth mating, number of eggs laid, egg hatchability, and adult longevity. We also compared the effect of multiple mating imposed on males or females separately with the effect of that imposed on both sexes simultaneously (both sexes treatment). The incidence of mating of females and males that mated three times (3-mated females and males) was significantly lower than for females and males that mated twice or once (2-mated or 1-mated females and males). The incidence of mating of 1-mated moths (both sexes) was significantly higher than for 2-mated or 3-mated moths (both sexes). Two-mated or 3-mated females laid significantly more eggs with significantly higher hatchability than 1-mated females. Females that mated with 1-mated males (second male mating) or 2-mated males (third male mating) laid significantly fewer eggs than those that mated with virgin males (first male mating). Females laid significantly more eggs after the second and third matings for moths of both sexes than after the first mating for moths of both sexes. The mechanisms of improvement and decline of female reproductive performance when multiple mating was imposed on males or females are also discussed in relation to the reproductive biology of C. medinalis.     

Behavioural dynamics of biparental care in the dung beetle Onthophagus taurus

In the dimorphic dung beetle Onthophagus taurus major males provide assistance during offspring provisioning. We examined the behavioural dynamics of biparental care to quantify directly how males and females allocate time to parental and nonparental behaviours and to determine whether parents adjust their level of investment relative to their partner's contribution. Females allocated more of their time budget to parental behaviours than males. The proportion of time females allocated to parental behaviours increased after oviposition while that of a male decreased. Male paternity assurance behaviours were negatively associated with male and female parental behaviours. Theoretical models predict that the investment provided by the members of a cooperative pair should be negatively correlated and that any shortfall of one parent should be partially compensated for by the other. In the absence of a male, unassisted females allocated more time to parental care, and performed more parental behaviours. However, compensation was incomplete as unassisted females performed fewer parental behaviours than pairs, resulting in significantly lighter brood masses (the egg and its associated dung supply). Males performed more parental behaviours when paired with small females, and small females more than large females. Contrary to prediction, the investments provided by males and females in a cooperative pair were positively correlated. Males coordinated their parental behaviours with the females rather than acting independently. Since parental behaviours were directly related to the weight of brood masses, the observed parental interactions will have important fitness consequences in this species. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Sex Discrimination via Anal Gland Secretion in a Territorial Monogamous Mammal

Monogamy in mammals is relatively unusual, only occurring in 3–5% and to an even lesser degree in rodents. Monogamy exists in two forms, facultative which evolved due to female dispersion and obligate when male care is needed for offspring survival. Most monogamous rodents use scent marking as a form of territorial defence to obtain exclusive access to vital resources. Intrasexual competition occurs in many species to maintain pair bonds and also to signal presence within a territory. In this study, we tested the hypothesis that resident obligate monogamous Eurasian beaver (Castor fiber) can discriminate between the sexes when investigating anal gland secretions (AGS) from unknown (stranger) mated pairs. We predicted that, due to intrasexual competition, dominant territorial male and female beavers will display a stronger response to AGS of same‐sex conspecifics. Territorial intrusion by mated pair strangers was simulated by the formation of experimental scent mounds (ESM) with AGS. Our results showed that both sexes (1) displayed a stronger response to the male AGS in sniffing duration, (2) physically responded for longer durations towards male AGS and (3) overmarked male AGS more frequently. We infer that obligate monogamous mammals can determine the sexual identity of intruding conspecifics of a mated pair via AGS and that intruding males may pose more of a threat to resident males via intrasexual competition and resident females due to the long‐term costs, such as infanticide and abortion, of a new dominant male.     

Mating experience weakens starvation tolerance in the seed bug Togo hemipterus (Heteroptera: Lygaeidae)

Organisms are exposed to various stresses caused by environmental fluctuations. One of the most common stresses is the shortage of food. Individuals of many species must survive periods of starvation. There appears to be a trade‐off between reproduction and survival. When residual reproductive value declines for an individual, life‐history theory predicts an increase in current reproductive investment. Current reproductive investment differs between virgin and mated individuals. It is likely that mating experience influences starvation tolerance. However, few studies have investigated sex differences in the effect of mating experience on starvation tolerance or clarified the causes of reductions in starvation tolerance in both sexes. In the present study, these questions are investigated using the seed bug Togo hemipterus (Heteroptera: Lygaeidae).The results of the present study demonstrate that mating is costly for both sexes. Mated males show very short survival times and a daily reduction in weight, and daily energy expenditures are significantly greater in mated males than in virgin males. It is possible that starvation increases the mating effort of males, such as behavioural activities and the amount of time spent searching for females. A trade‐off between survival duration and lifetime fecundity is found in virgin females. However, there is no trade‐off in mated females, which have very short survival times. Whether male seminal substances contribute to the short survival times of mated females is considered. This is the first report demonstrating the influence of sex and mating experience on starvation tolerance. Sex‐specific causes for reductions in starvation tolerance are discussed.     

Sex Allocation in Response to Maternal Condition: Different Tactics of Care-Giving by Male and Female Zebra Finches

While behavioral ecologists have studied sex allocation for 30 yr, almost nothing is known about the roles of male vs. female parents in making secondary allocation decisions in species with bi‐parental care. To investigate possible sex differences in tactics in zebra finches, we manipulated the condition of one cohort of females by trimming their flight feathers; a second cohort of females and all males were left intact. Focal observations of feeding behavior revealed that manipulated females did not reduce their feeding time, but rather focused their efforts on rearing sons, while control females allocated effort to rearing daughters. Males mated to manipulated females tended to focus effort to daughters, while control males showed no consistent response. Manipulated females experienced a lower reproductive rate, primarily because of a low daughter rate, relative to control females. Manipulated females also laid smaller clutches and had longer inter‐clutch intervals, which suggest that their ability to metabolize and allocate essential proteins from pectoral muscle mass during the egg‐laying period was impaired. To our knowledge, this study is the first to show that parents of both sexes make active sex allocation decisions when the physiological condition of one parent is manipulated; males do not simply adopt female allocation patterns, but make complementary adjustments. Our results have implications for interpretation of handicapping studies that seek to understand the dynamics of inter‐sexual negotiations of care‐giving. Investigation of sex differences in sex allocation tactics in species with limited dietary protein, such as granivorous birds, is a potentially important direction for future research.     

Reproductive behaviour of mated and virgin females of a solitary sawflyDineura virididorsata

Sex ratios of the arrhenotokous sawflyDineura virididorsata Retz. (Hymenoptera: Tenthredinidae) are variable among populations in the field, but are biased in favour of females. We have concluded elsewhere that the sex ratios of this species cannot be satisfactorily explained by any general sex ratio model that is available. Here we present data on the different behavioural traits that are relevant to interpretation of the evolution of female-biased sex ratios under local mate competition. We report on oviposition behaviour, mating behaviour and the patterns of movement of ovipositing virgin and mated females. Our results show that adult emergence is synchronous and females will mate daily with different males (in the laboratory, at least). This would decrease the relatedness of offspring should it also take place in the field. In addition, ovipositing females (mated and virgin) leave their site of release in a relatively short time and deposit few eggs relative to their capabilities, so siblings are unlikely to be clumped. The time that virgins spend in deposition of an egg is not different from that spent by mated females, so there is no differential ‘investment’ in this regard. These behavioural observations agree with our previous conclusion that the preconditions of current sex ratio theories are not met byD. virididorsata.     

Mating Effect on Sex Pheromone Production of the Oriental tobacco budworm,Helicoverpa assulta

This study was undertaken to clarify the suppression phenomenon of sex pheromone production after mating and its relationship to the physiological mechanism in adult females of Helicoverpa assulta, and determine the mating factor from males causing depletion of sex pheromonc production. Sex pheromone production of H. assulta females was mostly terminated in 3 hours after mating. Mated females maintained with a low titer of sex pheromone until 3 days when it started to increase again, which showed a characteristic of species mating more than once. The mated female again produced pheromone upon injection of pheromone biosynthesis activating neuropeptide (PBAN) or extracts of brain-suboesophageal ganglion complexes (Br-Sg) of mated female, which were shown similar pheromonotropic activities as compared with virgin females. These results indicated that the mating did not inhibit the receptivity of pheromone gland itself and PBAN biosynthesis in suboesophageal ganglion of the mated females. And it seems to support that the depletion of sex pheromone production is responsible for blocking of PBAN release from head. To investigate the mating factor from adult males, when extracts of reproductive organs of male were injected into hemocoel of virgin females evoking depletion of sex pheromone production as shown in mated female. The results suggest that a chemical substance(s) from the male reproductive organs could be responsible for the loss of sex pheromone biosynthesis in H. assulta.     

Discrimination reversal learning reveals greater female behavioural flexibility in guppies

Behavioural flexibility allows an animal to adapt its behaviour in response to changes in the environment. Research conducted in primates, rodents and domestic fowl suggests greater behavioural persistence and reduced behavioural flexibility in males. We investigated sex differences in behavioural flexibility in fish by comparing male and female guppies (Poecilia reticulata) in a reversal learning task. Fish were first trained on a colour discrimination, which was learned equally rapidly by males and females. However, once the reward contingency was reversed, females were better at inhibiting the previous response and reached criterion twice as fast as males. When reward reversing was repeated, males gradually reduced the number of errors, and the two sexes had a comparable performance after four reversals. We suggest that sex differences in behavioural flexibility in guppies can be explained in terms of the different roles that males and females play in reproduction.     

雌星豹蛛性信息素的行为学证据

采用星豹蛛(Pardosa astrigera)成熟雄蛛求偶时潜伏时间、静止时间、身体震动和第一墩步足伸展次数等行为参数,利用行为学方法测定了不同性别、日龄和生殖状态的星豹蛛雌蛛释放的拖丝对雄蛛求偶行为的影响。结果表明,雄蛛第一对步足伸展和身体震动等典型求偶行为是进行星豹蛛性信息素生物测定的可靠评价指标。星豹蛛雄蛛能通过拖丝上的性信息素辨别星豹蛛的性别、日龄和生殖状态。雄蛛在成熟3周未交配雌蛛拖丝处理过滤纸上潜伏时间和静止时间都相应最短,在交配未产卵雌蛛、雌亚成蛛和成熟雄蛛拖丝上时间中等,在卵孵化雌蛛拖丝处理滤纸上潜伏时间和静止时间都相应最长。成熟3周未交配雌蛛和交配未产卵雌蛛释放的拖丝都能引起雄蛛第一对步足伸展和身体震动等典型求偶行为,雄蛛对成熟3周未交配雌蛛拖丝典型求偶行为的频率都相应高于交配未产卵雌蛛。卵孵化雌蛛释放的拖丝虽也能引起雄蛛第一对步足伸展行为,但其伸展频率显著降低;而其它拖丝都不能引起雄蛛典型求偶行为。     

Parental investment asymmetries of a globally endangered scavenger: unravelling the role of gender,weather conditions and stage of the nesting cycle

ABSTRACT

Capsule: In Egyptian Vultures Neophron percnopterus, both sexes invested similar parental effort throughout the breeding period. However, there was variation in the degree of intensity of parental care during some stages of the breeding period, suggesting that sex-role specialization exists for some activities.

Aims: To quantify parental care behaviour of Egyptian Vultures for the first time and to examine the role of sex, weather conditions, and stage of nesting cycle on breeding ecology.

Methods: We monitored 15 nests of Egyptian Vultures to analyse parental care investment. We collected data on nine different behavioural parameters/activities per sex, which were recorded throughout the entire breeding period. Variation in parental investment was analysed using generalized linear mixed models.

Results: Females invested more effort in incubation/brooding (61.45% for females and 31.54% for males) and egg turning (0.45?events/h for females and 0.37?events/h for males) while males contributed more to nest material delivery to the nest (0.67?deliveries/h for males and 0.14?deliveries/h for females). Conversely, both sexes invested the same effort in nestling attendance (21.89% for females and 21.21% for males) and food provisioning (0.28?items/h for females and 0.25?items/h for males). Furthermore, parental investment was not affected by weather, especially during critical moments such as incubation/brooding, however, changeover rate was positively related to temperature.

Conclusion: Our results suggest that, in the Egyptian Vulture, one sex is not entirely responsible for a particular task and the compensatory effort of the other mate is required. Finally, our findings indicate that major events such as incubation onset and hatching caused important shifts in the patterns of parental investment.     

Differential movement and activity patterns of sexes in a biparental beetle during the reproductive season

1. Biparental care is stabilised if parents perform different tasks during care. Specialised parental roles may require different time and energy budgets that in turn are expected to influence the activity and space use of sexes. 2. Here we investigate movement patterns of the biparental Lethrus apterus beetle using a grid of pitfall traps in their natural habitat. 3. Sexes of the burrow building L. apterus perform different roles during caregiving, as females collect most of the leaves, which serve as food for the offspring while paired males stay mostly in the burrow. We hypothesised that sex differences in mate search and parental activities are reflected in movement patterns. 4. We found that females frequently travelled short distances, whereas males were detected less often but when detected, they travelled significantly longer distances than females. 5. Our results are consistent with the notion that efficient parental food provisioning requires more localised movement and activity patterns. Furthermore, the long distance movements of some males may indicate active mate searching behaviour.     

The Sicker Sex: Understanding Male Biases in Parasitic Infection,Resource Allocation and Fitness

The “sicker sex” idea summarizes our knowledge of sex biases in parasite burden and immune ability whereby males fare worse than females. The theoretical basis of this is that because males invest more on mating effort than females, the former pay the costs by having a weaker immune system and thus being more susceptible to parasites. Females, conversely, have a greater parental investment. Here we tested the following: a) whether both sexes differ in their ability to defend against parasites using a natural host-parasite system; b) the differences in resource allocation conflict between mating effort and parental investment traits between sexes; and, c) effect of parasitism on survival for both sexes. We used a number of insect damselfly species as study subjects. For (a), we quantified gregarine and mite parasites, and experimentally manipulated gregarine levels in both sexes during adult ontogeny. For (b), first, we manipulated food during adult ontogeny and recorded thoracic fat gain (a proxy of mating effort) and abdominal weight (a proxy of parental investment) in both sexes. Secondly for (b), we manipulated food and gregarine levels in both sexes when adults were about to become sexually mature, and recorded gregarine number. For (c), we infected male and female adults of different ages and measured their survival. Males consistently showed more parasites than females apparently due to an increased resource allocation to fat production in males. Conversely, females invested more on abdominal weight. These differences were independent of how much food/infecting parasites were provided. The cost of this was that males had more parasites and reduced survival than females. Our results provide a resource allocation mechanism for understanding sexual differences in parasite defense as well as survival consequences for each sex.     

Rearing Temperature Influences Adult Response to Changes in Mating Status

Rearing environment can have an impact on adult behavior, but it is less clear how rearing environment influences adult behavior plasticity. Here we explore the effect of rearing temperature on adult mating behavior plasticity in the butterfly Bicyclus anynana, a species that has evolved two seasonal forms in response to seasonal changes in temperature. These seasonal forms differ in both morphology and behavior. Females are the choosy sex in cohorts reared at warm temperatures (WS butterflies), and males are the choosy sex in cohorts reared at cooler temperatures (DS butterflies). Rearing temperature also influences mating benefits and costs. In DS butterflies, mated females live longer than virgin females, and mated males live shorter than virgin males. No such benefits or costs to mating are present in WS butterflies. Given that choosiness and mating costs are rearing temperature dependent in B. anynana, we hypothesized that temperature may also impact male and female incentives to remate in the event that benefits and costs of second matings are similar to those of first matings. We first examined whether lifespan was affected by number of matings. We found that two matings did not significantly increase lifespan for either WS or DS butterflies relative to single matings. However, both sexes of WS but not DS butterflies experienced decreased longevity when mated to a non-virgin relative to a virgin. We next observed pairs of WS and DS butterflies and documented changes in mating behavior in response to changes in the mating status of their partner. WS but not DS butterflies changed their mating behavior in response to the mating status of their partner. These results suggest that rearing temperature influences adult mating behavior plasticity in B. anynana. This developmentally controlled behavioral plasticity may be adaptive, as lifespan depends on the partner’s mating status in one seasonal form, but not in the other.     

Incubation effort in relation to male attractiveness in zebra finches Taeniopygia guttata

The division of labour in parental care between the two sexes varies between and within species. In birds, parents have been shown to invest more into egg production and nestling care when paired with an attractive rather than an unattractive mate, as predicted by the differential allocation hypothesis. Here we investigate variation in the female's and male's share of incubation behaviour, a vital, and costly, period of parental care during which the embryo is vulnerable to perturbations in developmental conditions. We manipulated the attractiveness of male zebra finches Taeniopygia guttata , using red or green leg-rings. To simulate their natural social environment we allowed them to breed in outdoor aviaries. All males within an aviary were given the same coloured ring to avoid ring-colour related assortative mating. Males within a colony, however, were still expected to show some variation in attractiveness with the earliest laying females possibly pairing with the most attractive males. Indeed we found that both factors played a role in explaining female incubation effort. Among females mated to red ringed males, earlier laying females contributed significantly more to incubation than late laying females, but no such pattern was found in females mated to green ringed males. Overall, there were no differences in the level of incubation provided by both parents between treatment groups, suggesting some compensation within the pair. Hatching success was correlated with a pair's total incubation effort. These results suggest that variation in the division of parental care between the sexes is in agreement with both increased effort of females mated with attractive males, and females compensating for the reduced effort of attractive males seeking further mating opportunities. These two factors can act at the same time in natural populations and both should be considered when explaining variation in division of labour between the sexes.     

Effects of parent age at mating on reproductive response of Glyptapanteles flavicoxis (Hymenoptera: Braconidae), a larval parasitoid of the gypsy moth (Lepidoptera: Lymantriidae)

Glyptapanteles flavicoxis (Marsh) (Hymenoptera: Braconidae) is a gregarious larval parasitoid of the Indian gypsy moth Lymantria obfuscata (Walker) (Lepidoptera: Lymantriidae), that is believed to have potential for inundative releases against gypsy moth populations, because it can be reared in large numbers with few hosts. Unfortunately, sex ratios in laboratory reared G. flavicoxis are usually male-biased, hindering efforts to mass release this species for biological control by making the production of females costly. Because parental age at time of mating is known to affect the sex ratio in some Braconidae, we crossed haploid males and virgin females at 0, 1, 4, 9, and 16 d old with at least 10 trials for each of the 25 combinations. Numbers and sex ratios of progeny produced by females each day were recorded. Both progeny and sex ratios (percentage of females) among progeny produced by ovipositing females of G. flavicoxis decreased markedly over time, so only the first days production need be used in mass rearing. The reduction in the proportion and numbers of females among progeny as females aged is consistent with sperm depletion. Approximately 30% of females in all age classes mated to newly emerged males (day 0) produced all male progeny, whereas only 10-15% of those mated to older males failed to produce any daughters. When crosses with only male progeny were excluded from the analysis, females mated to males 1 d old had higher sex ratios in progeny than those mated to males in other age classes. In addition, females mated the day that they emerged tended to have progeny with the highest sex ratios.     

Biparental care in incipient colonies of the dampwood termiteZootermopsis angusticollis Hagen (Isoptera: Termopsidae)

Parental care and division of labor in male and female primary reproductives in incipient colonies of the dampwood termite Zootermopsis angusticolliswere studied by recording repertory sizes, behavior frequencies, time budgets, and transition probabilities of acts. Repertory sizes for males and females were identical, and behaviors were performed at similar frequencies. Time budget data also suggested equal participation in parental care by both sexes. Transition probability analysis suggested that males and females did not organize their behavior in roles. We therefore found no indication of sex-based division of labor during colony foundation and the incipient stages of colony growth. The lack of behavioral specialization may be due to the fact that acts might be related and/or have more than one function, and different tasks are not spatially associated. Our observations are consistent with the hypothesis that monogamous mating systems in termites involve mate assistance.