Aggression in female Red-Winged Blackbirds: A strategy to ensure male parental investment

Summary Female Red-winged Blackbirds (Agelaius phoeniceus) are often aggressive towards conspecific females during the breeding season. We hypothesize that the function of female-female aggression in this species is to guard the nonshareable portion of the male's parental investment.The investment-guarding hypothesis predicts that a female should be more aggressive toward another female evincing interest in mating with the territory-owning male than toward a female simply perching within the male's territory. Results of mount presentations to females with active nests confirmed this prediction. Nesting females attacked a stuffed conspecific female mounted in a precopulatory, soliciting posture significantly more often than a mount in a normal, perched posture.The male's nonshereable parental care consists of provisioning his young, and most of this care is invested in the brood of his primary (first-to-nest) female. It is therefore predicted that primary females should be more aggressive than secondary (later-nesting) females. Female mount presentations also confirmed this prediction. Primary females attacked the soliciting mount significantly more often than secondary females.     

Early maternal investment in male and female African elephant calves

Summary The suckling behaviour of 130 freeranging elephant calves aged between birth and 4.5 years old was examined in Amboseli National Park, Kenya. Analyses of frequencies of suckling and durations of suckling bouts showed that males attempted to suckle more often, were more successful at their attempts, and as a result were estimated to have a higher milk intake than did female calves. Mothers were equally tolerant of their sons' and daughters' demands to suckle at young ages, but were less tolerant of their older sons' demands. The growth rates of males based on hind footprint length were faster than those of females from birth onwards. During drought years with low food availability, male calf survivorship in the first year was lower than that of female calves. During wet years, there was little difference between sexes in survivorship. It appeared that during dry years mothers were unable to sustain milk production at a level that met the metabolic requirements of their sons, and as result male calves were more likely to die. Females with a surviving son tended to have a longer interbirth interval than did females with a surviving daughter. We suggest that greater early maternal investment in male calves occurs because, in the highly-competitive polygynous mating system of elephants, size in adult male elephants is an important factor in mating success.     

Pheromone-based female mate choice and its effect on reproductive investment in a spitting spider

Numerous studies have focused on whether organisms can signal or perceive pheromones and use chemical signals in species and mate recognition. Recently, there have been an increasing number of studies investigating whether pheromones are used in mate choice. Yet, little attention has been paid in exploring the effects of pheromone-based mate choice on reproductive investment. We first tested this hypothesis by providing virgin Scytodes sp. females with a choice between two virgin males in the presence of chemical signals alone and found strong evidence of an odor-based mate preference. We then examined the consequences of the odor-based mate choice by allowing female Scytodes sp. that had previously made an odor-only mate choice to mate with preferred and non-preferred males, respectively. We measured the success of copulation, mortality of male, pre-oviposition interval, egg-sac weight, egg weight, fecundity, fertility, embryonic period, and size of offspring at hatching. Females that mated with the preferred males produced significantly heavier egg sacs that contained more and larger eggs with a greater fertility. Significantly more non-preferred males than preferred males were killed by spitting. However, pre-oviposition interval, embryonic period, and hatchling size were not affected by female mate choice. This study is the first to demonstrate that female spiders are able to regulate their highly valuable reproductive investment based solely on chemical signals.     

Dominance rank,female reproductive synchrony,and male reproductive skew in wild Assamese macaques

In groups with multiple males, direct mate competition may select for the evolution of dominance hierarchies that sort males into a queue for access to fertile females. The priority-of-access (PoA) model proposed by Altmann in 1962 makes explicit predictions about the resulting paternity distribution based on an interaction between male dominance rank and the overlap of female receptive phases. Here, we investigated whether the logic of the PoA model predicted the distribution of paternity across ranks in a seasonal breeder with high reproductive synchrony over six consecutive mating seasons. We studied 18 males that resided in a group of wild Assamese macaques (Macaca assamensis) in their natural habitat at Phu Khieo Wildlife Sanctuary, Thailand, between 2006 and 2011 with 5 to 13 conceptions per season. We assessed whether mate guarding increased paternity success, described “short-term” deviations from predicted paternity distribution, and examined how these are related to the number of competitors and fertile females. We determined genetic paternity of 43 (93 %) offspring born into the study group and found reproductive skew to be relatively low with 29 % alpha male paternity in accordance with the high degree of female reproductive synchrony observed. Short-term deviations from expected paternity distribution over ranks were not explained by the number of resident males or the number of conceiving females or their interaction. Within the limits of this study, these results suggest that even if males cannot discern female fertile phases, if reproduction is seasonal, and if reproductive synchrony is high, males may also compete directly over access to females.     

Effects of spermatophores on male and female monarch butterfly reproductive success

Summary I present the results of experiments designed to measure the effects of spermatophores produced by male monarch butterflies on male and female reproductive success. There was wide variation in the number of matings by captive males, suggesting the potential for strong sexual selection on males. Male lifespan was not affected by total number of matings, nor did it differ between males that were allowed to mate and those not exposed to females. Two effects of spermatophores on female behavior or fecundity are reported; (1) Females that received large spermatophores delayed remating longer than those receiving small ones. (2) Females allowed to mate several times laid more eggs than singly-mated females. The relative importance of these effects is discussed in relation to monarch mating patterns.     

Costs of an immune challenge and terminal investment in a long-lived bird

An induced immune challenge can have two counteracting effects on an individual's reproductive investment. (1) The resource demand could increase to "fuel" the immunologic reaction, which in turn can lead to an adaptive decrease in investment in resource-costly activities, such as reproduction. One the other hand, (2) the individual could assume that the immune activity it experiences is indicative of a serious infection. The latter can lead to an adaptive increase in reproductive investment in response to the reduced prospects of survival and future reproduction, so called "terminal investment." To measure such life-history-related consequences of increased immune activity, one group of incubating female Common Eiders (Somateria mollissima) was injected with a nonpathogenic antigen (sheep red blood cells, SRBC) while controls were injected with sterile saline. The eider is a long-lived sea-duck. Females, who incubate the eggs and care for young without assistance from the male, engage in facultative anorexia during incubation leading to a large reduction in body mass. Eiders can abandon their young to other females at the cost of reduced young survival. The immune challenge resulted in a larger mass loss, a prolonged incubation period, and reduced return rate, demonstrating both short- and long-term costs of immune challenge. Additionally, in response to what might have been interpreted as reduced survival chances in immune-challenged females, these females more often tended their own brood after hatching, despite having suffered higher costs during incubation.     

Factors affecting male and female reproductive success in a chipmunk (Tamias sibiricus) with a scramble competition mating system

While sexual selection in mammals with female defense systems has historically received considerable attention, much less is known about the factors affecting reproductive success in mammalian species with scramble competition mating systems. Using mark–recapture techniques and DNA microsatellite loci to assign parentage, we examined the mating system and factors affecting the variation of the annual reproductive success in an introduced population of the Siberian chipmunk, Tamias sibiricus, a small, monomorphic, solitary squirrel. Our analyses showed that breeding females were spatially dispersed, setting the stage for a scramble competition mating system. Male reproductive success was positively associated with the size of the home range. The strength of sexual selection on this behavioral trait was very strong, equaling previous estimates for morphological traits in female defense mating systems. These findings suggest that a behavioral trait, space use, strongly influences the annual reproductive success in males, which is consistent with expectations in a scramble competition mating system. In both sexes, reproductive success was influenced by habitat, with twice as many juveniles produced in semi-open than in closed habitat, possibly due to differences in food availability between the two habitats.     

Preference for male traits in female wolf spiders varies with the choice of available males,female age and reproductive state

Individual variation in female preference for male traits may influence mate choice, especially if benefits and costs of choosiness vary with the range of available males or reproductive timing. We examined variation in female preference for male leg tuft size in Schizocosa ocreata (Hentz) wolf spiders with video playback. Dichotomous (simultaneous) choice test experiments included all possible combinations of four stimuli (i.e., modified versions of the same video male stimulus): average tuft size (control), reduced (−25%), enlarged (+25%), and no tufts (removed). Females exhibited a directional preference for larger tuft size independent of the nature of the choice (except for reduced tufts vs no tufts where no difference was seen). Female preference in the short term (over a period of 4 days) was also highly repeatable for control vs reduced tufts, but not for control vs enlarged tufts. Responses of females in ‘no-choice’ presentations of a single (control) male stimulus varied with age post-maturity; females were less receptive in weeks 1 and 2, highly receptive at week 3, and less thereafter. Mated females were least receptive and most aggressive towards a male stimulus. Females offered choices repeatedly at different ages post-maturity consistently preferred the control male vs reduced tufts over all 3 weeks but varied in their preference for enlarged tufts vs control male. In the first 2 weeks, females preferred the enlarged tuft male stimulus, but showed no preference by the third week. Females tested in week 4 showed no preference in either choice. Results suggest that the potential interaction between female preference for male traits and female reproductive timing may be a critical consideration in mate choice.     

Age,experience and reproductive performance in a long-lived bird: a hormonal perspective

The ultimate factors influencing age-specific reproductive performance in birds have been widely discussed, and several hypotheses have been suggested to explain why young/inexperienced breeders have lower reproductive success than older individuals. In comparison, proximate factors and, particularly, hormonal mechanisms influencing age-related reproductive performance have received lesser attention. In this paper, we examined how baseline levels of corticosterone and prolactin, two hormones involved in reproduction, changed with age and experience in a long-lived bird, the Black-browed albatross (Thallasarche melanophris) during the brooding stage. Corticosterone levels were not linked to age, whereas prolactin levels increased until individuals reached 15 years of age. First-time breeders had higher corticosterone levels and lower prolactin levels than experienced ones. Corticosterone levels were not correlated with breeding experience among experienced birds, whereas prolactin levels slightly increased with advancing experience. Among experienced breeders, there was no effect of individual quality on corticosterone and prolactin levels. Baseline corticosterone and prolactin levels were respectively, positively and negatively correlated to time spent fasting/brooding on the nest. Moreover, the probability of successfully fledging a chick was negatively related to corticosterone levels, but not to prolactin levels. Elevated corticosterone levels in first time breeders may serve as evidence for depleted body reserves resulting from lower foraging/brooding capabilities and therefore support the constraint hypothesis. Low prolactin levels in young/inexperienced birds may be interpreted either as evidence for their lower breeding capacities (constraint hypothesis) or for their limited breeding investment (restraint hypothesis). Finally, we report, for the first time, the hormonal changes associated with the onset of senescence. The very old and most experienced birds, which had the lowest probability of successfully fledging a young, displayed elevated corticosterone levels and low prolactin levels, possibly indicating a degradation of breeding skills and/or a disruption of the endocrine system in senescent birds.     

The relationship between presumed gamete contribution and parental investment in a communally breeding bird

Summary Measures of relative gamete contribution were related to degree of parental investment for both males and females from six groups of communally breeding pukeko (Porphyrio porphyrio). If there is a fitness cost to participation in parental duties, then for males parental investment should relate to degree of confidence of paternity while for females investment should relate to the number of eggs laid. Data were analysed in two ways: (1) by looking at individuals of the same sex within the same group to see if males that had a greater probability of paternity and females that had greater number of eggs in a nest subsequently invested relatively more in parental duties and (2) by pooling the data from the six groups and determining if there was a positive correlation between measures of presumed gamete contribution and amount of parental investment. The results from both types of analyses are variable and give no clear support for a positive relationship between presumed gamete contribution and parental investment even though a possible fitness cost is suggested. Factors which might influence investment are discussed.     

Racketed tail of the male and female turquoise-browed motmot: male but not female tail length correlates with pairing success,performance, and reproductive success

Both males and females of many avian species maintain elaborate plumage traits, and elaborate monomorphic plumage may convey adaptive benefits to one or both sexes as inter- or intraspecific signals. Both sexes of the turquoise-browed motmot (Eumomota superciliosa) are elaborately plumed with long racket-tipped tail. I investigated whether the racketed tail functions as a sexually selected signal in one or both sexes by testing the predictions that males and/or females with the largest tails have: (1) greater pairing success, (2) greater reproductive performance (clutch-initiation date, clutch size, and hatching success), and (3) greater reproductive success. Yearling males with longer denuded rachises (wires) on the central tail feathers had greater pairing success. In addition, adult males with longer wires paired with females who laid larger clutches, had greater hatching success independent of clutch size, and fledged more young. There was no relationship between female tail plumage and pairing success, reproductive performance, or fledgling success. These results are consistent with the hypothesis that male tail plumage functions as a mate choice or status signal, but that the tail of the female does not function in a sexually selected context. I discuss alternative hypotheses for the evolutionary maintenance of the elaborate female tail plumage.     

Innate responses to male sexual harassment in female mosquitofish

Male mosquitofish are very persistent in their sexual activity and harass any female they encounter. Gravid females pay a large tribute to this intense male sexual activity in terms of reduced foraging efficiency. Previous observations have demonstrated that gravid females, when chased by a male, dilute male harassment by moving closer to other females to form shoals. They also approach other males to promote male competition, and when males differ in size, they preferentially target large males, whose harassment is less intense. In this study, we tested whether the modulation of females’ social preferences in response to male harassment is innate or learned. We tested social preference in three groups of females that differed in experience of sexual harassment and in the factors affecting it. Females of the first group were reared without any sexual experience, and pregnancy was induced through artificial insemination. The second group was composed of naive females kept singly with a male; these females experienced sexual harassment but were prevented from experiencing the effects of male–male competition and shoaling on the amount of male sexual harassment. In the third group (controls), females were reared in multi-male, multi-female groups and could experience the modulating effects of social interactions on sexual harassment. When exposed to a harassing male, females of the three groups immediately reduced their distance from another female, approached a group of males or moved toward the larger of two available males. Moreover, the results for these three groups of females were similar to those obtained in wild-caught females that were tested in the same three tests in a previous study (Dadda et al. An. Behav., 70:463–471, 2005). This suggests that the strategies adopted by females in response to male sexual harassment do not need to be learned through specific experience of the social contexts.     

Optimality of parental investment: The influence of nursing on reproductive success of mother and female young house mice

Summary The amount of parental milk investment determines not only the number of young the parent can produce, but also affects the offspring's fitness. The antagonism between quantity and quality of offspring was investigated in laboratory mice.In nursing first litters containing 2,4, ..., 14 pups, mothers invested increasing amounts of milk. This mainly extended the intervals between their first and second litters and slightly increased the size of their second litters, whilst third litters were not affected. In the female young, the decreasing amount each individual received as a result of increasing litter size led to delayed birth and reduced the size of their first litters. The intervals between their first and second litters and the size of the second litters were also affected, although to a lesser extent.Taking these results into consideration on a standard lifetime pattern of reproduction, the effects were calculated on net reproductive rates as an expression of fitness. Death rates were assumed that referred to population equilibrium or population growth. In both situations, the fitness curves of mothers and young showed that the actual level of milk investment and the level yielding optimal reproduction were in agreement. There was no significant deviation towards increased investment, as might result from parent-offspring conflict.     

Variance in male and female reproductive success in a harem-polygynous mammal,the black-tailed prairie dog (Sciuridae: Cynomys ludovicianus)

Summary Black-tailed prairie dogs are colonial rodents that live in contiguous social groups called coteries. A typical coterie contains one adult (2 years old) male, three or four adult females, and several yearlings and juveniles of both sexes. A large coterie sometimes contains two or more adult males. Using detailed behavioral observations on 164 females (of which 160 copulated) and data from four polymorphic loci from parents and offspring of 121 litters, we examined the black-tail mating system.Most females (101/164=62%) copulated with a single adult male, and only 3 of the 102 litters with 2 offspring (3%) showed unequivocal evidence of multiple paternity. Adult males usually copulated with several different adult females.In one-male coteries, females usually copulated exclusively with the resident adult male (RAM) (82/112=73%); this trend was confirmed by electrophoresis of blood smaples. In multimale coteries, each female frequently copulated with at least two different RAMs (28/52=54%); in 4 of 5 multimale coteries (80%) which produced two or more litters whose paternities could be unequivocally resolved by electrophoresis, two different RAMs each sired at least one litter.Of the 164 females, 30 (18%) copulated with both the RAM (or one of the RAMs, in multimale coteries) and an extracoterie adult male, but only 3 (2%) copulated exclusively with an extracoterie adult male. Electrophoresis showed that 9 of 121 litters (7%) were sired by an extracoterie adult male.Intersexual comparisons of annual reproductive success and lifetime reproductive success both indicate that black-tails are polygynous (i.e., that variance in reproductive success is greater for males than for females).     

Determinants of male reproductive success in wild long-tailed macaques (Macaca fascicularis)—male monopolisation, female mate choice or post-copulatory mechanisms?

One of the basic principles of sexual selection is that male reproductive success should be skewed towards strong males in species with anisogamous sex. Studies on primate multi-male groups, however, suggest that other factors than male fighting ability might also affect male reproductive success. The proximate mechanisms leading to paternity in multi-male primate groups still remain largely unknown since in most primate studies mating rather than reproductive success is measured. Furthermore, little research focuses on a female’s fertile phase. The aim of this study was to investigate the relative importance of male monopolisation and female direct mate choice for paternity determination. We also investigated the extent to which paternity was decided post-copulatory, i.e. within the female reproductive tract. We used a combined approach of behavioural observations with faecal hormone and genetic analysis for assessment of female cycle stage and paternity, respectively. The study was carried out on a group of wild long-tailed macaques living around the Ketambe Research Station, Gunung Leuser National Park, Indonesia. Our results suggest that both male monopolisation and post-copulatory mechanisms are the main determinants of male reproductive success, whereas female direct mate choice and alternative male reproductive strategies appear to be of little importance in this respect. Female cooperation may, however, have facilitated male monopolisation. Since paternity was restricted to alpha and beta males even when females mated with several males during the fertile phase, it seems that not only male monopolisation but also post-copulatory mechanisms may operate in favour of high-ranking males in long-tailed macaques, thus reinforcing the reproductive skew in this species.     

Altered responses to female odors in parasitized male mice: neuromodulatory mechanisms and relations to female choice

There is accumulating evidence that females may preferentially select parasite-free or -resistant males. Minimal attention has, however, been paid to the mate preferences and responses of the parasitized male hosts themselves. Here, we considered the effects of parasitic infection on male host mate responses, the neuromodulatory correlates of these responses, and the relations of these responses to female mate choice. Using an odor “preference” test, we examined the effects of different stages of an acute, sub-clinical infection with the naturally occurring, enteric, single host, protozoan parasite, Eimeria vermiformis, on the responses of male mice, Mus musculus domesticus, to the odors of estrous females along with the responses of uninfected females to the parasitized males. At 4 days post-infection (non-infective, pre-patent stage) E. vermiformis-infected male mice showed a significantly decreased preference for the odors of estrous females, whereas at 10 days post-infection (infective, patent stage) infected males showed a significantly increased preference for the odors of estrous females. Parasitized males displayed no significant changes in their responses to the odors of non-estrous females, supporting effects on the reproductively related responses of the host. In parallel, estrous females displayed a reduced interest in the odors of infected males. Least interest was expressed in the odors of the patent, infective males, consistent with the avoidance of contagion. Using selective opioid peptide receptor agonists and antagonists we found evidence that enhanced kappa opioid peptide (e.g., dynorphin) activity was related to the decreased sexual interest of the pre-infective males, while augmented delta opioid peptide (e.g., enkephalin) activity was associated with the enhanced responses of the infective males to females. We further showed that acute kappa opiate administration reduced the responses of uninfected males to females and that uninfected females displayed modified responses to the odors of uninfected males subject to acute modifications of opioid activity. We suggest that these differential shifts in endogenous opioid activity in the parasitized males are associated with and, or related to alterations in neuro-immune and endocrine functions. These findings show that parasitic infection can have, depending on the stage of infection and associated neuromodulatory changes, either significant facilitatory or inhibitory effects on male host preferences for and responses to females. Received: 22 April 1996 / Accepted after revision: 15 March 1997     

Plasma sex steroid profiles and the seasonal reproductive cycle in male and female winter flounder,Pleuronectes americanus

The pattern of seasonal gonadal development and variations in plasma sex steroids were investigated in adult male and female winter flounder, Pleuronectes americanus (Walbaum), from Conception Bay Newfoundland beginning August 1987 to December 1988. The winter flounder reproductive cycle can be divided into five consecutive phases of relative reproductive activity including: (1) rapid gonadal recrudescence in the fall (August–December); (2) continued slow gondadal growth in females, or maintenance of the well developed gonads in males, during the winter (December–February); (3) a prespawning phase of gonadal maintenance in the spring (March–April); (4) spawning early in the summer (May–June) after the female gonads reach peak weight; and (5) the summer postspawned period (June–August) when the gonads remain regressed. Female gonadal recrudescence in August is characterized by small increases in plasma estrogen levels and recruitment of small oocytes (150 m) into yolk accumulation. For the winter months, estradiol-17 levels in the plasma remain stable, approximately 15 ng ml^-1, until rising again together with testosterone to peak hormone levels just prior to spawning in conjunction with the highest seasonal values for the gonadosomatic index (GSI) and oocyte diameter. After spawning, the ovaries are regressed and sex steroids in the plasma fall to very low levels. Rapid seasonal recrudescence of the gonads in males is evident from rising GSI values, which reach a maximum in October, and from substantial early seasonal increases in plasma testosterone and 11-ketotestosterone. Afterwards, although GSI values subsequently decline presumably reflecting the process of spermiogenesis and/or onset of spermiation in some males, the testes remain relatively well developed with the presence of sperm throughout the spawning season. As the proportion of spermiating males increases, the plasma levels of the androgenic steroid hormones rise to peak circulating levels from April to June. At the end of the summer reproductive season, the testes of post-spawned males become regressed and plasma testosterone and 11-ketotestosterone levels fall, reaching the lowest seasonal values.     

Age-related changes in reproductive effort of male bison

Summary The behavior of male bison during the rut was studied, to test the prediction that reproductive effort should increase with age. Because bull bison do not show parental care, the major component of reproductive effort is competition with other bulls to obtain copulations with estrous females. Data were collected on activity budgets during the pre-rut and rut seasons, on interactions with other bulls, and on proximity to tended females during the rut. Participation in the rut started at ages 5–6. Older bulls showed greater percent time active and less percent time eating than younger bulls. The older bulls also had higher counts of dominance and fight-related interactions and lower counts for behavior associated with submission than did the younger bulls. Risk taken seemed to increase with age since bulls aged 6 to 12 participated more frequently in dangerous activities associated with fighting. Within the 6 to 12 year old group, activity patterns changed little with age. However, from the interaction information, peak ages of 8 to 11 years appeared. For the activities: back up, chase, clash, head nod, jump away, move away, run away, run toward, push heads, dominant, and risk, 11 and 12 year olds more closely resembled younger bulls. This suggests a reproductive effort curve that is flat at zero until about age 6, increases to age 8, flattens out again to around age 10, and then decreases somewhat. Reproductive effort in male bison thus generally conforms with life-history theory predictions.