Male reproductive success in free-ranging feral horses

In the social organization of feral horses, adult males compete to monopolize groups or bands of females, sometimes called harems. Alternative male strategies are to remain alone or with other bachelors or, less commonly, to accept subordinate status within a harem. The hypothesis that dominant harem stallion status confers a reproductive advantage was tested in free-ranging feral horses. The presence of foals in harems headed by vasectomized (VSX) versus intact stallions was used to assess the ability of these stallions to control reproduction in their harems. Of harems headed by VSX stallions, 17 and 33% contained foals during years 2 and 3 post-treatment, respectively. In contrast, 86 and 80% of harems headed by non-VSX stallions contained foals in those years. Acquisition of pregnant mares appeared more likely than sneak copulations by bachelor stallions to account for foals in harems with a single stallion. However, most foals were born into harems that included a subordinate stallion, an occurrence that was undoubtedly exacerbated by the extended breeding season resulting from the sterility of the harem stallion. Thus, in comparing alternative reproductive tactics, bachelors appeared less successful than subordinate stallions within a harem. However, the highest reproductive success was achieved by the harem stallion, further demonstrating that alternative tactics are not equally profitable. Received: 13 July 1999 / Received in revised form: 24 July 1999 / Accepted: 24 July 1999     

Are low infidelity rates in feral horses due to infanticide?

A growing number of studies conducted on diverse taxa have shown that extra-pair/group paternity is higher than what would be predicted from behavioral observations alone. While it may be beneficial for females to mate with multiple males, this often results in offspring not sired by the behavioral father, which could influence offspring survival, especially in social mammals. Feral horses (Equus caballus) maintain stable social relationships over several years, usually with one stallion defending a harem band of unrelated mares against other males. Sneak copulations by subordinate males have been observed and mares sometimes change bands, both of which can result in foals sired by males other than the dominant band stallion. We measured female fidelity in free-ranging feral horses in 23 bands, with 51 foals over four foaling seasons and tested offspring paternity against parental behaviors. We used 12 polymorphic microsatellite loci and the program CERVUS 2.0 to determine and exclude potential sires. The majority of mares remained in the band with the sire of their foal resulting in most foals being sired by the band stallion. Most foals that were not sired by the band stallion were born in the year after a round-up and we could not determine if they were the result of band changing or sneak copulations. Foals born into a band without their sire had lower survival rates and mothers were significantly more protective of foals not sired by the band stallion. These findings suggest that band stability increases the reproductive success of mares and support the importance of infanticide risk in equid social structure.     

Aging and improving reproductive success in horses: declining residual reproductive value or just older and wiser?

In many mammalian species, female success in raising offspring improves as they age. The residual reproductive value hypothesis predicts that each individual offspring will be more valuable to the mother as she ages because there is less conflict between the current and potential future offspring. Therefore, as mothers age, their investment into individual offspring should increase. Empirical evidence for an influence of declining residual reproductive value on maternal investment is unconvincing. Older mothers may not invest more, but may be more successful due to greater experience, allowing them to target their investment more appropriately (targeted reproductive effort hypothesis). Most studies do not preclude either hypothesis. Mare age significantly influenced maternal investment in feral horses living on the North Island of New Zealand. Older mares, that were more successful at raising foals, were more protective for the first 20 days of life, but less diligent thereafter. Total maternal input by older mothers did not seem to be any greater, but was better targeted at the most critical period for foal survival and a similar pattern was observed in mares that had lost a foal in the previous year. In addition, older mothers were more likely to foal in consecutive years, supporting the hypothesis that they are investing less than younger mares in individual offspring. Therefore, older mothers seem to become more successful by targeting their investment better due to experience, not by investing more in their offspring. Received: 22 September 1999 / Received in revised form: 30 October 1999 / Accepted: 6 November 1999     

Promiscuous behaviour disrupts pregnancy block in domestic horse mares

Based on questionnaires from horse breeders, we found that bringing a pregnant mare which had been mated away from home into a vicinity of a familiar male who was not the father of her foetus increased probability of pregnancy disruption. These mares aborted in 31% of cases, while none of those mated within the home stable aborted. Repeated sexual activity either by a stallion or dominant gelding from the normal home group was observed shortly after the mare came from away-mating. Pregnant mares isolated from home males by a fence were even seen soliciting them over the fence. We speculate that, once returned to the home “herd”, and introduced to familiar males, mares were more likely to terminate their pregnancy to save energy and avoid likely future infanticidal loss of their progeny by dominant male(s) of the home social group. This is a newly discovered phenomenon where a mare manipulates the male’s paternity assessment by promiscuous mating. It may explain a common increased incidence of foetal loss in domestic horses occurring in nearly 40% of pregnancies. We conclude that the common practice of transporting the mare for mating and then bringing her back to an environment with males, stallions or geldings, which did not sire the foetus, is the main cause of high percentages of pregnancy disruption in domestic horses.     

Forced copulation as a conditional alternative strategy in camel crickets

In the animal kingdom, males typically acquire reproductive access to females through courtship, nuptial gifts, or combat. In many species, small or poor-condition males that are unable to produce an attractive display, proffer an adequate mating gift, or dominate in male–male competition may resort to forced copulation. Forced copulation has been previously demonstrated in the camel cricket Pristoceuthophilus marmoratus and is uniquely facilitated by male hind leg weaponry that functions in both intrasexual competition and sexual coercion. Because male leg armaments are positively allometric, the largest males would seem best equipped to force-copulate, although they might be the most attractive to females, and so least likely to adopt a forced copulation strategy. In order to resolve this potential paradox and determine which males are in fact most likely to force-copulate, we manipulated male body size using diet and performed mating trials. Results showed that (1) male size increased with diet quality, (2) females appear to prefer larger males, and (3) smaller males were more likely to attempt forced copulation. Our results thus suggest that forced copulation is a condition-dependent alternative mating tactic used by small males unattractive to females.     

Variations in the birth sex ratio and neonatal mortality in a natural herd of horses

Variations in birth sex ratios and sex differences in juvenile mortality occur in a number of mammalian species, and in many cases have been linked to resource availability. Most of these biases in offspring sex ratios concern polygynous species with pronounced sexual dimorphism, and where females only are philopatric. Data on species with unusual life-history strategies, such as slight sexual dimorphism or dispersal by both sexes, are of particular interest. In this study of a natural herd of horses (Equus caballus) which experienced an eruptive cycle, and therefore a period of nutritional stress, male offspring had higher neonatal mortality rates in nutritionally poor years than in good ones, whereas “year quality” had no effect on the mortality of female offspring; year quality could therefore be used by mares as predictor of sex-specific offspring survival. We show that the environmental conditions that predicted lower survival of males were negatively related to their production: the birth sex ratio the following year was female-biased; and mares were less likely to produce a son when they had produced a son the preceding year. There was no significant effect of mother's parity, age or rank, or the timing of conception or birth on offspring sex ratios. The mechanism leading to biases in the birth sex ratio could have been the loss of male embryos by mares that did not foal. As there was no evidence for selective abortion of male foetuses in females that did foal the next year, it is not necessary to invoke maternal adjustment, though this remains a possibility. Finally, there was a suggestion that male offspring were more costly to raise than females, since mothers that reared a son in poor years tended to experience an increase in the interbirth interval between their two subsequent offspring. Received: 28 December 1996 / Accepted after revision: 27 July 1997     

Direct and indirect estimates of sperm precedence and displacement in the dragonfly Sympetrum danae (Odonata: Libellulidae)

Summary The non-territorial dragonfly Sympetrum danae shows a high degree of post-copulatory mate-guarding, which suggests precedence of the last male's sperm. Irradiated male techniques revealed that the last male that mates with a female fertilizes an average of 95% (100% in 28% of the cases) of the clutch laid immediately after copulation, irrespective of any previous matings. Sperm volume estimates in both sexes and sperm opacity changes in females at 7 stages of copulation and oviposition indicated that males remove 41–87% of previously deposited sperm from the female's storage organs during the first 5 min of the copulation. Ejaculation takes place during the remaining 10–15 min. We maintain that indirect estimates of precedence in libellulids, based on sperm volume changes, always understimate reality because of the supplementing effect of a first-in, last out mechanism. Because males are skilled at achieving sperm precedence, they are forced to be good guards as well, since a risk of take-overs exists. Females are believed to benefit from contact guarding because it results in a lowered risk of male harassment and predation as well as a lower energy expenditure during oviposition.     

A female evolutionary response when survival is at risk: male harassment mediates early reallocation of resources to increase egg number and size

One unexplored area in sexual conflict studies is the female physiological costs and possible resource reallocation that accompany evolutionary costs due to male harassment. Using females of the damselfly Hetaerina americana, I first investigated whether male harassment affected female mating rate and survival and explored whether such effects induced a resource allocation from immunity (in the form of phenoloxidase activity) and muscular fat reserves to egg number and size. Using two seasons that differed in male harassment, it was found that the higher the male harassment, the fewer are the female matings and the lower is the female survival. These results were corroborated using an experimental approach in which a situation of high male harassment was induced. It was also found that when the first mating takes place and at high male harassment, females had more reduced phenoloxidase activity and fat reserves and tended to lay most of the eggs they produce in their lifetime and these were considerably large. However, at low male harassment, egg number and size were more equally produced across matings. Females under high male harassment seemed to suffer the survival costs but may show a plastic evolutionary response of reallocating resources to egg traits to maximize fitness.     

Mixed reproductive strategy and mate guarding in a semi-colonial passerine,the swallow Hirundo rustica

Summary Both male and female swallows Hirundo rustica have a mixed reproductive strategy (parental care for offspring and extra-pair couplations). Mate guarding protects females from male harassment and male swallows from being cuckolded. Females hide their fertile period by copulating successfully with their mates for an extended period during first clutches. Males guard in the pre-fertile period, when many unpaired males are present. Early breeding swallows guard more than late breeders since more sexual chases of females by non-mate males take place in the early period. Solitarily breeding females experience few chases by strange males; copulation frequency, length of copulation period and mate guarding is adjusted to a lower level than among colonial birds. Male guarding activity is more intense in the fertile than in the pre-fertile period. Guarding reduces success of extra-pair copulation attempts.Three female swallows each paired and copulated with a single male and later changed to a new male before starting to breed. Extra-pair copulations most often take place between neighbouring swallows in the fertile period of the female. Many old, early breeding males and many young, late breeding females participate in extra-pair couplations. Successful extra-pair copulations peak in the fertile period contrary to success of pair copulations which does not change during the copulation period.     

Status,reproductive success and fitness in Cape mountain zebra (Equus zebra zebra)

Summary Demographic data relating to herd size and stability are given for a population of Cape mountain zebra (Equus zebra zebra) under longterm observation. Temporal dispersion patterns of male and female offspring differed and were independent of the mother's status. Dispersion in females appeared to be related to physiological state, and dispersion in both sexes was related to age rather than changes in parental behaviour. Reproductive success of dominant and subordinate mares was equal and independent of age and social and reproductive variables. Fitness of dominant mares, however, was significantly higher than that of subordinates, the latter having a higher foal mortality, part of which could be attributable to dominants' aggression. The fitness of all males born was 1.6:1 compared with all females. Dominant mares produced significantly more daughters than sons. This trend was not found for subordinates. Mother's status was positively correlated with dominant status in her female offspring but not related to the subsequent status of her sons. Daughters had a more than twice as great a chance of breeding than sons. For maximum fitness gains, therefore, dominant mares should produce more daughters, since a high proportion of these would also have high status and fitness. This tendency is reflected in the sex ratio skewed towards females found for dominant mares.     

Social grouping and maternal behaviour in feral horses (<Emphasis Type="Italic">Equus caballus</Emphasis>): the influence of males on maternal protectiveness

The risk of infant injury or mortality influences maternal behaviour, particularly protectiveness. Mares are found in bands with a single stallion or bands with more than one stallion in which paternity is less certain. We investigated maternal behaviour in relation to band type. Mares in bands with more than one stallion were more protective of their foals, particularly when stallions and foals approached one another. The rate of aggression between the stallion and foal was a significant predictor of maternal protectiveness, and mare protectiveness was significantly correlated with reduced reproductive success in the subsequent year. Mares that changed band types with a foal at foot, or had their band type experimentally altered, were more protective of their foal in multi-stallion bands than they were in single-stallion bands. Equids are unusual amongst ungulates in that infanticide and feticide have been reported. Both occur where paternity has been uncertain, and equid social structure is similar to other species in which infanticide has been reported. Stallions benefit from infanticide as the mare has greater reproductive success in the subsequent year. Stallion aggression is a significant modifier of mare behaviour and maternal effort, probably due to the risk of infanticide.     

Love bites: male fang use during coercive mating in wolf spiders

Evolutionary conflicts of interest between the sexes are common, as mating tactics and strategies that increase fitness benefits for one sex may incur costs for the other. As a consequence, antagonistic coevolution between the sexes often results in a complex arms race between male persistence and female resistance. Coercive mating (e.g., forced copulation) likely benefits males by increasing the probability of mating; however, costs to females may be high, including injury or even death. Here, we report on a study of the use of fangs by males of the wolf spider Schizocosa ocreata (Hentz) during forcible mating with resistant females, which results in hemolymph loss and scar tissue formation in females. Microscopic inspection revealed that fang wounds (evidenced by scar tissue) were absent on unmated (virgin) females but were found on mated females and were significantly more frequent in coercively mated females. Experimental fang immobilization studies found no difference in copulation success between experimental (fangs immobilized) and control (sham manipulation) males. However, males that had use of their fangs had significantly longer copulation duration. These findings represent a previously unknown male mating tactic in spiders and an unrecognized potential cost of mating for female spiders.     

Mate guarding and copulation behaviour in monogamous and polygynous blue tits: do males follow a best-of-a-bad-job strategy?

This study investigates the importance of mate guarding for males and females in the facultatively polygynous blue tit Parus caeruleus. We present observational data in combination with a paternity analysis using DNA fingerprinting to show that (1) male blue tits guard their mate, since they stay closer to their mate, initiate fewer flights and follow their mate more often during the female's presumed fertile period; (2) polygynous males do not suffer more from lost paternity despite lower mate guarding; (3) in monogamous pairs there is either no relation or a positive relation (depending upon the variable measured) between measures of mate guarding intensity and the proportion of extra-pair young in the nest; and (4) monogamous males that are more often followed by their fertile female suffered less from lost paternity. We conclude that, despite mate guarding, paternity seems to be largely under female control and unattractive males guarding their mate are making the best of a bad situation. Experimental evidence is provided showing that when males were temporarily removed from their territory, their mate suffered from increased harassment from neighbouring males that intruded in the territory and tried to copulate with the female. Almost all of these copulation attempts were unsuccessful because females refused to copulate. We conclude that mate guarding may be beneficial for females because harassment by neigbouring males is prevented.     

Copulatory tactics in relation to sperm competition in Rocky Mountain bighorn sheep

Summary Sperm competition arises in the bighorn mating system because receptive ewes are some-times defended and mated in sequence by a series of increasingly dominant rams and because subordinate (coursing) rams commonly succeed in forcing matings with defended ewes. To identify potential components in a male strategy of sperm competition, the social circumstances and schedule of copulation among free-ranging bighorns were investigated. Three such components emerged: (i) Defending and coursing rams copulated with estrous ewes at extraordinarily high rates, presumably to increase the proportion of their own versus rivals' sperm present at fertilization by simple numerical preponderance at insemination. (ii) Coursing but not defending rams concentrated copulations later in the period of estrus (sensu this study). Evidence from domestic sheep suggests that this may correspond to the period of maximum female fertility. A greater risk and/or reduced benefit associated with forced versus cooperative copulation may force coursing rams to restrict copulation to periods of highest fertility, whereas defending rams may gain even from copulations outside this period. (iii) Defending rams copulated at unusually high rates immediately after bouts of successful coursing. Such retaliatory copulations were associated with reduced amounts of courtship and were more likely to occur if the defending ram had copulated infrequently recently. The number of retaliatory copulations was independent of the number of preceding forced copulations, probably because defending rams rationed ejaculates in anticipation of future coursing. The function of retaliation is not clear, although the literature on domestic sheep suggests the possibility of some form of mechanical displacement of rival sperm. Unlike females of many ungulate species, bighorn ewes were typically receptive to cooperative copulation over a period of days. Although persistent courtship by defending rams may coerce such extended receptivity, it is also possible that ewes, facing unavoidable forced matings by coursing rams, gain genotypic benefits for offspring by providing prolonged copulatory access to relatively dominant defending rams.     

Male preference for large females and assortative mating for body size in the sand lizard (Lacerta agilis)

Summary Male sand lizards (Lacerta agilis) are polygynous and guard individual females for several hours to days after copulation. Even though the copulation itself only lasts 2–4 min, the total time that a male invests per female is considerably more and may constitute a substantial investment during a mating season. In such situations, when male copulation frequency is constrained, or when variation in female fecundity is high, mate choice by males may be adaptive. Large body size in female sand lizards is correlated with higher fecundity. In choice experiments performed in the laboratory, male sand lizards preferred to court large females rather than small females. In addition, when there was little difference in size between the females in the experiment, the males visited the two females more often before they started to court the preferred female. The results from a field study during 1984 and 1987–1990 showed that females are non-aggressive, have small neighboring home ranges (c. 100 m²) and may share burrows and sites for thermoregulation. This means that females can be found close together and thus gives males the opportunity to choose a mate. Assortative mating with respect to size was observed in a natural population, as well as a limited number of direct choices of females by males. These results support the results of the choice experiment.     

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.     

Reproductive status influences group size and persistence of bonds in male plains zebra (<Emphasis Type="Italic">Equus burchelli</Emphasis>)

Animal groups arise from individuals’ choices about the number, characteristics, and identity of associates. Individuals make these choices to gain benefits from their associations. As the needs of an individual change with its phenotype, so too we expect the nature of its associations to vary. In this paper, we investigate how the social priorities of male plains zebra (Equus burchelli) depend on reproductive state. An adult male is either a bachelor, and lacking mating access, or a stallion defending a harem. Multiple harems and bachelor males aggregate in larger herds. Herds frequently split and merge, affording males opportunities to change associates. Over a 4-year period, we sampled the herd associations in a population of 500–700 zebras. To isolate the effects of reproductive state on male social behavior, we account for potential confounding factors: changes in population size, grouping tendencies, and sampling intensity. We develop a generally applicable permutation procedure, which allows us to test the null hypothesis that social behavior is independent of male status. Averaging over all individuals in the population, we find that a typical bachelor is found in herds containing significantly more adults, bachelors, and stallions than the herds of a typical stallion. Further, bachelors’ bonds with each other are more persistent over time than those among stallions. These results suggest that bachelors form cohesive cliques, in which we may expect cooperative behaviors to develop. Stallion–stallion associations are more diffuse, and less conducive to long-term cooperation. This contribution is part of the special issue “Social Networks: new perspectives” (Guest Editors: J. Krause, D. Lusseau and R. James).     

Condition-dependent control of paternity by female purple martins: implications for coloniality

Proposed causal links between extra-pair copulation (EPC) and colony formation in socially monogamous birds hinge on the question of which sex controls fertilizations. We examined in colonial purple martins Progne subis (1) whether EPCs were forced or accepted by females, and (2) the degree to which apparently receptive females were able to obtain EPCs against their mates’ paternity defenses. Paternity analyses of multilocus DNA fingerprinting confirmed previous findings of a marked relationship between age class and extra-pair fertilizations (EPFs), with young males losing paternity of 43% (n = 53) of their putative offspring compared to 4% (n = 85) by old males. All assignable extra-pair offspring were sired by old males, with one male obtaining most EPFs each year. Contrary to the hypothesis that EPCs are forced, EPF frequency within age class did not increase with seasonal increases in the number of males per fertile female. Whereas the male control hypothesis predicted that the male age class that mate-guarded more would be cuckolded less, the reverse was true: young males guarded significantly more intensely. The male age class difference in cuckoldry could not be explained by the possibility that young and inexperienced females (which are usually paired to young males) were more vulnerable to forced copulation because EPFs were unrelated to female age. These findings suggest that females (1) pair with old males and avoid EPCs, or (2) pursue a mixed mating strategy of pairing with young males and accepting EPCs from old males. The receptivity to EPCs by females paired to young males put them in conflict with their mates. Two factors determined the paternity achieved by young males: (1) the relative size of the male to the female, with young males achieving much higher paternity when they were larger than their mates, and (2) the intensity of mate-guarding. Both variables together explained 77% of the variance in paternity and are each aspects of male-female conflict. Given female receptivity to EPCs, mate-guarding can be viewed as male interference with female mating strategies. We conclude that EPCs are rarely or never forced, but the opportunity for females paired to young males to obtain EPCs is relative to the ability of their mates to prevent them from encountering other males. Evidence of mixed mating strategies by females, combined with other features of the martin mating system, is consistent with the female-driven “hidden lek hypothesis” of colony formation which predicts that males are drawn to colonies when females seek extra-pair copulations. Received: 23 March 1995/Accepted after revision: 14 January 1996     

Cloacal erection promotes vent apposition during forced copulation in the New Zealand stitchbird (hihi): implications for copulation efficiency in other species

Cloacal protuberances (CP) in male birds result from spermatic engorgement of storage tubules around the cloaca during the breeding season. We examined seasonal changes in the volume and orientation of the CP in the New Zealand stitchbird Notiomystis cincta. The male stitchbird has one of the largest recorded CPs for any species (max = 1,570 mm³), with CP volume increasing by almost 400% between the non-breeding and breeding seasons. While sperm competition has been positively correlated with the magnitude of CP storage in other species, no evidence previously existed for the CP improving copulation efficiency. By measuring the relative orientation of the CP throughout the year, we show that not only does the CP increase in size as males become sexually active, it also changes its orientation by approximately 60°. This results in it shifting from facing posteriorly to becoming almost perpendicular to the abdomen. This cloacal erection improves the apposition of the male and female cloacal openings during face-to-face forced copulation in this species. This provides the first reported evidence supporting the copulation efficiency hypothesis of the avian CP. While the magnitude of seasonal changes in female cloacal volume was similar to males, female cloacal orientation remained virtually unchanged across seasons. This difference between the sexes is likely to reflect differing selection pressures for optimizing sperm transfer. In females, a posterior-facing cloaca is ideal for both waste evacuation and sperm reception, whereas, for the male, a posterior-facing cloaca is well suited for waste evacuation, but possibly hinders sperm delivery. Changes in male cloacal orientation from the non-breeding to the breeding season are a likely reflection of conflict in this dual function. Evidence of changes in CP orientation in another passerine species suggests this phenomenon is widespread and also important for understanding related fields such as sperm competition, forced copulation and constraints on the evolution of the avian intromittent organ.     

Re-copulation and post-copulatory mate guarding increase immediate female reproductive output in the dragonfly Nannophya pygmaea Rambur

After copulation, male Nannophya pygmaea dragonflies mate guard by hovering over ovipositing females and repelling conspecific males. Copulation is not always a prerequisite for oviposition in the females of this species because females can store the sperm received during previous visits/copulations. An oviposition episode consists of several bouts of oviposition separated by periods of perching. We conducted two types of male-removal experiments to examine the effects of mating and post-copulatory mate guarding on the oviposition behaviour of females. In the first experiment, we removed all males from the habitat to eliminate the effect of re-copulation, mate-guarding and harassment by males. In the second experiment, we removed males immediately after copulation to eliminate the effects of guarding and other post-copulatory male-female interactions. We compared these experimental data with data obtained under natural conditions. The dipping rate in an oviposition bout was not influenced by copulation or guarding. However, guarded females made more dips per episode than did solitary females. The proportion of time actually spent ovipositing (total bout duration/oviposition episode duration) of guarded females was higher than that of solitary females. Solitary females often oviposited in more than one territorial site, while guarded females usually oviposited within a single territorial site during an oviposition episode. Because males tend to hold territories at sites where egg survival is high, guarded females (and the male guardian) benefit from guarding in terms of egg hatchability. The possible benefits for solitary females are discussed.