Female choice of large males in the treefrog Hyla chrysoscelis: the importance of identifying the scale of choice

Summary In this study, I detected the presence of female choice for larger males in the treefrog Hyla chrysoscelis. Mated males were larger than unmated males at the scales females could have assessed males: local groups (males found within 2 meters of a mated pair) and the nightly chorus. In addition, females were observed to initiate amplexus with the larger males in local groups. Mated males were larger than unmated males for two of the four seasons studied. Seasonal mating success was also analyzed with a multiple regression model that included size and chorus attendance (nights spent at the breeding site). The partial selection coefficients, which represent the relative magnitude of directional selection, were significant for size in only one season and for chorus attendance in all four seasons. Therefore, there are two possible advantages to being a large male: increased likelihood of obtaining a mate due to female preference, and increased likelihood of mating even if no female preference because of a greater number of opportunities to obtain a mating if matings occurred at random.     

Polygyny in the red-winged blackbird: do females prefer monogamy or polygamy?

Five hypotheses have been proposed to explain polygyny in the red-winged blackbird (Agelaius phoeniceus). We categorized the hypotheses into three groups based on female preference for unmated versus monogamously mated males: (1) the “polygyny threshold” model, “sexy son” hypothesis and the “asynchronous settlement” model, which assume that females prefer unmated males to mated males on breeding situations of homogeneous quality; (2) the “neutral mate choice” hypothesis, which assumes that females have no preference; and (3) the “cooperative female choice” model, which assumes that females prefer monogamously mated males to unmated males. We tested the direction of female preference in two field experiments. In both experiments, newly settling females were given a choice of two adjacent territories, one defended by an unmated male and the other by a monogamously mated male. Male mating status was randomized with respect to the variation in territory quality and male quality. Early in the breeding season, significantly more females settled with the unmated males than with the mated males. Although more females settled with the unmated males than with the mated males late in the breeding season, the difference was no longer significant. Female settlement late in the season appeared to be related to the tenure of the resident females: the new females avoided territories where the resident females were in early stages of their nesting, but settled on territories where the resident females were in late stages. The pattern of female settlement shows that females prefer unmated males to mated males. The preference is consistent with the polygyny threshold model, sexy son hypothesis and the asynchronous settlement model, and inconsistent with the neutral mate choice hypothesis and the cooperative female choice model. For this reason, the latter two hypotheses are unlikely to explain the occurrence of polygyny in our population of red-winged blackbirds. Received: 1 December 1994 / Accepted after revision: 28 October 1995     

Time constraints and multiple choice criteria in the sampling behaviour and mate choice of the fiddler crab,Uca annulipes

Active female sampling occurs in the fiddler crab Uca annulipes. Females sample the burrows of several males before remaining to mate in the burrow of the chosen partner. Females time larval release to coincide with the following nocturnal spring tide and must therefore leave sufficient time for embryonic development after mating. Here we show how this temporal constraint on search time affects female choosiness. We found that, at the start of the sampling period (when time constraints are minimal), females selectively sample the larger males in the population. Towards the end of the sampling period (when the temporal constraints increase the costs of sampling), females are less selective. Furthermore, we suggest that the number of males sampled (and other indices of ‘‘sampling effort’’) may not be reliable indicators of female choosiness and may not reflect the strength of female mating preferences under certain conditions. Burrow quality also emerged as an important criterion in final mate choice. Burrow structure potentially influences reproductive success, and mate acceptance based on burrow structure appears to involve a relatively invariant threshold criterion. Since there is no relationship between male size and burrow quality, females are using at least two independent criteria when choosing potential mates. We envisage mate choice as a two-stage process. First, females select which males to sample based on male size. They then decide whether or not to mate with a male based on burrow features. This sampling process explains how two unrelated variables can both predict male mating success. Received: 23 March 1995/Accepted after revision: 14 January 1996     

Display rate by male fairy-wrens (Malurus cyaneus) during the fertile period of females has little influence on extra-pair mate choice

Empirical and theoretical studies have only recently begun to examine how females use complex multi-component displays when selecting mates. Superb fairy-wrens are well suited to the study of female choice because females have control over extra-group paternity and cuckold their mates at high rates, while males possess a variety of sexually selected traits. Available evidence suggests that females base their extra-group mate choice on the timing of male moult into breeding plumage or the onset of display. However, males continue to perform elaborate displays throughout the season, and direct most displays to females during their fertile period. We therefore conducted focal observations on fertile females to quantify the frequency of male display and used microsatellite genotyping to compare the role of display rate during the breeding season and the timing of male moult on female mate choice. We show that the addition of data on male display rate does not improve our ability to predict which males obtain extra-group paternity. The timing of male moult into breeding plumage remains the only predictor of male extra-group reproductive success. Nevertheless, we found that males displayed more to females that were unable to select extra-group mates on the basis of the timing of moult or the onset of display. This raises the possibility that there are circumstances when females use display rate to discriminate between potential extra-group sires. Overall this study supports the theoretical prediction that females are more likely to base their mate choice on reliable indicators of male quality such as fixed morphological traits and displays of endurance, in this case an early moult into breeding plumage and the performance of an elaborate display during the winter, than a flexible behavioural trait such as display rate during the breeding season. Received: 26 January 2000 / Revised: 1 August 2000 / Accepted: 26 August 2000     

How reliable are behavioral cues for assessment of male mating status in polyterritorial wood warblers,Phylloscopus sibilatrix?

It has been suggested that polyterritoriality in birds, i.e. mated males defending two spatially separated territories, deceives females into accepting polygyny against their best interest. Mated males singing on secondary territories should make it difficult for new arriving females to assess the mating status of unmated and mated males advertising to females. However, there have been objections to the deception hypothesis, suggesting that differences in territorial behavior between males of different mating status may make correct assessment possible. Polyterritorial males frequently leave their secondary territory to visit the female in the primary territory, thereby revealing their mating status to females in search of mates. The aim of this study is to investigate how reliable behavioral cues are in assessing the mating status of males in polyterritorial wood warblers. Our study shows that singing activity of mated males on secondary territories and unmated males was similar in the morning, while there was a significant difference in the afternoon. Unmated males spent, on average, more time on their territories than mated males spent on their secondary territories. However, there was considerable variation in both unmated and mated males in the frequency in which they left their territories. For instance, unmated males left their territories quite frequently early in the season compared to later. Hence, females must be able to interpret singing behavior and a male leaving his territory differently depending on time of the season and time of the day in order to correctly assess the mating status of advertising males. Estimating the increase in probability of finding an unmated male by using behavioral cues showed that females could increase the probability of finding an unmated male compared to random choice with respect to mating status. The uncertainty was, however, still considerable. The most important factor affecting the probability of finding an unmated male was the proportion of mated males on secondary territories compared to unmated males. Our study suggests that presence on a territory is not a reliable cue for the assessment of male mating status in polyterritorial wood warblers.     

Why do female pied flycatchers mate with already mated males: deception or restricted mate sampling?

The polygyny threshold model suggests that females make an optimal choice between mated and unmated males. However; in birds in which males provide parental care, the fitness of secondary females is often lower than expected from this model. This has been explained by the deception hypothesis, which states that males hide their mating status and deceive females into polygyny. Yet there is no direct evidence that secondary females are unaware of male mating status when they settle. Alternatively, females settle with mated males as a result of mate competition and costs of searching. We used videofilming at nestboxes defended by males to study mate sampling of female pied flycatchers Ficedula hypoleuca. The females visited on average only 2.74 males (range 1–8, n = 43). Most (16 of 19) of the polygynous matings occurred because females had only visited mated males, or the unmated males visited became occupied by competitors during the sampling period. Among females that could choose between both mated and unmated males, the majority (13 of 16) settled with unmated males. These results lend little support to the deception hypothesis but are consistent with the view that females are able to detect male mating status but sometimes settle with mated males because of cost of searching. Prospecting females seemed willing and able to suffer the cost of fighting with aggressive primary females in the males' secondary territory if no alternative mating options were available. In addition to male mating status, females took male quality (plumage colour, age) into account in mate choice but the former appeared to be the more important. Correspondence to: T. Slagsvold     

The role of sexual imprinting and the Westermarck effect in mate choice in humans

Positive sexual imprinting is a process by which individuals use the phenotype of their opposite-sex parent as a template for choosing mates and is suggested to play an important role in human mate choice. In contrast, negative imprinting, or ??The Westermarck Effect??, is characterized by individuals developing a strong sexual aversion to others with whom they lived closely in infancy and early childhood. In this review, we evaluate the literature on their effects on mate choice in humans. We find little evidence to support positive imprinting in humans because the studies either have serious design flaws, do not exclude effects of heritable mating preferences, or do not account for several possible alternative explanations. Instead, it seems that the opposite phenomenon, negative sexual imprinting, has some support from natural experiments which have found that individuals avoid mating with those with whom they lived closely in infancy and early childhood. However, it seems that early association does not produce a strong-enough aversion to completely annihilate sexual desire, probably because the mind uses multiple kinship cues to regulate inbreeding avoidance. Thus, it appears that the evidence for both types of imprinting is fairly weak in humans. Thus, more studies are needed to test the role of sexual imprinting on mate choice in humans, especially those measuring interactions between positive and negative imprinting.     

What do pairing patterns in common tern, Sterna hirundo, recruits reveal about the significance of sex and breeding experience?

In general, reproductive output in long-lived bird species increases in older compared to younger individuals. Therefore, experienced mates should be advantageous for first-time breeders. To examine requirements and consequences of experienced pair mates we investigated the first pair bonds of common terns, Sterna hirundo, recruiting to their natal colony. We found that male recruits were usually the same age as their mates, whereas female recruits were usually the younger member of the pair. In order to acquire experienced mates, it was necessary for males to arrive early in the year of recruitment. Mates with 2 or more years of breeding experience were only attainable by male recruits characterised by greater body mass and age. Female recruits arrived more than 1 week later than their experienced mates and significantly later in the season than male recruits paired with experienced females. In general, females first bred at a younger age than males, and neither the female recruits body mass nor their age was related to the level of experience of their first mate. These sex-specific differences in obtaining an experienced mate did not result in different levels of reproductive success between the sexes. Male and female recruits with mates with 2 or more years breeding experience benefited from having experienced mates: they had greater reproductive success. First-time breeders paired with mates with only 1 year of breeding experience did not differ from pairs where both members were breeding for the first time in terms of reproductive success, but clutches were larger. Our results illustrate not only different prerequisites for males and females, but also males need for experienced mates. Delayed male breeding (postponing breeding for another year), supposed to be a negative trait, and high body mass, supposed to be a trait of superior individual quality, can be combined in some individuals, improving reproductive success and showing that breeding common terns use a range of tactics to begin reproduction.Communicated by F. Trillmich     

Breeding synchrony and extra-pair mating in red-winged blackbirds

Using data from a 6-year paternity study of red-winged blackbirds, I tested the hypotheses that increased nesting synchrony should either promote extra-pair mating by increasing the advantage of extra-pair mating to females, or decrease extra-pair mating by constraining males from seeking extra-pair copulations. Contrary to these hypotheses, the occurrence of extra-pair paternity did not vary with nesting synchrony over the breeding season, or vary with the number of synchronous nests within territories or within marshes, or with nesting order on territories. However, for nearly all nests with extra-pair young, there were fewer females synchronous with that nest on the cuckolder's territory than on the territory of the cuckolded male. This “advantage” of a synchrony difference was less pronounced for older males that cuckolded younger males, particularly when the two males were not neighbors. Collectively, these results suggest that breeding synchrony affects extra-pair mating by affecting mate guarding, but that breeding synchrony alone can not be used to predict which females are more likely to engage in extra-pair mating, nor with which extra-pair males they will mate. Understanding why extra-pair mating by older males is less affected by breeding synchrony may explain much about both the proximate and ultimate causes of extra-pair mating in red-winged blackbirds. Received: 7 June 1996 / Accepted after revision: 25 November 1996     

Badge size in the house sparrow Passer domesticus

Summary I studied female mate choice in house sparrows Passer domesticus in relation to the size of the black throat patch of males (badge size), which is a signal of dominance status, and to territory quality. Males with large badges obtained a mate earlier in spring than males with small badges. Males which remained unmated during the entire breeding season had smaller badges than mated males, even when controlling for the effect of other morphological traits and age. Largebadged males had territories with more breeding sites than small-badged males. Territories defended by males with large badges primarily had hole nest sites, which were safe from predators, and nestlings rarely fell from hole nests. Females were implanted with estradiol to induce copulation solicitation displays. Females responded more strongly to a male taxidermic mount and song than to song alone, and they responded more strongly and frequently to male mounts with large badges. Female house sparrows chose mates on the basis of male badges and perhaps on the basis of the quality of the nesting territories offered.     

Environmental,social, morphological,and behavioral constraints on opportunistic multiple paternity

Multiple mating and multiple paternity in polytocous species have been mostly studied from an adaptive (i.e., cost–benefit) perspective. Disease, time, energy, and the risk of injuries are well-known costs of multiple mating, yet from both male and female perspectives, a number of genetic and non-genetic benefits have also been identified. The effects of environmental conditions and individual-specific behavior, however, are much less well understood. Using a long-term study on yellow-bellied marmots (Marmota flaviventris), we evaluated the impacts of environmental variation, social structure, female body mass, and female docility (a personality trait) on the occurrence of multiple paternity. Multiple paternity was influenced by environmental constraints, social constraints, a female’s personality, and her body mass at emergence from hibernation. Personality and mass effects were detected only when environmental or social conditions were favorable. Our results suggest that multiple paternity is mainly limited by the opportunity to have access to multiple mates and is influenced by costs or mate choice because heavier females were more likely to have litters with multiple sires than smaller ones. Future studies in other species might benefit from considering environmental constraints when studying multiple paternity.     

Mate choice in the brown thornbill (<Emphasis Type="Italic">Acanthiza pusilla</Emphasis>): are settlement decisions,divorce and extrapair mating complementary strategies?

In socially monogamous birds, females may express mate preferences when they first select a breeding partner, through divorce and subsequent breeding dispersal to a new partner and through extrapair mating. We examined settlement patterns, divorce and breeding dispersal in a sedentary Australian passerine, the brown thornbill (Acanthiza pusilla), in relation to two traits known to influence extrapair paternity (male age and male size). Settlement decisions, divorce and territory switching behaviour were all female strategies that reduced their likelihood of breeding with 1-year-old males. Females preferred to settle in territories with 2+ -year-old males, were more likely to divorce 1-year-old males, and only switched territories if they had an opportunity to form a new pair bond with an old male. In contrast, female settlement and divorce decisions were not influenced by male size. Female thornbills obtain a direct benefit from preferring older males as social mates because breeding success improves with male age in brown thornbills. Nevertheless, divorce rates in this species were low (14% of pair bonds were terminated by divorce), and individuals rarely switched territories following the death of a mate. Both of these mating strategies appeared to be primarily constrained by the distance adults moved to initiate a new pair bond (1–2 territories) and by the limited availability of unpaired older males in the immediate neighbourhood.Communicated by M. Webster     

Sexual selection and the function of a melanin-based plumage ornament in polygamous penduline tits <Emphasis Type="Italic">Remiz pendulinus</Emphasis>

Melanin-based ornaments are often involved in signaling aggression and dominance, and their role in sexual selection is increasingly recognized. We investigated the functions of a melanin-based plumage ornament (facial ‘mask’) in male Eurasian penduline tits Remiz pendulinus in the contexts of male–male aggression, mating success, and parental care. The penduline tit is a passerine bird with a unique mating system in which both sexes may mate with several mates in a breeding season, and one (or both) parent deserts the clutch. Our study revealed that mask size of males is more likely an honest signal used by females in their mate choice decisions than a trait involved in male–male competition. First, mask size increased with both age and body condition, indicating that the mask may signal male quality. Second, males with larger masks paired more quickly and had more mates over the breeding season than males with smaller masks. Third, we found no evidence that male mask size signals male–male aggression or dominance during competitive encounters. The increased mating success of large-masked males, however, did not translate into higher reproductive success, as nestling survival decreased with mask size. Therefore, we conclude that there is either no directional selection on male mask size or males with larger masks receive indirect, long-term benefits.     

Long-term mate and territory fidelity in neotropical buff-breasted wrens (Thryothorus leucotis)

Among sedentary species, mate and site fidelity may be imposed by year-round territoriality and high annual survivorship, both of which theoretically lead to few opportunities to move between mates and breeding territories. We examined mate and territory fidelity in sedentary neotropical buff-breasted wrens (Thryothorus leucotis) in relation to the opportunities to divorce (vacancies on neighbouring territories), breeding experience and site of origin (within vs outside local breeding population). Overall, buff-breasted wrens showed high rates of mate and territory fidelity between years, with individuals significantly more faithful to their territories than to their partners. Mate and territory fidelity was not forced upon buff-breasted wrens, as most individuals had opportunities to switch mates and territories. However, this was not uniform among all individuals. Individuals that had bred together displayed 100% mate and territory fidelity despite the fact that males and females in almost all experienced pairs had opportunities to switch mates and territories. Local recruits (philopatric offspring that acquired territories in the local breeding population) also displayed mate and territory fidelity, but fidelity occurred because recruits rarely had the opportunity to divorce prior to initiating their first breeding attempt. By contrast, almost half of all territorial individuals that originated outside the local breeding population (immigrants) had opportunity to divorce, with approximately two thirds of them switching mates and territories. Divorces occurred within the first 5 months of pairing, suggesting that immigrants may have sampled mates and territories before forming permanent partnerships.     

Female reactions to male absence after pairing in the pied flycatcher

Mating with an already mated pied flycatcher (Ficedula hypoleuca) male is costly for a female. Two hypotheses explain why some females still mate with already mated males. The deception hypothesis suggests that some females mate with already mated males since it is difficult to assess perfectly the mating status of a male with separate territories (polyterritoriality). The search cost hypothesis states that females are aware of male mating status but the costs of searching for an unmated male exceed costs associated with the status of secondary female. One potential cue that could disclose a male's mating status is the existence of brief visits to the primary territory by polyterritorial males. To mimic such visits I removed the male from the territory for 60 min soon after female settlement. Only few females abandoned their mates as a consequence of male removal even though they had available unmated males close by. This result is most consistent with the deception hypothesis. Females did not use occasional male absence as a cue to avoid presumably polyterritorial males and in this respect they were not perfect in assessing males.     

Supply and demand: causes and consequences of assortative mating in common terns <Emphasis Type="Italic">Sterna hirundo</Emphasis>

Although assortative mating is widespread among long-lived monogamous birds, the underlying mechanisms still remain unclear in many species. In this study, we analysed assortative mating by age and body mass in the common tern Sterna hirundo with special regard to mate choice decisions of newly paired birds. To assess whether assortative mating by age is an active decision involving either homotypic or directional preferences or rather a passive process due to restricted availability of potential mates, we analysed the influence of age-dependent arrival date and cohort size. Furthermore, we looked for direct benefits in terms of reproductive success. Common terns mated assortatively by age and arrival date but not by body mass at arrival. Assortative mating by age was age dependent and was detected mainly in younger birds, whereas birds older than 8 years rather re-paired with younger ones. The availability of same-aged mates was restricted by cohort size and arrival date. We found no general evidence for better reproductive performance of same-aged pairs. Instead, relative reproductive success was related to own age and the relative age of mate: It should be advantageous for any bird to acquire an old mate; hence, assortative mating by age seems to be beneficial only for old terns but not for young ones. Age-assortative mating in common terns occurred by both passive and active processes, which are not mutually exclusive. Our results do not indicate a homotypic but a directional preference and support the theory that high-quality (older/experienced) individuals tend to mate assortatively if same-aged mates are available, which leaves low-quality (young/inexperienced) individuals to mate among themselves.     

Does female aggression prevent polygyny? An experiment with pied flycatchers (Ficedula hypoleuca)

The female aggression hypothesis states that resident females may be able to prevent polygyny by behaving aggressively towards intruding females. A critical test of the hypothesis is to provide prospecting females with a choice between displaying mated males some of which have initial mates with artificially reduced levels of aggressiveness. Here we present a mate choice experiment on pied flycatchers Ficedula hypoleuca. The species is a cavity nester, and resident females were prevented from behaving aggressively by enclosing them within their own nestboxes: narrowing the entrance hole so that they could not escape but could still let their head out and have some contact with their mate. This treament had only a minor influence on male behaviour. We studied whether the experimental males were better able to attract a new female than a control group of mated males. Four predictions from the female aggression hypothesis were supported. (1) Mating success of control males was positively related to the distance between their primary and secondary territory. (2) For experimental males, mating success was unrelated to interterritorial distance. (3) Experimental males had higher mating success than control males when the interterritorial distance was short but (4) not when it was long. Experimental males had much lower mating success than unmated males, as would be expected if prospecting females are able to discover male mating status from cues other than visits by primary females to their mates' secondary nest sites. Received: 5 January 1998 / Accepted after revision: 30 December 1998     

Mate choice in the face of both inbreeding and outbreeding depression in the intertidal copepod Tigriopus californicus

 In species vulnerable to both inbreeding and outbreeding depression, individuals might be expected to choose mates at intermediate levels of genetic relatedness. Previous work on the intertidal copepod Tigriopus californicus has repeatedly shown that crosses between populations result in either no effect or hybrid vigor in the first generation, and hybrid breakdown in the second generation. Previous work also shows that mating between full siblings results in inbreeding depression. The present study again found inbreeding depression, with full sibling mating causing significant fitness declines in two of the three populations assayed. In the mate choice assays, a single female was combined with two males. Despite the costs of both inbreeding and outbreeding, mate choice showed clear inbreeding avoidance but no clear pattern of outbreeding avoidance. This lack of outbreeding avoidance may be attributed either to the temporary increase in fitness in the F₁ generation or to the absence of selection for premating isolation in wholly allopatric populations with infrequent migration. If this inability to avoid unwise matings is common to other taxa, it may contribute to the problem of outbreeding depression when allopatric populations are mixed together. Received: 18 May 1999 / Accepted: 25 January 2000     

Operational sex ratios and behavioural sex differences in a pipefish population

In the pipefish Syngnathus typhle, only males brood embryos in specially developed brood pouches, supplying oxygen and nutrients. Laboratory studies have shown that this elaborate paternal care has led to sex-role reversal in this species: males limit female reproductive rate, females are the primary competitors for mates and males exercise greater selectivity in accepting mates. In the first field study of this pipefish, we describe mating behaviour in the wild and test the hypothesis that temporal variations in the operational sex ratio (OSR) determine sex differences in mating behaviour. Our study comprised two reproductive seasons of two sequential mating periods each, the latter separated by a lengthy interval of male brooding. During mating periods, females displayed to all males without wandering and males moved about searching for females, without reacting to all females. The OSR was least female-biased (or even male-biased) at the onset of the breeding season, when most pipefish were simultaneously available to mate, but became strikingly female-biased as males' pouches were filled. The OSR remained substantially female-biased during the second mating period, because few males became available to remate at any one time. As hypothesised, female-biased OSRs resulted in more female-female meetings. As well, females were above the eelgrass more often than brooding males, thus exposing themselves to conspecifics and/ or predators. In the second year, males arrived earlier than females on the breeding site and male pregnancies were shorter, because of higher water temperatures, so rematings occurred earlier. Males met more often during that year than the previous one, but male competitive interactions were still not observed. The field results support laboratory studies and demonstrate that behaviours associated with female-female competition are more prominent when the OSR is more female-biased. Correspondence to: A. Vincent     

The costs of male parental care and its evolution in a neotropical frog

Summary Parental care is practiced exclusively by males of the Puerto Rican frog, Eleutherodactylus coqui. Males brood clutches of direct-developing eggs in non-aquatic nest sites and defend eggs against cannibalistic nest intruders. Here, I report on energetic and mating costs incurred by males that provide parental care, and suggest how these proximate costs affect male fitness and the evolution of male parental care in this species. Energetic costs are small for brooding males in comparison to non-brooding, calling males. Brooding males had a higher frequency of empty stomachs and lost small, but significant, fractions of their initial body mass during parental care. Abdominal fat bodies of brooding males during the middle third of parental care were significantly smaller than those of calling males; those of males brooding eggs in earlier or later stages were not different. The mating cost of parental care is greater. Most brooding males cease calling during parental care. However, gravid females are available (i.e., known to mate) on most nights during the principal breeding season; hence non-calling males miss potential opportunities to mate. A mating cost was estimated by calculating nightly mating probabilities for calling males in a plot where nightly calling male densities and daily oviposition schedules were known. On average, a male exhibiting normal calling behavior would be expected to obtain a new mate once every 35.7 days. Hence a brooding male that ceased calling for a 20-day parental care period would miss, on average, 0.56 additional mates. Males that were more successful than average in attracting mates could miss up to 1.63 matings. A marginal value model (Fig. 1) is used to analyze the net effect on male fitness of parental care benefits and costs in E. coqui (Fig. 3). The model indicates that males garner the highest reproductive success by providing care from oviposition through hatching. There is no stage during the pre-hatching period at which a desertion strategy would yield higher reproductive success. In fact, the model suggests that males should provide full parental care even in the face of much higher mating costs than currently obtain in the system.