Reproductive Trade-Offs and Direct Costs for Males in Arthropods

Until 30 years ago, the emphasis on reproductive costs for males was mainly on costs related to mate searching, courtship and fighting with rival males. However, costs for males are substantial and varied and often resemble the more thoroughly studied female reproductive costs. Costs can be referred to as trade-off costs, where investment in reproductive activity comes at the expense of another important activity or fitness component. Investment in reproduction at the expense of longevity and future reproduction is the ultimate cost, because it affects fitness directly. In contrast, flawed performance (e.g., of the immune system) is perceived as a mechanistic trade-off, because it affects fitness indirectly through a mediator (i.e., parasites). Finally, direct costs refer to direct measurements of the energy expenditure during involvement in reproduction-related activities. Both direct and mechanistic trade-off costs often result in decreased longevity compared to unmated males (an ultimate cost). Males incur costs during different reproductive phases: before copulation, when producing sperm, while searching for, courting and copulating with females, and subsequently when guarding females or taking care of offspring. This synthesis follows previous pioneering reviews addressing specific aspects of male costs, but strives to summarize all known male reproductive cost types more comprehensively, including their classification. We suggest several directions for targeted future research. While costs for males have been fairly well described, it is now necessary to uncover the ecological and evolutionary factors responsible for differences between closely related species and systems and to better link between directly-measured costs, mechanistic trade-off costs and ultimate trade-off costs.     

Sex-biased terminal investment in offspring induced by maternal immune challenge in the house wren (Troglodytes aedon)

The reproductive costs associated with the upregulation of immunity have been well-documented and constitute a fundamental trade-off between reproduction and self-maintenance. However, recent experimental work suggests that parents may increase their reproductive effort following immunostimulation as a form of terminal parental investment as prospects for future reproduction decline. We tested the trade-off and terminal investment hypotheses in a wild population of house wrens (Troglodytes aedon) by challenging the immune system of breeding females with lipopolysaccharide, a potent but non-lethal antigen. Immunized females showed no evidence of reproductive costs; instead, they produced offspring of higher phenotypic quality, but in a sex-specific manner. Relative to control offspring, sons of immunized females had increased body mass and their sisters exhibited higher cutaneous immune responsiveness to phytohaemagglutinin injection, constituting an adaptive strategy of sex-biased allocation by immune-challenged females to enhance the reproductive value of their offspring. Thus, our results are consistent with the terminal investment hypothesis, and suggest that maternal immunization can induce pronounced transgenerational effects on offspring phenotypes.     

Parental investment of male two-spotted goby, Gobiusculus flavescens (Fabricius)

The trade-off between parental care and feeding was studied in the male two-spotted goby (Gobiusculus flavescens F.). Two temperatures, 8.5 degrees C and 13.0 degrees C, were used, with five replicates at each temperature, in order to determine whether temperature influenced parental behaviour. In each replicate, two males and four females were introduced to an aquarium, where the males chose between two nests and courted the females. In each replicate, one male spawned. After spawning, the males guarded the eggs until hatching. The guarding males' behaviour was recorded with a video camera twice a day (15 min each time), once before and once after they were fed. The male's condition (c-factor) was calculated at the start of the experiment and after egg hatching. The eggs were spawned in an artificial nest (half of a PVC tube), and attached to the nest in a single layer. The areas with eggs (representing brood size) were marked after spawning and the fry counted after hatching (which was used to calculate area hatched). Numbers of prey eaten (plankton) and number of aggressive encounters between the guarding male and the other fishes were recorded. Time spent in the nest and time used on fertilisation, fanning and cleaning were estimated and related to egg age, brood size, hatching success, temperature and food availability (no food or food).The results showed that feeding (expected to influence future reproduction) decreased and parental expenditure (current reproduction) increased, as the eggs developed (became closer to independence). Parental expenditure was significantly higher at 13.0 degrees C than at 8.5 degrees C, presumably due to higher oxygen demands by the eggs, and a greater risk of egg-infections. The c-factor of the males guarding eggs decreased over time, in contrast to the non-guarding males' c-factor. Guarding males' aggressiveness decreased as the eggs got older, but increased just before hatching. A possible explanation for this could be the decreasing intrusion by the non-guarding male and females caused by high aggressive behaviour by the guarding male early in the brood cycle. The exploitation of the nest (percentage of total nest area covered by eggs) seemed to determine the amount of parental expenditure and loss of condition, while brood size (area of eggs) had no effect.     

Nest-building activity and laying date influence female reproductive investment in magpies: an experimental study

Nest size or nest-building activity has recently been hypothesized to be a postmating sexually selected signal in monogamous birds: females may assess a male's parental quality and willingness to invest in reproduction by his participation in nest building. Females may thus adjust their reproductive effort (i.e. clutch size) not only to their own abilities but also to those of their mates. We investigated whether female magpies, Pica pica, use nest-building activity rather than nest size to adjust their reproductive effort during replacement breeding attempts. After we removed their first clutch, high-quality pairs that built a large nest for the first clutch were more capable of building a replacement nest and females adjusted their clutch size in relation to the time it took to build the nest rather than nest size. We also found support for the hypothesized trade-off between clutch size and egg size in magpies. In replacement clutches females decreased clutch size and increased egg volume, thereby probably improving the survival probability of their offspring in less favourable conditions.Copyright 2002 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour     

Is Mate Provisioning Predicted by Ornamentation? A Test with Northern Cardinals (Cardinalis cardinalis)

Male birds of many species feed their mates during courtship and incubation. The amount of food provided can be substantial and even essential for successful reproduction in some species, and can influence female nest attentiveness in many others. Additionally, mate provisioning may predict later nestling feeding rates. Females may thus benefit from being able to determine male provisioning effort. We assessed the expression of several ornaments, known to indicate condition in male northern cardinals (Cardinalis cardinalis), and compared these with mate provisioning rates, nestling feeding rates, and nest attentiveness. We found that male ornamentation may not be indicative of mate provisioning rates. Mate provisioning rate did not co‐vary with reproductive success, male feedings to nestlings, or nest attentiveness of females. However, females which were fed more often during incubation tended to provision nestlings less. Reduced female parental effort following extensive incubation feeding may be indicative of females using incubation feeding to assess future male parental effort. Male hormonal condition that favors high rates of nestling provisioning may be a proximate cause of mate provisioning during incubation, even in the absence of selection, favoring high rates of mate provisioning. Both sexes may have capitalized on this unselected behavior.     

Costs of reproduction and terminal investment by females in a semelparous marsupial

Evolutionary explanations for life history diversity are based on the idea of costs of reproduction, particularly on the concept of a trade-off between age-specific reproduction and parental survival, and between expenditure on current and future offspring. Such trade-offs are often difficult to detect in population studies of wild mammals. Terminal investment theory predicts that reproductive effort by older parents should increase, because individual offspring become more valuable to parents as the conflict between current versus potential future offspring declines with age. In order to demonstrate this phenomenon in females, there must be an increase in maternal expenditure on offspring with age, imposing a fitness cost on the mother. Clear evidence of both the expenditure and fitness cost components has rarely been found. In this study, we quantify costs of reproduction throughout the lifespan of female antechinuses. Antechinuses are nocturnal, insectivorous, forest-dwelling small (20-40 g) marsupials, which nest in tree hollows. They have a single synchronized mating season of around three weeks, which occurs on predictable dates each year in a population. Females produce only one litter per year. Unlike almost all other mammals, all males, and in the smaller species, most females are semelparous. We show that increased allocation to current reproduction reduces maternal survival, and that offspring growth and survival in the first breeding season is traded-off with performance of the second litter in iteroparous females. In iteroparous females, increased allocation to second litters is associated with severe weight loss in late lactation and post-lactation death of mothers, but increased offspring growth in late lactation and survival to weaning. These findings are consistent with terminal investment. Iteroparity did not increase lifetime reproductive success, indicating that terminal investment in the first breeding season at the expense of maternal survival (i.e. semelparity) is likely to be advantageous for females.     

Major benefits of guarding behavior in subsocial bees: implications for social evolution

Parental care is a behavior that increases the growth and survival of offspring, often at a cost to the parents' own survival and/or future reproduction. In this study, we focused on nest guarding, which is one of the most important types of extended parental care; we studied this behavior in two solitary bee species of the genus Ceratina with social ancestors. We performed the experiment of removing the laying female, who usually guards the nest after completing its provisioning, to test the effects of nest guarding on the offspring survival and nest fate. By dissecting natural nests, we found that Ceratina cucurbitina females always guarded their offspring until the offspring reached adulthood. In addition, the females of this species were able to crawl across the nest partitions and inspect the offspring in the brood cells. In contrast, several Ceratina chalybea females guarded their nests until the offspring reached adulthood, but others closed the nest entrance with a plug and deserted the nest. Nests with a low number of provisioned cells were more likely to be plugged and abandoned than nests with a higher number of cells. The female removal experiment had a significantly negative effect on offspring survival in both species. These nests frequently failed due to the attacks of natural enemies (e.g., ants, chalcidoid wasps, and other competing Ceratina bees). Increased offspring survival is the most important benefit of the guarding strategy. The abandonment of a potentially unsuccessful brood might constitute a benefit of the nest plugging behavior. The facultative nest desertion strategy is a derived behavior in the studied bees and constitutes an example of an evolutionary reduction in the extent of parental care.     

Incubation feeding by male Scarlet Tanagers: a mate removal experiment

ABSTRACT Incubation feeding, where males feed their mates, is a common behavior in birds and may improve female condition, nest attentiveness, and nesting success. We used behavioral observations and a temporary mate removal experiment to test the female nutrition hypothesis for incubation feeding by male Scarlet Tanagers (Piranga olivacea). All males (N= 20) were observed incubation feeding and fed females both at the nest (x? 1.36 trips/h) and away from the nest (x? 20.1 trips/h). Male feeding rate off‐nest was negatively correlated with the duration of female foraging bouts and positively correlated with the total time females spent incubating per hour. Eggs were predated at seven of 19 (37%) nests, but nest survival during incubation was not related to either female incubation behavior or male feeding rate. During temporary removal experiments (N= 12), female Scarlet Tanagers remained on the nest significantly longer and did not have longer foraging bouts. An unexpected outcome of the removal experiments was a dramatic change in female vocal behavior. All 12 experimental females gave chik‐burr calls during the male‐removal experiments (x? bout length = 11.7 min), but during normal observation periods only six of 20 females at the incubation stage gave chik‐burr calls (x? bout length = 0.7 min, N= 20). Our results suggest that female tanagers likely gain nutritional benefits from incubation feeding, but male feeding may not improve immediate reproductive success. Nine of 54 (17%) nestlings in five of 17 broods (29%) were extra‐pair young (EPY), indicating that males could potentially benefit from incubation feeding via mate retention and fidelity as well as, or instead of, through immediate gains in reproductive success. Our study indicates that females benefit from incubation feeding and do not simply passively accept food from their mates, but instead may influence male feeding rates through direct (e.g., mate following and vocalizing) and indirect (the threat of mate abandonment or cuckoldry) means.     

Parental effort of kestrels (Falco tinnunculus) in nest defense: effects of laying time, brood size, and varying survival prospects of offspring

We studied the nest defense behavior of Eurasian kestrels (Falcotinnunculus) towards a stuffed pine marten (Martes martes) througha 3-year vole cycle (1990–92) in western Finland. Survivalprobability of offspring decreases with a later start of breeding,and, therefore, early breeders should protect their offspringmore than late ones. We found this true for males during theincubation period, but not for females. In addition, we expectedthe nest defense intensity to increase with offspring number.During the incubation period, this was true for females, butnot for males. During the nestling phase, parents did not adjusttheir defense effort to natural or manipulated (by one to twoyoung) brood size. Survival prospects of kestrel offspring werehighest in the increasing vole year 1991 and lowest in the decreasingvole year 1992, and, therefore, we expected the defense activityof kestrels to follow the same trend. However, the oppositeresult appeared true for females with a similar tendency formales. Most hypotheses predicting avian nest defense behaviorwere not supported by our data. Temporally heterogeneous environmentand low degree of nest-site tenacity of migratory kestrels maymake them unfamiliar with environmental variation and survivalprospects of their offspring. Therefore, fitness benefits ofparental care are not predictable, and kestrels may thus adjusttheir parental effort to their own future reproductive potential(i.e., number of future breeding attempts), rather than to somecurrent investment indicator, like offspring age and number.     

What makes a nest-building male successful? Male behavior and female care in penduline tits

Why do females increase parental effort when caring for theoffspring of attractive males? First, attractive males may bepoor fathers so that their females are compelled to increasetheir own contribution in order to fledge some young (the partner-compensationhypothesis). Second, females mated to attractive males may bewilling to increase their parental effort to reap high indirectbenefits for their offspring, and in turn males can decreasetheir own contribution (the differential allocation hypothesis[DAH]). We investigated these hypotheses in the penduline titRemiz pendulinus, a small passerine bird that has sequentialpolygamy by both sexes and strict uniparental care either bythe male or the female. We focused on two sexually selectedmale traits: nest size and nest-building behavior. We show thatmale care is unrelated to nest-building behavior, whereas femalesare more likely to care for the offspring of those males thatspend more time nest building. Females also more likely carefor the offspring of males that build large nests. Consequently,the reproductive success of males increases with nest size andnest-building behavior. Our results are consistent with theDAH and suggest that nest-building behavior and nest size areunder postmating sexual selection in penduline tits.     

Temporal and spatial complexity of maternal thermoregulation in tropical pythons

Parental care is a widespread adaptation that evolved independently in a broad range of taxa. Although the dynamics by which two parents meet the developmental needs of offspring are well studied in birds, we lack understanding about the temporal and spatial complexity of parental care in taxa exhibiting female-only care, the predominant mode of parental care. Thus, we examined the behavioral and physiological mechanisms by which female water pythons Liasis fuscus meet a widespread developmental need (thermoregulation) in a natural setting. Although female L. fuscus were not facultatively thermogenic, they did use behaviors on multiple spatial scales (e.g., shifts in egg-brooding postures and surface activity patterns) to balance the thermal needs of their offspring throughout reproduction (gravidity and egg brooding). Maternal behaviors in L. fuscus varied by stage within reproduction and were mediated by interindividual variation in body size and fecundity. Female pythons with relatively larger clutch sizes were cooler during egg brooding, suggesting a trade-off between reproductive quantity (size of clutch) and quality (developmental temperature). In nature, caregiving parents of all taxa must navigate both extrinsic factors (temporal and spatial complexity) and intrinsic factors (body size and fecundity) to meet the needs of their offspring. Our study used a comprehensive approach that can be used as a general template for future research examining the dynamics by which parents meet other developmental needs (e.g., predation risk or energy balance).     

Evolution of passerine incubation behavior: influence of food, temperature, and nest predation

Abstract.— Incubation behavior is one component of reproductive effort and thus influences the evolution of life-history strategies. We examined the relative importance of body mass, frequency of mate feeding, food, nest predation, and ambient temperature to explain interspecific variation in incubation behavior (nest attentiveness, on- and off-bout durations, and nest trips per hour) using comparative analyses for North American passerines in which only females incubate. Body mass and frequency of mate feeding explained little variation in incubation behavior. We were also unable to detect any influence of food; diet and foraging strategy explained little interspecific variation in incubation behavior. However, the typical temperature encountered during reproduction explained significant variation in incubation behavior: Species breeding in colder environments take shorter bouts off the nest, which prevents eggs from cooling to temperatures below the physiological zero temperature. These species must compensate for shorter off-bouts by taking more of them (thus shorter on-bouts) to obtain needed energy for incubation. Nest predation also explains significant variation in incubation behavior among passerines: Species that endure high nest predation have evolved an incubation strategy (long on- and off-bouts) that minimizes activity that could attract predators. Nest substrate explained additional variation in incubation behavior (cavity-nesting birds have shorter on-bouts and make more frequent nest trips), presumably because nest predation and/or temperature varies among nest substrates. Thus, nest predation can influence reproductive effort in a way previously not demonstrated–by placing a constraint on parental activity at the nest. Incubating birds face an ecological cost associated with reproductive effort (predation of entire brood) that should be considered in future attempts to explain avian life-history evolution.     

Natural variation in stress response is related to post-stress parental effort in male house sparrows

The central life-history trade-off between current and future reproductive effort seems to be mediated by corticosterone in birds. However, still little is known about how naturally occurring corticosterone levels during an acute stress may influence subsequent parental behavior. In this study we observed the parental behavior of free-living male house sparrows (Passer domesticus) both before and after they were subjected to a standard capture–handling stress. We investigated the relationships between corticosterone levels, pre- and post-stress parental behavior, while we statistically controlled for a number of other variables using a multivariate regression method, the path analysis. We found that males' baseline feeding rate predicted the body mass of the nestlings, indicating that male parental care is directly linked to fitness. Corticosterone levels were not explained by baseline feeding rate, but both baseline and stress-induced corticosterone levels had a negative influence on the males' post-stress feeding behavior. Moreover, males with large bib size had a stronger stress response and lower post-stress feeding rate than small bibbed males. These results indicate that naturally occurring variation in baseline and stress-induced corticosterone levels may influence subsequent parental decisions: individuals mounting a robust stress response are likely to reduce their parental commitment. Parental effort may be regulated in a complex manner, with corticosterone mediating the life-history trade-off between current reproduction and survival. However, different resolutions of this trade-off were apparent only following the stress, therefore the ability to modulate the stress response and maintain parental care in stressful situations may be important in life-history evolution.     

Reduced immunocompetence and cost of reproduction in common eiders

Immunocompetence may be especially important in long-lived species where infectious organisms may have detrimental effects upon future reproductive value of hosts. The resource demand for immunocompetence may reduce resource availability for reproduction and a trade-off between these traits has therefore been proposed. Capital breeders, such as the common eider (Somateria mollissima), rely upon accumulated body reserves during reproduction. Eiders lose more than 40% of pre-breeding body mass during egglaying and incubation and many females abandon their ducklings to other females after hatching. Results from our observational study show that levels of leukocytes (i.e., lymphocytes, heterophils and heterophil/lymphocyte ratio) are not related to body mass early in the incubation period. However, eider females with low initial body mass showed signs of immunosuppression (i.e., decreased late levels of lymphocytes) and increased response towards stressors (i.e., increased heterophil/lymphocyte ratio) later in the incubation period. Moreover, females with low lymphocyte levels more frequently abandoned their brood, and females abandoning young had an increased return rate to the next breeding season. However, among brood abandoning females return rate was lower for the females with low lymphocyte levels. These results suggest that immunosuppression, as indicated by low lymphocyte levels, is a reproductive cost that may be partly compensated for by abandoning young.     

The Effect of an Experimental Brood Reduction on Male Desertion in the Panamanian Blue Acara Cichlid Aequidens coeruleopunctatus

Investment in present vs. future reproduction is a life-history trade-off faced by many animals. Because males generally pay a higher cost from lost mating opportunities than females, males are expected to react more strongly to changes in brood value. We examined the effect of an experimental brood reduction on male desertion in the substrate-brooding biparental cichlid Aequidens coeruleopunctatus under field conditions. We tested the prediction that brood reduction should decrease the duration of male care and examined the effect of brood reduction on the quality of male and female parental care. Our results show that males with reduced broods stopped providing parental care earlier than males with control broods. Males with reduced broods, however, also stayed longer with their broods as the season progressed. Brood reduction did not decrease daily investment in male or female parental care. We conclude that males trade off present and future reproduction by changing the duration but not the quality of parental care. The longer duration of male care in the experimental group later in the season suggests that the trade-off between present and future reproduction changes as the season progresses because the payoffs of desertion progressively decrease.     

The effect of paternal investment on female fertility intention in South Korea

Life history theory views reproduction as an outcome of resource allocation. The allocation of resources such as parental investments of time, energy and material resources involves trade-offs between number of offspring and timing of reproduction. Within the framework of mammalian parental investment, the outstanding feature of human reproduction is the high level of paternal care. Although empirical evidence suggests that human paternal investment may have evolved as a reproductive strategy to reduce infant and child mortality rates, the effects of actual paternal investment, including allocating time to child care, on female reproductive decisions have received relatively little attention. We examined the trade-off from two perspectives using a representative sample of married South Korean women aged 20–44 in 2005 (n=977). First, paternal investment in domestic labor, including child care and housework, was expected to be associated with women's preference regarding future reproduction. Second, relative paternal investment was expected to increase women's preference for future reproduction, especially among employed women. We found that increased paternal investment in child care and housework remarkably enhanced women's intention to have a second child, especially among employed women. In addition, although family members provide a low percentage of child care in South Korea, such help is likely to be a useful resource for second childbirth among employed women. Somewhat expectedly, older age and longer time since first birth had negative effects on women's second-child intention. There is growing evidence that, in the lowest fertility societies, paternal investment may be an essential resource for promoting future reproductive behavior of women, especially employed women.     

Have your cake and eat it too: male sand gobies show more parental care in the presence of female partners

Traditionally, male parental effort and mate attraction effortare expected to be in conflict as they compete for the sameresource budget. However, the quality of care provided by themale may be of a direct benefit to females and may provide animportant mate choice cue. In a laboratory experiment, we examinedhow males modified their parental behavior with respect to matingopportunity by allowing male sand gobies to mate with a singlefemale either in a big or small nest (a constraint on futuremating potential). We then exposed half of these males to thevisual stimulus from additional females and recorded male eggfanning and nest building (two components of care), courtshipbehavior, and reproductive success through out the brood cycle.We found that males fanned longer and more frequently and didmore nest construction in the presence of females and in bignests. Males guarding large nests courted females more thandid males guarding small nests. All males consumed eggs duringthe brood cycle, but complete clutch cannibalism was most frequentwhen males were guarding small nests in the absence of females.The pattern of filial cannibalism that we observed suggeststhat males prematurely terminated care when their reproductivepotential was low, that is, when there was little nest spacefor additional mating and no mates present. We found no supportfor a trade-off between mate attraction and parental care. Indeed,taken together our results suggest that males may use parentalcare as a courtship strategy and that males who invest in mateattraction also have higher parental effort.     

Mate desertion by male Great Reed Warblers Acrocephalus arundinaceus at the end of the breeding season

Male Great Reed Warblers usually take part in the care of offspring as nest defenders and by feeding young, but at the end of the breeding season they desert their mates with eggs or nestlings. Deserted females continue offspring care. Desertion does not depend on the male's mated status (polygynous or monogamous) nor on his past breeding success. Deserted females compensate for the loss of their partners by increasing the frequency of food-bringing, resulting in a reduction in the amount of time the nestlings are brooded. Although desertion may lead to increased rates of offspring mortality through predation, breeding success of deserted females was high, especially if the male assisted during the early stages. Deserters pay costs by giving up the chance of additional matings and by lowering the reproductive success of existing mates. Male warblers reduce the former cost by choosing the season of desertion and the latter is lowered by the female's high parental ability. A deserter was found to start moulting while his mate was still feeding his nestlings, and an earlier start to the moult may be the primary benefit that he gains. Male Great Reed Warblers usually take part in the care of offspring as nest defenders and by feeding young, but at the end of the breeding season they desert their mates with eggs or nestlings. Deserted females continue offspring care. Desertion does not depend on the male's mated status (polygynous or monogamous) nor on his past breeding success. Deserted females compensate for the loss of their partners by increasing the frequency of food-bringing, resulting in a reduction in the amount of time the nestlings are brooded. Although desertion may lead to increased rates of offspring mortality through predation, breeding success of deserted females was high, especially if the male assisted during the early stages. Deserters pay costs by giving up the chance of additional matings and by lowering the reproductive success of existing mates. Male warblers reduce the former cost by choosing the season of desertion and the latter is lowered by the female's high parental ability. A deserter was found to start moulting while his mate was still feeding his nestlings, and an earlier start to the moult may be the primary benefit that he gains. Male Great Reed Warblers usually take part in the care of offspring as nest defenders and by feeding young, but at the end of the breeding season they desert their mates with eggs or nestlings. Deserted females continue offspring care. Desertion does not depend on the male's mated status (polygynous or monogamous) nor on his past breeding success. Deserted females compensate for the loss of their partners by increasing the frequency of food-bringing, resulting in a reduction in the amount of time the nestlings are brooded. Although desertion may lead to increased rates of offspring mortality through predation, breeding success of deserted females was high, especially if the male assisted during the early stages. Deserters pay costs by giving up the chance of additional matings and by lowering the reproductive success of existing mates. Male warblers reduce the former cost by choosing the season of desertion and the latter is lowered by the female's high parental ability. A deserter was found to start moulting while his mate was still feeding his nestlings, and an earlier start to the moult may be the primary benefit that he gains. Male Great Reed Warblers usually take part in the care of offspring as nest defenders and by feeding young, but at the end of the breeding season they desert their mates with eggs or nestlings. Deserted females continue offspring care. Desertion does not depend on the male's mated status (polygynous or monogamous) nor on his past breeding success. Deserted females compensate for the loss of their partners by increasing the frequency of food-bringing, resulting in a reduction in the amount of time the nestlings are brooded. Although desertion may lead to increased rates of offspring mortality through predation, breeding success of deserted females was high, especially if the male assisted during the early stages. Deserters pay costs by giving up the chance of additional matings and by lowering the reproductive success of existing mates. Male warblers reduce the former cost by choosing the season of desertion and the latter is lowered by the female's high parental ability. A deserter was found to start moulting while his mate was still feeding his nestlings, and an earlier start to the moult may be the primary benefit that he gains. Male Great Reed Warblers usually take part in the care of offspring as nest defenders and by feeding young, but at the end of the breeding season they desert their mates with eggs or nestlings. Deserted females continue offspring care. Desertion does not depend on the male's mated status (polygynous or monogamous) nor on his past breeding success. Deserted females compensate for the loss of their partners by increasing the frequency of food-bringing, resulting in a reduction in the amount of time the nestlings are brooded. Although desertion may lead to increased rates of offspring mortality through predation, breeding success of deserted females was high, especially if the male assisted during the early stages. Deserters pay costs by giving up the chance of additional matings and by lowering the reproductive success of existing mates. Male warblers reduce the former cost by choosing the season of desertion and the latter is lowered by the female's high parental ability. A deserter was found to start moulting while his mate was still feeding his nestlings, and an earlier start to the moult may be the primary benefit that he gains.     

Does nest predation pressure influence the energetic cost of nest guarding in a teleost fish?

The energetic costs of providing parental care are widely documented, but rarely do studies consider the role of environmental variation (e.g., predation pressure) in this context. Here, we tested if variation in nest predation pressure influenced the energetic costs of parental care in smallmouth bass (Micropterus dolomieu), a teleost fish species that provides lengthy paternal care. First, we documented that nest predation pressure varied among the six lakes studied and the relative predation pressure ranking was consistent across a three year period. We used a combination of traditional proximate body composition (PBC) analyses and electromyogram (EMG) telemetry to quantify activity costs of nesting fish across these populations. The traditional approach revealed declines in energy stores across the parental care period but showed no evidence of an increased energetic cost to parents from populations with higher nest predation pressure. Comparing the distribution of EMG data from the two extremes of predation pressure revealed that males from the site of highest predation spent more time at higher EMG levels relative to the parents from the lake of lowest predation pressure. Although not statistically significant, males from the site of highest predation pressure also spent 21?C24?% of their time burst swimming when guarding young offspring compared to 10?C11?% for males at the site of lowest predation pressure. These differences in overall activity, a large contributor to the energy use of fish, may translate into longer recovery times and decreased future reproductive opportunities.