Evidence of selection for egg crypsis in conspicuous nests

ABSTRACT The value of egg coloration as crypsis, once accepted as a general principle, has recently been questioned because most experiments have failed to show that egg coloration deters predation. The nest‐crypsis hypothesis postulates that, among species that build conspicuous nests, selection for egg crypsis is relaxed or absent because visually searching predators detect nests prior to eggs. I tested the nest‐crypsis hypothesis using the large, relatively conspicuous nests of American Robins (Turdus migratorius), and eggs that differed markedly in color that were collected from the nests of Red‐winged Blackbirds (Agelaius phoeniceus), Brewer's Blackbirds (Euphagus cyanocephalus), and Yellow‐headed Blackbirds (Xanthocephalus xanthocephalus). Each nest (N= 22) received a clutch of each species during three sequential predation trials that were 16 d in duration. The order of clutch presentation was randomized for each nest. Survival trends for Brewer's and Yellow‐headed Blackbirds were similar, and higher than those for clutches of Red‐winged Blackbirds. By the end of trials, overall survival of the three clutch types was roughly equivalent. However, clutches of Red‐winged Blackbird eggs, the most conspicuous egg type to the human eye, were discovered sooner by predators. Because the experimental design controlled for effects of nest crypsis, nest location, and nest size, this difference in egg survival can be attributed to differences in egg pigmentation. Thus, my results support a role for egg coloration as camouflage in conspicuous nests.     

Optimizing surveys for marsh songbirds: does timing matter?

Analysis of data from point counts, a common method for monitoring bird population trends, has evolved to produce estimates of various population parameters (e.g., density, abundance, and occupancy) while simultaneously estimating detection probability. An important consideration when designing studies using point counts is to maximize detection probability while minimizing variation in detection probability both within and between counts. Our objectives were to estimate detection probabilities for three marsh songbirds, including Marsh Wrens (Cistothorus palustris), Swamp Sparrows (Melospiza georgiana), and Yellow‐headed Blackbirds (Xanthocephalus xanthocephalus), as a function of weather covariates and to evaluate temporal variability in detection probability of these three species. We conducted paired, unlimited radius, 10‐min point counts during consecutive morning and evening survey periods for our three focal species at 56 wetlands in Iowa from 20 April to 10 July 2010. Mean detection probabilities ranged from 0.272 (SE = 0.042) for Marsh Wrens to 0.365 (SE = 0.052) for Swamp Sparrows. Time of season was positively correlated with detection probability for Swamp Sparrows, but was negatively correlated with detection probability for Yellow‐headed Blackbirds, suggesting that detection probability increased during the breeding season for Swamp Sparrows and was highest early in the breeding season for Yellow‐headed Blackbirds. Understanding how detection probabilities of marsh songbirds vary throughout the breeding season allows targeted survey efforts that maximize detection probabilities for these species. Furthermore, consistent detection probabilities of marsh songbirds during morning and evening survey periods mean that investigators have more time to conduct surveys for these birds, allowing greater flexibility to increase spatial and temporal replication of surveys that could provide more precise estimates of desired population parameters.     

Effects of parents and Brown‐headed Cowbirds (Molothrus ater) on nest predation risk for a songbird

Nest predation limits avian fitness, so ornithologists study nest predation, but they often only document patterns of predation rates without substantively investigating underlying mechanisms. Parental behavior and predator ecology are two fundamental drivers of predation rates and patterns, but the role of parents is less certain, particularly for songbirds. Previous work reproduced microhabitat‐predation patterns experienced by Yellow Warblers (Setophaga petechia) in the Mono Lake basin at experimental nests without parents, suggesting that these patterns were driven by predator ecology rather than predator interactions with parents. In this study, we further explored effects of post‐initiation parental behavior (nest defense and attendance) on predation risk by comparing natural versus experimental patterns related to territory density, seasonal timing of nest initiation, and nest age. Rates of parasitism by Brown‐headed Cowbirds (Molothrus ater) were high in this system (49% nests parasitized), so we also examined parasitism‐predation relationships. Natural nest predation rates (NPR) correlated negatively with breeding territory density and nonlinearly (U‐shaped relationship) with nest‐initiation timing, but experimental nests recorded no such patterns. After adjusting natural‐nest data to control for these differences from experimental nests other than the presence of parents (e.g., defining nest failure similarly and excluding nestling‐period data), we obtained similar results. Thus, parents were necessary to produce observed patterns. Lower natural NPR compared with experimental NPR suggested that parents reduced predation rates via nest defense, so this parental behavior or its consequences were likely correlated with density or seasonal timing. In contrast, daily predation rates decreased with nest age for both nest types, indicating this pattern did not involve parents. Parasitized nests suffered higher rates of partial predation but lower rates of complete predation, suggesting direct predation by cowbirds. Explicit behavioral research on parents, predators (including cowbirds), and their interactions would further illuminate mechanisms underlying the density, seasonal, and nest age patterns we observed.     

Nest attendance by tropical and temperate passerine birds: Same constancy,different strategy

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The Effect of Male Incubation Feeding,Food and Temperature on the Incubation Behaviour of New Zealand Robins

Because of finite resources, organisms face conflict between their own self‐care and reproduction. This conflict is especially apparent in avian species with female‐only incubation, where females face a trade‐off between time allocated to their own self‐maintenance and the thermal requirements of developing embryos. We recorded incubation behaviour of the New Zealand robin (Petroica longipes), a species with female‐only incubation, male incubation feeding and high nest predation rates. We examined how male incubation feeding, ambient temperature and food availability (invertebrate biomass) affected the different components of females’ incubation behaviour and whether incubation behaviour explained variation in nest survival. Our results suggest that male incubation feeding rates of 2.8 per hour affect the female’s incubation rhythm by reducing both on‐ and off‐bout duration, resulting in no effect on female nest attentiveness, thus no support for the female‐nutritional hypothesis. The incubation behaviours that we measured did not explain nest survival, despite high nest predation rates. Increased ambient temperature caused an increase in off‐bout duration, whereas increased food availability increased on‐bout duration. While males play a vital role in influencing incubation behaviour, female robins attempt to resolve the trade‐off between their own foraging needs and the thermal requirements of their developing embryos via alternating their incubation rhythm in relation to both food and temperature.     

Sex‐Specific Parental Care in Response to Predation Risk in the European Roller,Coracias garrulus

Sublethal effects of predation constitute an important part of predation effects, which may modulate prey population and community dynamics. In birds, the risk of nest predation may cause a reduction in parental activity in the care of offspring to reduce the chance of being detected by predators. In addition, parents may modify their parental food allocation preferences within the brood in response to predation risk. Our aim in this study was to evaluate the effects of risk of nest predation on parental care and within‐nest food allocation in the European Roller (Coracias garrulus), an asynchronously hatching bird. We manipulated brood predation risk by placing a snake model near the nests that simulates the most common nest predator in the Mediterranean region. Our results show that males but not females increased their provisioning rate when they were exposed to the model and that despite this, nestlings’ body mass decreased in response to this temporary increase in predation risk. We did not find evidence that parents changed their food allocation strategy towards senior or junior nestlings in their nests in response to predation risk. These results show that the European roller modifies parental care in response to their perception of predation risk in the nest and a sex‐specific sensitivity to the threat, which suggests a different perception of offspring reproductive value by parents. Finally, our results show that changes in parental behaviour in response to nest predation risk might have consequences for nestling fitness prospects.     

Effect of alarm calling by male Red‐winged Blackbirds on nestling begging and female provisioning behavior

ABSTRACT Nestling begging and parental provisioning can attract nest predators and reduce reproductive success, so parents and their offspring might be expected to respond adaptively by minimizing predator‐attracting cues when predators threaten nests. Male Red‐winged Blackbirds (Agelaius phoeniceus) are well known for their antipredator alarm calls that contain information about the approach of potential nest predators. We examined the begging behavior of nestlings and the provisioning behavior of females in response to antipredator alarm calls of males to test the adaptive response hypothesis. Playback experiments provided no evidence that alarm calls function to switch off vocal begging; nestlings were equally likely to beg vocally during playback and control periods. Video recordings showed that male alarm calling had no significant effect on inappropriate vocal begging (in the absence of an adult), but significantly reduced the incidence of spontaneous calling (in the absence of begging). Adult females responded to male antipredator alarm calls by delaying their provisioning visits. In addition, although having no significant effect on use of nest‐arriving calls by females, male alarm calling significantly reduced their use of nest‐leaving calls. We conclude that nestling and female Red‐winged Blackbirds respond to male alarm calls in ways that might reduce the risk of predation, but nestlings beg vocally when females arrive to feed them, regardless of male alarm calling, perhaps to avoid a competitive disadvantage with broodmates.     

Mothers vigilantly guard nests after partial brood loss: a cue of nest predation risk in a paper wasp

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Introduced mammalian predators induce behavioural changes in parental care in an endemic New Zealand bird

The introduction of predatory mammals to oceanic islands has led to the extinction of many endemic birds. Although introduced predators should favour changes that reduce predation risk in surviving bird species, the ability of island birds to respond to such novel changes remains unstudied. We tested whether novel predation risk imposed by introduced mammalian predators has altered the parental behaviour of the endemic New Zealand bellbird (Anthornis melanura). We examined parental behaviour of bellbirds at three woodland sites in New Zealand that differed in predation risk: 1) a mainland site with exotic predators present (high predation risk), 2) a mainland site with exotic predators experimentally removed (low risk recently) and, 3) an off-shore island where exotic predators were never introduced (low risk always). We also compared parental behaviour of bellbirds with two closely related Tasmanian honeyeaters (Phylidonyris spp.) that evolved with native nest predators (high risk always). Increased nest predation risk has been postulated to favour reduced parental activity, and we tested whether island bellbirds responded to variation in predation risk. We found that females spent more time on the nest per incubating bout with increased risk of predation, a strategy that minimised activity at the nest during incubation. Parental activity during the nestling period, measured as number of feeding visits/hr, also decreased with increasing nest predation risk across sites, and was lowest among the honeyeaters in Tasmania that evolved with native predators. These results demonstrate that some island birds are able to respond to increased risk of predation by novel predators in ways that appear adaptive. We suggest that conservation efforts may be more effective if they take advantage of the ability of island birds to respond to novel predators, especially when the elimination of exotic predators is not possible.     

Sexual pigmentation and parental risk‐taking in yellow warblers Setophaga petechia

Adult‐directed predation risk imposes important behavioral constraints on parents and might thus alter relationships between costly sexual ornaments and parental performance. For instance, under low predation risk, highly ornamented individuals might display better parental performance than others, as predicted by ‘good parent’ models of sexual selection. However, under high risk of predation, highly ornamented individuals might abandon parental effort if conspicuous to predators, or if social partners are more willing to take parental risks when paired with highly ornamented mates. We experimentally elevated perceived adult‐directed predation risk near nests to explore how carotenoid‐ and phaeomelanin‐based pigmentation in both sexes relate to parental risk‐taking for offspring in the yellow warbler Setophaga petechia. Compared to other males, males with more intense carotenoid‐based pigmentation maintained higher levels of paternal effort under predation risk at highly concealed nests, but reduced nestling provisioning rate more at exposed nests. Further, when faced with predation risk, females with more phaeomelanin‐based pigmentation reduced nestling provisioning rate less than other females, regardless of nest concealment. Females displayed higher parental effort across treatments when paired to males with more colorful carotenoid pigmentation. However, birds did not reduce parental effort under risk less when paired to a highly ornamented mate, suggesting that predation risk did not accentuate differential allocation. Males did not take fewer parental risks than females. Results indicate that nest concealment modifies parental risk‐taking by males with colorful carotenoid‐based pigmentation, and suggest that female melanin‐based pigmentation may indicate boldness and greater a propensity to take parental risks.     

Timing,frequency, and duration of incubation recesses in dabbling ducks

Nest attendance is an important determinant of avian reproductive success, and identifying factors that influence the frequency and duration of incubation recesses furthers our understanding of how incubating birds balance their needs with those of their offspring. We characterized the frequency and timing (start time, end time, and duration) of incubation recesses for mallard (Anas platyrhynchos) and gadwall (Mareca strepera) hens breeding in Suisun Marsh, California, USA, and examined the influences of day of year, ambient temperature at the nest, incubation day, and clutch size on recess frequency and timing using linear mixed models. Mallard, on average, took more recesses per day (1.69 ± 0.80, mean ± standard deviation) than did gadwall (1.39 ± 0.69), and 45% of mallard nest‐days were characterized by two recesses, while only 27% of gadwall nest‐days were characterized by two recesses. Mallard morning recesses started at 06:14 ± 02:46 and lasted 106.11 ± 2.01 min, whereas mallard afternoon recesses started at 16:39 ± 02:11 and lasted 155.39 ± 1.99 min. Gadwall morning recesses started at 06:30 ± 02:46 and lasted 91.28 ± 2.32 min, and gadwall afternoon recesses started at 16:31 ± 01:57 and lasted 192.69 ± 1.89 min. Mallard and gadwall started recesses earlier in the day with increasing ambient temperature, but later in the day as the season progressed. Recess duration decreased as the season progressed and as clutch size increased, and increased with ambient temperature at the nest. The impending darkness of sunset appeared to be a strong cue for ending a recess and returning to the nest, because hens returned to their nests earlier than expected when recesses were expected to end after sunset. Within hens, the timing of incubation recesses was repeatable across incubation days and was most repeatable for mallard afternoon recesses and on days in which hens took only one recess. Hens were most likely to be away from nests between 04:00 and 07:00 and between 16:00 and 19:00; therefore, investigators should search for nests between 07:00 and 16:00. Our analyses identified important factors influencing incubation recess timing in dabbling ducks and have important implications for nest monitoring programs.     

Incubation recess behaviors influence nest survival of Wild Turkeys

In ground nesting upland birds, reproductive activities contribute to elevated predation risk, so females presumably use multiple strategies to ensure nest success. Identification of drivers reducing predation risk has primarily focused on evaluating vegetative conditions at nest sites, but behavioral decisions manifested through movements during incubation may be additional drivers of nest survival. However, our understanding of how movements during incubation impact nest survival is limited for most ground nesting birds. Using GPS data collected from female Eastern Wild Turkeys (n = 206), we evaluated nest survival as it relates to movement behaviors during incubation, including recess frequency, distance traveled during recesses, and habitat selection during recess movements. We identified 9,361 movements off nests and 6,529 recess events based on approximately 62,065 hr of incubation data, and estimated mean nest attentiveness of 84.0%. The numbers of recesses taken daily were variable across females (range: 1?7). Nest survival modeling indicated that increased cumulative distance moved during recesses each day was the primary driver of positive daily nest survival. Our results suggest behavioral decisions are influencing trade‐offs between nest survival and adult female survival during incubation to reduce predation risk, specifically through adjustments to distances traveled during recesses.     

Discriminating uniparental and biparental breeding strategies by monitoring nest temperature

Birds exhibit a wide diversity of breeding strategies. During incubation or chick‐rearing, parental care can be either uniparental, by either the male or the female, or biparental. Understanding the selective pressures that drive these different strategies represents an exciting challenge for ecologists. In this context, assigning the type of parental care at the nest (e.g. biparental or uniparental incubation strategy) is often a prerequisite to answering questions in evolutionary ecology. The aim of this study was to produce a standardized method unequivocally to assign an incubation strategy to any Sanderling Calidris alba nest found in the field by monitoring nest temperature profiles. Using drops of >3 °C in nest temperature (recorded with thermistors) to distinguish incubation and recess periods, we showed that the number of recesses and the total duration of these recesses from 09:00 to 17:00 h UTC allowed us reliably (99.1% after 24 h and 100% when monitoring the nest for at least 4 days) to assign the incubation strategy at the nest for 21 breeding adults (14 nests). Monitoring nest temperature for at least 24 h is an effective method to assign an incubation strategy without having to re‐visit nests, thereby saving time in the field and minimizing both disturbance and related increase in predation risk of clutches. Given the advantages of our method, we suggest that it should be used more widely in studies that aim to document incubation strategies and patterns in regions where ambient temperatures are at least 3 °C below the median nest temperature.     

Nest monitoring does not affect nesting success of Whinchats Saxicola rubetra

It is important to assess the effect that research activities may have on animals in the wild, especially when key parameters, such as breeding success, could potentially be influenced by observer activity. For birds, some studies have suggested that nest monitoring can increase the chances of nest failure due to predation, whereas others suggest that human nest visits may actually deter mammalian predators. Nest monitoring visits can also influence breeding success more indirectly by altering parental provisioning behaviour. Here, the influence of monitoring activities on nest success was examined in a ground‐nesting grassland bird, the Whinchat Saxicola rubetra. During the egg phase, a sample of nests were not visited between the initial finding event and the estimated hatching date; instead, the nest status was assessed at a distance. Daily survival rates (DSR) for these nests were compared with that of nests visited every 2 days. During the nestling phase, the effects of observer nest visits on parental provisioning behaviour were determined. Nest visits were found not to affect egg DSR significantly, and parental provisioning was disrupted for a maximum of 20 min (0.52% of the nestling period) following an observer visit. Given the variation in response to nest visits across species, we suggest that consideration should be given to observer impact in all studies where predation risk is high. Here, we illustrate a method for researchers to assess the impact of their nest visits to ensure they are not biasing estimates of breeding success.     

Substrate and structure of ground nests have fitness consequences for an alpine songbird

Songbird nests are an important life‐history component with multiple functions, including the creation of a suitable microclimate for offspring development. Thus, functional nest characteristics may influence fitness correlates, such as nestling size traits, and may co‐vary with prevailing environmental conditions. We investigated among‐ and within‐female variation in nest substrate, lining and decoration structures with associated fitness consequences (hatching success, nestling size traits, nest survival) across two breeding seasons for an alpine population of Horned Lark Eremophila alpestris. We combined these observations with explicit measures of nest temperature to address the influence of nest characteristics on microclimate. Nests in heather substrate had the coldest microclimates compared with grass and bare‐ground substrate, but also the greatest nest survival rates (68% versus 37–44% in other substrates), indicating the potential for substrate use decisions to reflect a trade‐off between microclimate and nest survival in response to prevailing weather and predation risk conditions. Furthermore, nest lining and nest decoration patterns indicated some support for a thermoregulatory function. Nests that were lined with willow (Salix sp.) seed‐down were associated with larger, heavier nestlings and the use of down lining decreased in frequency as the season warmed up. Nest decoration placed in front of the nest (e.g. stones or dirt clumps varying in mass from 5.3 to 186.6 g) was positively associated with warmer nest microclimates. Females demonstrated high phenotypic flexibility, as 61–94% of the observed variance in nest characteristics was explained by within‐female rather than among‐female differences. Such flexible nesting behaviour suggests the capacity to adjust to changing environmental conditions to maintain vital fitness correlates such as nest survival and nestling size development.     

Parent birds assess nest predation risk: influence of cavity condition and avian nest predator activity

Skutch hypothesized that nest predators visually assess parental activities to locate a prey nest, whereas parents modify fitness‐related traits to reduce the probability of nest predation. We examined how cavity condition and parental activity interact with avian nest predators to shape the nest success of two coexisting parid species, marsh tits Poecile palustris and oriental tits Parus minor, breeding in nest‐boxes during the incubation period. Nest‐boxes were manipulated to create a prolonged risk of nest predation (entrance diameter 2.6 cm control vs 5.5 cm treatment) soon after clutch completion. To measure changes in parental behavior, we also simultaneously simulated a pulsed risk of nest predation, using sound playbacks of a coexisting control bird and an avian nest predator. We found that the parent tits merely responded the pulsed risk, presumably due to an environment with high avian nest predator encounters, compared to the prolonged risk. Instead, both species spent more time on vigilance at the nest, only under prolonged risk conditions. The activity of corvids near the nest‐box was higher in the marsh tit than that in oriental tits. This activity was also higher in the treatment nest box than that in the control nest‐box. Nest predation during the incubation period was higher in marsh tits than in oriental tits, presumably due to higher and more plastic vigilance in oriental tits, compared to marsh tits. Our results highlight that the differences in cavity condition and parental activities at the nests of two coexisting non‐excavators may contribute to differential nest predation by attracting avian nest predators.     

Predation risk of eggs and nestlings relative to nest‐site characteristics of the Bull‐headed Shrike Lanius bucephalus

Appropriate nest‐site selection is one of the most important ways to minimize loss of reproductive investment due to predation. We determined the environmental characteristics associated with nest predation during the incubation and nestling periods of arboreal nesting Bull‐headed Shrikes on the oceanic Minami‐Daito Island where the predator community has low species diversity and includes only three introduced mammals: Ship Rat Rattus rattus, Japanese Weasel Mustela itatsi and Feral Cat Felis catus. Egg predation declined with increasing grassland cover around nests, whereas nestling predation declined with increasing nest concealment and nest height. Our results suggest that effective nest‐site characteristics for avoiding nest predation differ during the incubation and nestling periods and are dependent on the predator species and their search strategies, at least in habitats with low predator species diversity.     

Testing sources of variation in nestling‐stage nest success of Florida Scrub‐Jays in suburban and wildland habitats

Human modification of habitats can reduce reproductive success by providing novel cues to which birds may respond with behaviors that are actually maladaptive in those environments. Ad libitum human‐provided foods may provide the perception that urban habitats are food‐rich even as natural food availability decreases. Similarly, human activity may increase the perception that predation risk is high even as natural predators may decrease in abundance. In response, birds may reduce parental care with a subsequent cost to successful reproduction. Florida Scrub‐Jays (Aphelocoma coerulescens) in suburban areas have lower nest success during the nestling period than do wildland jays, possibly the result of such maladaptive responses, but maybe because of ecological differences with wildlands. We manipulated adult perception of predation risk and the availability of nestling foods in suburban and wildland areas to determine if these factors influenced parental care and nestling begging, and if the behavioral responses of adults influence nest survival during the nestling stage. Experimentally increasing perception of predation risk reduced parental care by both suburban and wildland females, but did not influence care by males. Increasing food availability, but not predation risk, had little influence on parental care, but resulted in decreased nestling begging rates and an increase in the frequency (pitch) of begging calls in both habitats. However, neither parental care nor food availability influenced nest survival during the nestling stage. Instead, the presence of helpers was the most important variable in nest survival analyses, suggesting that habitat‐specific differences in nest survival during the nestling stage were not simply the result of maladaptive parental behavior or shortage of nestling food resources in the suburban habitat. The lack of helpers combined with ecological differences, such as the abundance of nest predators, may be why fewer nests of Florida Scrub‐Jays survive during this stage in suburban areas.     

Are House Wren Troglodytes aedon eggs unusually strong? Test of the predicted effect of intraspecific egg destruction

As a result of opposing selective forces, the external strength of avian eggs should be near some size-specific optimum. However, in certain situations there should be selection on females to lay unusually strong eggs. According to one hypothesis, intraspecific egg destruction should favour increased egg strength as a means of defence against conspecific intruders. This hypothesis predicts that House Wrens Troglodytes aedon , a species well known for its tendency to destroy conspecific clutches, should be under selection for unusually strong eggs. However, the intensity of selection for strong eggs should also be modified by efficacy of nest defence against conspecific intruders in a given species (i.e. efficient nest defence by the breeding pair should weaken selection for unusually strong eggs). The goals of our study were: (1) to establish whether House Wren eggs are stronger than expected for their size; (2) to determine which structural mechanisms are responsible for their unusual strength; and (3) to test a hypothesis that, between wren species, the efficacy of nest defence and the intensity of egg-destroying behaviour affect the intensity of selection for unusually strong eggs. Our results demonstrated that: (1) House Wren eggs are 1.9 times stronger than expected for their size; (2) their unusual strength is achieved mostly by their unusually thick shells; and (3) eggs of the House Wren (extensive paternal nest defence; male egg-destroying behaviour suppressed during incubation) are significantly weaker structurally than eggs of the Marsh Wren Cistothorus palustris (reduced paternal nest defence; male egg-destroying behaviour present throughout incubation). These results are consistent with the hypothesis that the intraspecific egg-destroying behaviour and the efficacy of nest defence by the breeding adults have played a role in the evolution of strength of House Wren eggs.     

Devoted fathers or selfish lovers? Conflict between mating effort and parental care in a harem‐defending arachnid

When there is a temporal trade‐off between mating effort and parental care, theoretical models predict that intense sexual selection on males leads to reduced paternal care. Thus, high‐quality males should invest more in mating effort because they have higher chances of acquiring mates, whereas low‐quality males should bias their investment towards parental care. Once paternal care has evolved, offspring value should also influence males’ decisions to invest in offspring attendance. Here, we performed a manipulation under field conditions to investigate the factors that influence male allocation in either mating effort or parental care. We predicted that facultative paternal care in the harem‐holding harvestman Serracutisoma proximum would be negatively influenced by male attractiveness and positively influenced by offspring value. We found that attractive males were less likely to engage in egg attendance and that the higher the perceived paternity, the higher the caring frequency. Finally, egg mortality was not related to caring frequency by males, but predation pressure was much lower than that recorded in previous studies with the same population. Thus, the benefits of facultative male care may be conditional to temporal variation in the intensity of egg predation. In conclusion, males adjust their investment in either territory defence or egg attendance according to their recent mating history and perceived paternity. Our findings suggest that exclusive paternal care can evolve from facultative paternal care only if the trade‐off between mating effort and parental care is circumvented.