Nest predation of grassland bird species increases with parental activity at the nest

Alexander Skutch predicted that nest predation will increase with activity at nests, and that predation should be greatest during the nestling stage when parents are feeding young. We tested this hypothesis using three ecologically similar grassland bird species nesting on the high altitude grasslands of Wakkerstroom, South Africa. Parental activity, measured as adult arrival and departure frequency from the nest, was greater during the nestling than incubation stage. Nest predation, however, did not increase with parental activity between these stages in all three study species. However, nest site effects could have confounded this result. We therefore conducted an experiment that controlled for parental activity (by reusing natural nests of the study species with artificial clutches) in order to test for nest site effects. Nests that had a high rate of predation when used by active parents had a correspondingly high rate of depredation when the same nests were reused with artificial clutches (i.e. after controlling for parental activity). This result supports the notion that variation in nest site quality is the primary factor affecting nest predation rate. We also tested whether high predation during incubation is related to nest site effects. Nest predation rates of experimental clutches placed in reused nests that had been originally (when active parents were present) depredated during incubation stage were significantly higher than those that were originally depredated during the nestling stage. Finally, once nest site effects were accounted for, nest predation showed a positive increase with parental activity during the nestling stage across species.     

Habitat-specific clutch size and cost of incubation in eiders reconsidered

The energetic incubation constraint hypothesis (EICH) for clutch size states that birds breeding in poor habitat may free up resources for future reproduction by laying a smaller clutch. The eider (Somateria mollissima) is considered a candidate for supporting this hypothesis. Clutch size is smaller in exposed nests, presumably because of faster heat loss and higher incubation cost, and, hence, smaller optimal clutch size. However, an alternative explanation is partial predation: the first egg(s) are left unattended and vulnerable to predation, which may disproportionately affect exposed nests, so clutch size may be underestimated. We experimentally investigated whether predation on first-laid eggs in eiders depends on nest cover. We then re-evaluated how nesting habitat affects clutch size and incubation costs based on long-term data, accounting for confounding effects between habitat and individual quality. We also experimentally assessed adult survival costs of nesting in sheltered nests. The risk of egg predation in experimental nests decreased with cover. Confounding between individual and habitat quality is unlikely, as clutch size was also smaller in open nests within individuals, and early and late breeders had similar nest cover characteristics. A trade-off between clutch and female safety may explain nest cover variation, as the risk of female capture by us, mimicking predation on adults, increased with nest cover. Nest habitat had no effect on female hatching weight or weight loss, while lower temperature during incubation had an unanticipated positive relationship with hatching weight. There were no indications of elevated costs of incubating larger clutches, while clutch size and colony size were positively correlated, a pattern not predicted by the ‘energetic incubation constraint’ hypothesis. Differential partial clutch predation thus offers the more parsimonious explanation for clutch size variation among habitats in eiders, highlighting the need for caution when analysing fecundity and associated life-history parameters when habitat-specific rates of clutch predation occur.     

Clutch predation in great tinamous Tinamus major and implications for the evolution of egg color

Egg camouflage has been found to reduce predation in several ground‐nesting species. Therefore, the evolution of eggs that lack camouflage in ground nesting birds is puzzling. Even though clutch predation in the tropics is high, tinamous are the only tropical ground‐nesting birds that do not build a nest and do not lay cryptic eggs. I studied predation of great tinamou clutches in a lowland tropical forest and found that risk of predation was higher during incubation when the eggs are covered by the parent, than during laying when they are exposed, suggesting that predators primarily use cues from the incubating males to locate the clutch and not cues from the eggs. Clutch size had no effect on predation rate, even though larger clutches are more conspicuous to a human observer. Predation by visual cues is likely reduced during incubation by the camouflaged plumage and high nest attendance of males. If most predators use cues from the incubating male and not the eggs to locate clutches, then conspicuous egg color may have evolved in great tinamous as an intra‐specific signal. I evaluate hypotheses that may explain the maintenance of conspicuous egg color in tinamous.     

Reproductive tactics under severe egg predation: an eider’s dilemma

Parental defence against predators may increase offspring survival but entail other costs. Egg predation is frequent early in the laying sequence of the common eider, which differs in this and in several other ways from most other waterfowl. We test the hypothesis that permanent presence at the nest from the second or third egg is an adaptation for reducing egg predation in eiders. Two other alternative hypotheses for lower predation at later nest stages are early predation loss of the most vulnerable nests and seasonal decrease in predation risk. Analyses of predation rates at the one-egg and later stages refute these two alternatives. Early nest attendance by eider females is estimated to increase clutch survival by about 20% in four-egg and 35% in five-egg clutches, albeit probably at a cost of smaller clutch size.     

Causes of reduced clutch size in a tidal marsh endemic

We tested three hypotheses of clutch size variation in two subspecies of the swamp sparrow (Melospiza georgiana georgiana and M. g. nigrescens). Swamp sparrows follow the pattern of other estuarine endemics, where clutch size is smaller among tidal salt marsh populations (M. g. nigrescens) than their closest inland relatives (M. g. georgiana). Our results support predation risk and temperature, but not adult survival, as explanations of this pattern in swamp sparrows. Coastal nests were twice as likely to fail as inland nests, and parental activity around the nest site was positively related to clutch size at both sites. When brood size was controlled for, coastal adults visited nests less often and females vocalized less frequently during visits than inland birds, which may decrease nest detectability to predators. Coastal parents waited longer than inland birds to feed offspring in the presence of a model nest predator, but there was no difference in their response to models of predators of adults, as would be expected if coastal birds possessed increased longevity. Additionally, coastal females laid more eggs than inland females over a single season, following a within-season bet-hedging strategy rather than reducing within-season investment. Coastal territories experienced ambient air temperatures above the physiological zero of egg development more often, and higher temperatures during laying correlated with smaller clutches and increased egg inviability among coastal birds. Similar effects were not seen among inland nests, where laying temperatures were generally below physiological zero. Both subspecies showed an increase in hatching asynchrony and a decrease in apparent incubation length under high temperatures. Coastal individuals, however, showed less hatching asynchrony overall despite higher temperatures. Both air temperatures during laying and predation risk could potentially explain reduced clutch size in not only coastal plain swamp sparrows, but also other tidal marsh endemics.     

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.     

Nest predation is little affected by parental behaviour and nest site in two African Sylvia warblers

Parental behaviour and nest site are supposed to affect nest predation in birds. Few nest visits and high nest attentiveness are assumed to lead to low predation rates. Poorly concealed nests are thought to be more likely to be preyed upon than well concealed nests. Studies on the relationship between parental behaviour, nest site, and nest predation are rare and none have, so far, been conducted in the Afrotropics. We studied the effect of nest site, nest visitation rate, and nest attentiveness on the nest predation rate of the two tropical warblers Sylvia boehmi and Sylvia lugens in Kenya. Parental behaviour and predation on nests of 13 breeding pairs of both species were observed daily in two consecutive breeding seasons. In both species, parental activity at the nest was low [0.9 trips to the nest in 30 min during incubation, maximum 4.6 (S. boehmi) and 5.8 (S. lugens) trips to the nest in the nestling stage]. Predation rates in both species were high (Mayfield nest success 19.4 and 33.2%). Our analysis revealed only weak evidence for an effect of nest site, nest visitation rate, and nest attentiveness on the predation rate. It is suggested that smaller clutches of tropical in comparison to northern temperate birds result from lower feeding rates in tropical ecosystems with high predation rates (Skutchs hypothesis). The underlying mechanism could not be proven in this study.     

Nest height, nest concealment, and predator type predict nest predation in superb fairy-wrens (Malurus?cyaneus)

Three factors and their interaction effects are increasingly recognized as important determinants of nest predation: nest concealment, nest height, and predator type. The risk of nest predation is predicted to vary across these variables because of nest detectability and accessibility. In general, however, few studies examine how these three variables interact in relation to nest predation, focusing instead on either nest concealment or nest height (whereby predator identity is usually not known). In this study, we examine the role of nest concealment and nest height for nest survival using both artificial and natural nests in the superb fairy-wren (Malurus cyaneus). We indirectly identified potential predators through marks left on artificial eggs and footprints left on tracking tunnels. Predation level at artificial nests was lower than at natural nests, and this could be due to a failure of some nest predators to locate cryptic nests in the absence of cues provided by parental activity. Our results supported the prediction that exposed and concealed nests have different levels of nest predation, which can be explained by variation in predator type. Visual predators were only detected at exposed nests, and survival from visual predators was lower for high nests that were also exposed. However, olfactory predators were detected irrespective of nest height or nest concealment. Because rodents use olfaction to locate nests, this could explain the lack of association between nest concealment and predation outcome at low nests. In addition, rodent footmarks near nests were significantly associated with rodent tooth marks on eggs.     

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.     

Nest predators of Lance-tailed Manakins on Isla Boca Brava, Panamá

ABSTRACT.   Nest predation is often the primary cause of nest failure for passerines. Despite this, little is known about predation rates and the nest predators of birds in the tropics. I used video cameras to monitor seven Lance-tailed Manakin ( Chiroxiphia lanceolata ) nests on Isla Boca Brava, Panamá. One nest fledged young and six nests failed due to predation. I recorded five predation events involving four avian predators and one mammalian predator. Crested Oropendolas ( Psarocolius decumanus ) predated two nests and a Roadside Hawk ( Buteo magnirostris ) and a Black-chested Jay ( Cyanocorax affinis ) each predated one. The mammalian predator was a common opossum ( Didelphis marsupialis ). All avian predation was diurnal; the mammalian predation was nocturnal. My results suggest that tropical birds are subject to a diverse suite of nest predators, and that avian predators may be an important cause of nest failure at my study site.     

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.     

Nest predation and habitat change interact to influence Siberian jay numbers

We studied Siberian jays, breeding in northern Sweden, to examine the potential for interactions between nest predation and reduced vegetation heterogeneity around nest sites to cause a decrease in jay numbers. Parent behaviour and nests are highly cryptic in the species. Our 12-year data showed, however, that nests had a probability of only 0.46 to be successful and produce at least one nestling. Nest predation was intense and a main cause of nest failure. All predators that could be identified were visually oriented hunters, mostly other corvids able to colonize taiga forest only close to human settlements. Consistent with the idea that predators used visual cues, nest predation increased with parental activity, which suggests that predators used parental provisioning trips to locate nests. Furthermore, a reduction in daily nest survival rates with decreasing amount of nesting cover was more pronounced in areas with high corvid activity as predicted when cover mediates the hunting efficiency of visual oriented predators. Declining temperatures interacted with the effects of habitat characteristics to further reduce daily nest survival rates suggesting that parents were not able to increase nest visitation rates to satisfy the higher energy demands of their nestlings without endangering the nest. Our results identify a mechanism through which predation and human-induced reduction in nesting cover on a larger scale may interact to cause a reduction in Siberian jay numbers larger than expected from habitat loss alone.     

Winter territoriality in migrant Black Redstarts Phoenicurus ochrurus in the Mediterranean area

The nest predation hypothesis predicts that clutch size should be inversely related to the risk of nest predation. In this paper we analyse data on nest predation, clutch size and fledgling production in 128 Jackdaw nests in order to test if clutches are larger in the safer sites. The predation rate was positively correlated with the minimum nest-entrance dimensions (predation is greater in nests with large entrances). Clutch size was not related to the risk of nest predation, and our data do not therefore support the nest predation hypothesis.     

Age-specific nest-site preference and success in eiders

Variation in nest concealment is puzzling given the expected strong selection for safe nest sites. Selecting a concealed nest may decrease the risk of clutch predation but hinder parents from escaping predators, providing a possible solution to this paradox. Because the relative value of current versus future reproduction may vary with breeder age or state, nest concealment may also vary as a function of these attributes. We tested four predictions of the female and clutch safety trade-off hypothesis in eiders (Somateria mollissima): (1) nest concealment is negatively related to escape possibilities, (2) our capture rate of females is higher in covered nests, (3) egg predation is higher in open nests, and (4) overall nest success is unrelated to nest habitat. We also analysed nest microhabitat preferences and nest success relative to breeder age and body condition, controlling for nest spatial centrality. As expected, nest concealment and potential escape angle were negatively related, and capture by us, indicating female predation vulnerability, increased with nest cover. Clutch size was smaller in open nests, suggesting higher partial clutch predation, while it was larger among experienced and good-condition breeders. The probability of successful hatching was unrelated to nest habitat, positively associated with breeder experience, and negatively associated with hatching date. Experienced females selected more concealed and centrally located nests without sacrificing potential escape angles. The age-specific spatial distribution of nests on islands was unrelated to nest initiation dates, indicating no apparent competition. The age-specific preference of eiders for concealed nests may reflect declining reproductive value with age or confidence in surviving despite selecting a concealed nest. The apparently positive relationship between female age and survival and fecundity in eiders refutes the former alternative. Individual improvement in choosing safe nest sites, coupled with differential survival of individuals performing well, most likely explains age-specific nest-site preference and success.     

Nest survival in year‐round breeding tropical red‐capped larks Calandrella cinerea increases with higher nest abundance but decreases with higher invertebrate availability and rainfall

Nest survival is critical to breeding in birds and plays an important role in life‐history evolution and population dynamics. Studies evaluating the proximate factors involved in explaining nest survival and the resulting temporal patterns are biased in favor of temperate regions. Yet, such studies are especially pertinent to the tropics, where nest predation rates are typically high and environmental conditions often allow for year‐round breeding. To tease apart the effects of calendar month and year, population‐level breeding activity and environmental conditions, we studied nest survival over a 64‐month period in equatorial, year‐round breeding red‐capped larks Calandrella cinerea in Kenya. We show that daily nest survival rates varied with time, but not in a predictable seasonal fashion among months or consistently among years. We found negative influences of flying invertebrate biomass and rain on nest survival and higher survival of nests when nests were more abundant, which suggests that nest predation resulted from incidental predation. Although an increase in nest predation is often attributed to an increase in nest predators, we suggest that in our study, it may be caused by altered predator activity resulting from increased activity of the primary prey, invertebrates, rather than activity of the red‐capped larks. Our results emphasize the need to conduct more studies in Afro‐tropical regions because proximate mechanisms explaining nest predation can be different in the unpredictable and highly variable environments of the tropics compared with the relatively predictable seasonal changes found in temperate regions. Such studies will aid in better understanding of the environmental influences on life‐history variation and population dynamics in birds.     

Chick survival in relation to nest site: is the Antarctic petrel hiding from its predator?

In birds, the characteristics of the nest site may affect reproductive success. We found that shelter is an important characteristic of the Antarctic petrel (Thalassoica antarctica) nests because shelter prevents chick predation. However, the benefit of shelter was countervailed by melt water which mainly entered well-sheltered nests. Chick survival was monitored until the chick was left unattended for the first time. Late-hatched chicks had a higher survival probability than early-hatched chicks, possibly because late hatchers swamp the predator, the south polar skua (Catharacta maccormicki). Poorly sheltered nests tended to be occupied by parents with low body mass and late-hatched eggs. The results suggest that both shelter per se and parental characteristics may explain the relationship between predation risk and shelter. We need experiments to study the influence of nest site on reproductive success, and we need to map the frequency of melt water as a cause of reproductive failure.     

Relative effects of nest size and site on the risk of predation in open nesting passerines

Large nests may incur fitness cost in terms of conspicuousness to predators, but the effect of nest size on predation risk can be confounded by effects of nest site and parental characteristics. I examined relative effects of nest size and placement by experimentally exchanging subsets of inactive nests baited with artificial clutches, among three open-cup nesting passerine species characterized by different nest size, placement and predation rate. The prediction that increasing nest size (original nest replaced by nest of the larger species) would increase predation, while decreasing nest size (replacement by nest of the smaller species) would decrease predation, relative to control (replacement by conspecific nest) was not supported in any species. The prediction that predation should be higher for large nests compared to small ones, even after exchanging nests among species-specific sites, was not supported. Predation rate differed among species (combined site/nest effect) before manipulation, whereas only the effect of nest site was significant after manipulation. This means that predation differed between species-specific nest sites, irrespective of the nest placed at these sites, but not between large (thrush) and small (warbler) nests, irrespective of their placement. Results do not suggest that nest predation selects directionally for smaller nest size. This conclusion could be specific to the study system characterized by high nest densities and high predation rate.     

Effects of nest-site characteristics and parental activity on cowbird parasitism and nest predation in Brown-and-yellow Marshbirds

ABSTRACT Nest‐site selection and nest defense are strategies for reducing the costs of brood parasitism and nest predation, two selective forces that can influence avian nesting success and fitness. During 2001–2002, we analyzed the effect of nest‐site characteristics, nesting pattern, and parental activity on nest predation and brood parasitism by cowbirds (Molothrus spp.) in a population of Brown‐and‐yellow Marshbirds (Pseudoleistes virescens) in the Buenos Aires province, Argentina. We examined the possible effects of nest detectability, nest accessibility, and nest defense on rates of parasitism and nest predation. We also compared rates of parasitism and nest predation and nest survival time of marshbird nests during the egg stage (active nests) with those of the same nests artificially baited with passerine eggs after young fledged or nests failed (experimental nests). Most nests (45 of 48, or 94%) found during the building or laying stages were parasitized, and 79% suffered at least one egg‐predation event. Cowbirds were responsible for most egg predation, with 82 of 107 (77%) egg‐predation events corresponding to eggs punctured by cowbirds. Nests built in thistles had higher rates of parasitism and egg predation than nests in other plant, probably because cowbirds were most active in the area where thistles were almost the only available nesting substrate. Parasitism rates also tended to increase as the distance to conspecific nests increased, possibly due to cooperative mobbing and parental defense by marshbirds. The proportion of nests discovered by cowbirds was higher for active (95%) than for experimental (29%) nests, suggesting that cowbirds used host parental activity to locate nests. Despite active nest defense, parental activity did not affect either predation rates or nest‐survival time. Thus, although nest defense by Brown‐and‐yellow Marshbirds appears to be based on cooperative group defense, such behavior did not reduce the impact of brood parasites and predators.     

Nest site attributes and temporal patterns of northern flicker nest loss: effects of predation and competition

To date, most studies of nest site selection have failed to take into account more than one source of nest loss (or have combined all sources in one analysis) when examining nest site characteristics, leaving us with an incomplete understanding of the potential trade-offs that individuals may face when selecting a nest site. Our objectives were to determine whether northern flickers (Colaptes auratus) may experience a trade-off in nest site selection in response to mammalian nest predation and nest loss to a cavity nest competitor (European starling, Sturnus vulgaris). We also document within-season temporal patterns of these two sources of nest loss with the hypothesis that flickers may also be constrained in the timing of reproduction under both predatory and competitive influence. Mammalian predators frequently depredated flicker nests that were: lower to the ground, less concealed by vegetation around the cavity entrance and at the base of the nest tree, closer to coniferous forest edges and in forest clumps with a high percentage of conifer content. Proximity to coniferous edges or coniferous trees increased the probability of nest predation, but nests near conifers were less likely to be lost to starlings. Flickers may thus face a trade-off in nest site selection with respect to safety from predators or competitors. Models suggested that peaks of nest predation and nest loss to eviction occurred at the same time, although a competing model suggested that the peak of nest loss to starlings occurred 5 days earlier than the peak of mammalian predation. Differences in peaks of mammalian predation and loss to starlings may constrain any adjustment in clutch initiation date by flickers to avoid one source of nest loss.     

Experimental Evidence that Sexual Displays are Costly for Nest Survival

Nest predation is one of the most significant limitations for successful breeding of tropical passerines. Thus, parental strategies may include choosing appropriate nest sites and behaving in ways that minimize predation. Habitat characteristics that may influence nest success include degree of nest concealment, proximity to habitat edge, plant architecture as well as several others cited in the literature. However, few studies have examined display behavior as a factor that could also influence nest survival. We experimentally tested whether sexual motor displays served as a cue for visually oriented predators to locate artificial nests in a population of blue‐black grassquits Volatinia jacarina, a Neotropical passerine that exhibits a complex sexual display and is subjected to elevated rates of nest predation. We also evaluated the effect of nest substrate on survival. Predation rate was higher for nests within territories of displaying males relative to areas without displaying males and for nests placed in shrubs relative to grasses. Predation increased sharply in the third experimental replicate, at the end of the breeding season, which suggests that predators may develop a search image for nests or may become more abundant during specific periods of the season. Avian predators appear to be the most important nest predators. Results suggest that there may be a trade‐off between the increase in fitness derived from sexual displays of males to attract potential mates and the decrease owing to predation of active nests within their territories.