Accepting unrelated broods helps replacement male yellow-headed blackbirds attract mates

Replacement male yellow-headed blackbirds (Xanthocephalus xanthocephalus)did not destroy broods sired by the previous territorial maleand they showed no aggression toward females with unrelatedbroods. To test whether their tolerance of unrelated young wasmisdirected normal parental care, we removed males from experimentalterritories after primary nests were completed but before secondarynests were initiated. Replacement males fed young that theypresumably had sired in secondary nests and ignored foster youngin primary nests, whereas control males fed young in their primarynests. To identify potential benefits of accepting unrelatedyoung, we analyzed patterns of within-season breeding dispersaland of female settlement on territories following nest lossesto predators. Although some female yellowheads do renest onthe same territory following nest failures, the number of nestsinitiated on territories after a predation event was significantlylower than the number initiated on territories without predationover the same period of time. This implies that late-settlingfemales use the number of active or failed nests and/or thenumber of females on a territory when choosing where to breed.If replacement males that accepted unrelated offspring attractmore new females in the remainder of the current breeding seasonthan infanticidal males, then tolerance of unrelated young byreplacement males may be adaptive in some polygynous birds.     

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.     

Maternal Care in the Burrower Bug <Emphasis Type="Italic">Adomerus triguttulus</Emphasis>: Defensive Behavior

Some burrower bugs (Heteroptera: Cydnidae) show complex patterns of maternal care, including defense against predators and the provisioning of food to nymphs. Recently, the subsocial cydnid bugs have attracted the interest of researchers as model systems to study the behavioral ecology of parental investment. However, there have been few attempts to quantify the fitness benefits of maternal behavior other than provisioning. Here, we examined the maternal behavior of Adomerus triguttulus and its adaptive significance in terms of offspring survival in the field. A. triguttulus young depend on fallen nutlets of myrmecophorous mints, Lamium spp. Under field conditions, females attend offspring, from eggs to second instar nymphs, in nests on the ground under the litter. When disturbed, the females showed aggressive responses against the source of disturbance. The females often carried spherical clutches of eggs away from the nest when heavily disturbed. Female-removal experiments in the field demonstrated a defensive function of the female behavior; predators, such as ants, attacked egg clutches without females and the clutches often disappeared during the experiment. Egg clutches without females sometimes also suffered from fungal infection. Selective factors on maternal defensive behavior in A. triguttulus are discussed in terms of habitat properties possibly emerging from insect–plant associations.     

Do trade‐offs between predation pressures on females versus nests drive nest‐site choice in painted turtles?

Predation strongly influences reproductive behaviours because reproducing individuals must balance mortality risks to themselves and to their offspring. In many freshwater turtles, the nest predation risk decreases with nest distance from water, whereas the predation risk to females increases farther from water. To determine whether predation pressure influences the distance from water at which female turtles nest, we measured predation pressure on nesting females and on nests, as well as the distances of nests to water, in two populations of painted turtles. Using models, we found that female survival in both populations was high and did not vary with distance from water. Nest survival was also uncorrelated with nest distance to water, although it was significantly lower than adult survival in both populations and was only 1.2% in one population. Our results suggest that nest sites are not predictably safe from predators. Instead, turtles may hedge their bets by nesting over a wide range of distances from water because any distance is risky for nests and no distance is particularly risky for the nesting female. We suggest that other factors, such as suitable incubation conditions and/or post‐emergence hatchling survival, probably play a larger role than predation in driving nest‐site choice in painted turtles.     

Communal Relationships in a Social Spider Mite, Stigmaeopsis longus (Acari: Tetranychidae): An Equal Share of Labor and Reproduction between Nest Mates

The social spider mite, Stigmaeopsis longus (Saito), builds web‐nests and shares resources with fellow nest members. It does not show any distinct morphological castes. In this study, we observed whether there is any division of labor in reproductive and non‐reproductive tasks. Under experimental conditions, female density per nest did not affect per capita fecundity, even though the nest area allocated to an individual female, in which it feeds and oviposits, decreased with increased female density. Video observations on the behavior of either one or two females showed that there were no differences between these situations in the time budgets of all behaviors, nor in the nest‐weaving behavior of females. Furthermore, detailed behavioral analyses between the two situations suggested that S. longus females share reproduction and labor evenly with their nest mates, probably mediated through physical or chemical communication. Therefore, we concluded that the sociality of this mite species should be categorized as communal.     

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.     

Infanticide in great reed warblers: secondary females destroy eggs of primary females

In 1994–1995 artificial nests with attached model eggs were put into territories that were known to have been occupied by male great reed warblers,Acrocephalus arundinaceusin previous years. Because the eggs were made of soft plasticine, predators left peckmarks in them and this enabled us to identify predators by comparing peckmarks with reference marks made by various species. Previous field data had suggested that infanticidal behaviour existed in our study population, as nests of primary females suffered a three times higher rate of nest loss during the egg-laying period than nests of secondary and monogamous females. The presence of infanticide was supported by the experiment. Small peckmarks resembling those of a great reed warbler occurred almost exclusively in territories occupied by great reed warblers, in particular when a new female settled in the territory. The newly settled females built nests closer to depredated than non-depredated nests. That small peckmarks occurred when new females settled strongly suggests that it is secondary female great reed warblers that commit infanticide on eggs of primary females. Females of low harem rank are expected to gain from infanticidal behaviour because a low ranked female gets a higher proportion of male parental investment when the nest of the primary female fails.     

Nest Defence Behaviour of the Eurasian Kestrel (Falco tinnunculus) Against Human Predators

Man has become the main predator of many animal species. Because the characteristics of humans are quite distinct with respect to other terrestrial predators, the cost and benefits of defence behaviour may also differ. In this paper, we study the factors affecting nest defence behaviour of the Eurasian Kestrel ( Falco tinnunculus canariensis ) against a potential human predator throughout the reproductive cycle, as well as the balance of cost and benefits of this behaviour. The study population inhabits the island of Tenerife, and the nests are located on cliffs. The intensity of the defensive behaviour was unrelated to the frequency of human visits, prey abundance (Orthoptera, Coleoptera, lizards, birds and rodents), laying date, or number of offspring in the nest. Both males and females increased their defensive behaviour as the nesting period advanced, particularly when the chicks were older than 15 days. Moreover, the intensity of the defensive behaviour, especially of males, decreased when nests were more inaccessible. Although nest defence behaviour against humans appeared to be similar to those against other predators, the benefits are not clear because the probability of nest robbing was greater for these more aggressive pairs.     

Variation in social behavior within a spider mite genus, Stigmaeopsis (Acari: Tetranychidae)

The spider mites belonging to the genus Stigmaeopsis constructextremely dense oval woven roofs (web-nests) over depressionson the lower surface of host leaves and are known to have akind of sociality. The four species that occur on bamboo plantsin Japan show different nest areas. The nest area of Stigmaeopsislongus is the largest, followed by that of S. celarius, S. takahashii,and S. saharai in decreasing order. Smaller nests effectivelyprevent adults of several predator mite species from intruding.We hypothesized that variation in nest size reflects differentanti-intruder adaptations of this mite group in relation totheir sociality. The larger nest makers may adopt an alternativeanti-intruder strategy, namely, counterattack by a large group,so as to compensate for the disadvantage of large nests. S.longus and S. celarius adults effectively defended their largenests against potential predators, and the effects of nest defenseincreased with the number of individuals in a nest. S. takahashiiand S. saharai revealed no counterattack effect. Counterattackabilities that increase with the adult density, and thus, socialitymay compensate for the vulnerability of larger nests.     

Age of the incubating parents affects nestling survival: an experimental study of the herring gull Larus argentatus

The quality of conditions provided by avian parents will have consequences for both parental and offspring fitness. While many components of avian reproduction appear to vary with parental age, the effect of age on incubation has largely been ignored so far. In this study, we tested whether young herring gulls provide a different incubation environment from mature ones and whether this has consequences for offspring performance. Laying and rearing conditions were standardised using a cross-fostering protocol. Egg predation rates tended to be higher in the nests of young parents. However, nest site, nest construction and egg temperature during incubation did not vary with parental age. Overall, the duration of incubation was shorter in young compared to mature birds and this reflected the later laying date of the former, since incubation duration generally decreased across the season. However, male eggs incubated by young parents had longer incubation periods than predicted for their laying dates. In contrast, incubation length of female eggs incubated by young pairs, and of male and female eggs incubated by mature birds did not deviate from the expected for any given laying date. Offspring that had been incubated by young parents had considerably poorer survival than those incubated by mature pairs, despite being reared under standardized, favourable conditions (singly, by mature parents). This was due to increased mortality among female chicks that had been incubated by young parents. The chicks incubated as eggs by young and mature birds, which survived until fledging, did not differ in body mass and size growth, or body condition. The results of this study demonstrate that parental age can influence offspring performance via variation in incubation environment, and that females are more susceptible than males to conditions experienced during embryonic development.     

Intraspecific nest parasitism in the European starling Sturnus vulgaris

Biochemical genetic markers were used along with conventional methods (abnormal laying sequence/clutch size, unusual egg shape/pigmentation) to identify intraspecific nest parasitism at two British nestbox colonies of the European starling. Between 11 and 37% of first clutches were parasitized during 1977–1979. Parasitic females probably comprised all of the following categories: (1) paired females contesting a nestbox occupied by another pair; (2) previously paired females who had laid a clutch but had been unsuccessful; (3) unpaired females who had copulated with males that already had a mate and nest site; and (4) ‘professional’ nest parasites who distributed at lest some of their eggs in one or more nests other than their own. Although parasitized nests had higher clutch sizes, parasitism led to fewer host young fledging per egg laid, mainly through the eviction of eggs and subsequent nest desertion. Number of parasitic young fledged per egg laid was highest when eggs were laid synchronously with the host, when host clutches were larger, or a smaller number of parasite eggs were added to a nest, thus favouring parasites that distribute their eggs amongst a number of nests. A greater pressure on nest sites may have accounted for the higher levels of parasitism at the Aberdeen colony and for the greater number of parasite eggs laid in a nest. Although most parasitic female starlings appeared to be much less successful than non-parasitic ones, nest parasitism in the starling might evolve directly when one or more of the following advantages are present. (1) There are no constraints on the number of eggs a female may lay but there are constraints on the number of young she may feed adequately. (2) Female survival is increased by having fewer or no eggs/young to care for. (3) Current feeding conditions favour the survival of more young than would be produced by the most common clutch size. Intraspecific nest parasitism is considered to be a first stage in the evolution of interspecific nest parasitism.     

Effects of Mesopredators on Nest Survival of Shrub-Nesting Songbirds

ABSTRACT Although nest predation is often the single largest source of mortality in avian populations, manipulative studies to determine predator impacts on nest survival are rare, particularly studies that examine impacts of mid-size mammalian predators (hereafter, mesopredators) on nest survival of shrub-nesting birds. We quantified nest survival and identified nest predators of shrub-nesting songbirds within 4 large (approx. 40-ha) exclosures and 4 control sites within a longleaf pine (Pinus palustris) ecosystem. During 2003–2006, we located and monitored 535 shrub nests (222 with videography) for 4,804 nest-days to quantify daily nest survival and document predation events. We found no support for a treatment effect, suggesting mesopredators had little impact on daily nest survival (0.9303 in controls and 0.9260 in exclosures) of shrub-nesting songbirds. For the 5 most commonly monitored species, daily nest survival within species was constant. Our analysis suggested that shrub nests were most vulnerable during the nestling stage and presence of cameras on nests increased survival with the increase in survival being more pronounced during the incubation stage. We filmed 107 nest predation events, identifying predators at 88 nests. Of these 88 nests, snakes caused 33%, red imported fire ants (hereafter fire ants, Solenopsis invicta) 28%, raptors 17%, corvids 8%, mesopredators 6%, and small mammals 8% of nest predations. Cause-specific nest predation in controls and exclosures did not differ from expectation, providing evidence that compensatory predation did not occur. Nest predators differed from expectation with regard to nest stage; fire ants and raptors only depredated nests during the nestling stage. Presence of cameras had no effect on nest abandonment. Fire ants were the most prevalent nest predator, and nest predation by fire ants was only observed on nestlings, potentially reducing likelihood of renesting. Magnitude and timing of fire ant predation suggests that fire ants may be the most influential nest predator of shrub-nesting birds within the longleaf pine ecosystem. Our data suggest that controlling mesopredators will have no effect on nest success of shrub-nesting birds within longleaf pine forests.     

Why do Both Parents Incubate in the Kentish Plover?

Incubation by both parents is a common parental behaviour in many avian species. Biparental incubation is expected if the survival prospects of offspring are greatly raised by shared care, relative to the costs incurred by each parent. We investigated this proposition in the Kentish plover Charadrius alexandrinus, in which both parents incubate the clutch, but one parent (either the male or the female) usually deserts after hatching of the eggs. We carried out a mate‐removal and food supplementation experiment to reveal both the role of the sexes and food abundance in maintaining biparental incubation by removing either the male or the female from the nest for a short period of time. In some nests we provided supplementary food for the parent that remained at the nest to reduce the costs of incubation, whereas other nests were left unsupplemented. Although males spent more time on incubation after their mate had been removed, females’ incubation did not change. Notwithstanding the increased male incubation, total nest attentiveness was lower at uniparental nests than at biparental controls. However, incubation behaviour was not influenced by food supplementation. We conclude that offspring desertion during incubation is apparently costly in the Kentish plover, and this cost cannot be ameliorated with supplementary food.     

Handicapped females receive more feedings during incubation from their mates: support for the female nutrition hypothesis

The female nutrition hypothesis posits that provisioning intensity of incubating females by their mates may depend on female needs and ensure proper incubation and a corresponding high hatching and breeding success of breeding pairs. Here, we have handicapped female pied flycatchers Ficedula hypoleuca at the beginning of incubation by clipping two primaries on each wing and filmed nests during incubation and later nestling provisioning to estimate male involvement in incubation feeding at the nest and in offspring care. Incubation feeding was more frequent at late nests. Correcting for this seasonal effect, incubation feeding was significantly affected by treatment and twice as high at experimental as at control nests. There was no effect of the experiment on female incubation attendance. The handicap did not result in any effect on hatching and breeding success, nestling growth and male or female provisioning and mass at the end of the nestling period. Males adjust their incubation feeding activity at the nest to female energetic requirements during incubation.     

Microhabitat structure and avian nest predation risk in an open Argentinean woodland: an experimental study

We studied the effect of the general structure of the nest plant, especially the presence of thorns, and the structural homogeneity of the nest patch, on the vulnerability of nests to predation, using natural and artificial nests. Artificial nests placed in non-thorny plants had a significantly lower predation rate and higher daily survival rate than those in thorny plants. The addition of a ‘thorny microhabitat’ around the immediate proximity of nests placed in non-thorny plants did not have any effect on vulnerability of nests to predation. Conversely, natural nests were located in patches of habitat with a higher density of the species of plant that supported the nest compared to patches selected at random. However, daily survival rate was similar for natural nests placed in patches with a higher or lower density of the species of nest plant in the four bird species analysed. Similarly, survival of artificial nests did not increase with the presence of a higher number of plants similar to the nest plant in the nest patch. Thus, the observed patterns of survival for natural and artificial nests did not seem to support the potential prey-site hypotheses. Birds appeared to be the main nest predators in this ecosystem. Behavioural aspects of the identified predators and habitat structure could explain the lack of effect of thorns and nest patch characteristics on nest survival.     

Within-nest spawning-site preferences of female bluefin damselfish: the effect of early-stage eggs

In some fish species with exclusive paternal care females prefer to deposit their eggs in nests containing early-stage eggs and avoid those containing only late-stage eggs. Such female preference may also be shown when it comes to deciding where to lay within a given nest; that is, females may prefer to lay contiguously to younger rather than to older eggs. To understand if females benefit from such preferences by obtaining higher offspring survival, we conducted field studies with the bluefin damselfish, Abudefduf luridus. We investigated (1) female between-nest and within-nest spawning-site preferences relative to brood characters and (2) egg-hatching success related to male filial cannibalism. We found evidence that males entirely ate some of their clutches, especially the eggs received near the end of the breeding season. These were also more susceptible to being eaten when laid in nests containing only eggs in a late stage of development. Theoretically, these patterns of clutch loss should result in adaptive female preference for nests containing early-stage eggs, but we found no evidence for such a preference in female A. luridus. However, once females had chosen a nest they strongly preferred to lay contiguously to early-stage eggs than to late-stage eggs. Moreover, females benefited from this decision by obtaining higher survival rates for their clutches. We conclude that within-nest spawning-site preference of the female for early-stage eggs may have evolved as an adaptive response to a dilution effect on the risk of egg loss.     

Lack of costs associated with nest‐related behaviors in an arachnid with exclusive paternal care

Nest‐related behaviors may benefit males by increasing offspring survival and their attractiveness to females, but may also limit males’ foraging activity, increase their metabolic expenses, and expose them to increased mortality during nest attendance. Although intensively studied among birds and ectothermic vertebrates, the costs of nest‐related behaviors in arthropods remain poorly explored. Females of the Neotropical harvestman Zygopachylus albomarginis (Arachnida: Opiliones) lay eggs exclusively inside mud nests that are built, repaired, cleaned and defended by males, which may remain stationary and associated with the nest for up to five months. To assess energetic and survival costs of nest‐related behaviors in this arthropod species, we measured body condition of nesting and non‐nesting males and conducted a field capture‐mark‐recapture study to estimate their survival rates. Despite the long period of nest attendance, nesting males sustained good body conditions and presented higher survival rates than non‐nesting males and females. Two ecological conditions may play an important role modulating the costs of nest attendance in the species. First, high food supply in tropical rainforests may provide males with frequent access to food in the vicinity of their nests, reducing or eliminating the costs related to limited foraging opportunities. Second, predation pressure seems to be directly mostly to vagrant individuals, so that the more they move, the more likely they are to be singled out by predators. Taken together, our findings indicate that nest and offspring defense in Z. albomarginis provide numerous benefits, surprisingly imposing no evident cost to the males.     

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.     

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.     

Silk threads function as an 'adhesive cleaner' for nest space in a social spider mite

Individuals of the social spider mite Stigmaeopsis longus live communally in narrow, humid nests made from silk threads and display nest sanitation behaviour through the coordinated deposition of faeces. We used artificial dust to experimentally determine that females of this species use silk threads to perform regular cleaning of the nest space and eggs. We first learned that silk-weaving behaviour is not a by-product of nest building (nest reinforcement); rather, it is actively performed as a function of cleaning the living space and eggs. Furthermore, we determined the effectiveness of the attending females by artificially manipulating their natural habitat, which is clearly connected to the cleaning behaviour by parental females. As such, we have uncovered an extraordinary new role of silk threads as devices for cleaning the nest space and/or eggs. These results strongly indicate that special adaptations for maintaining clean habitats are essential for animals to evolve aggregative social lives.