Threat-sensitive anti-predator defence in precocial wader,the northern lapwing <Emphasis Type="Italic">Vanellus vanellus</Emphasis>

Birds exhibit various forms of anti-predator behaviours to avoid reproductive failure, with mobbing—observation, approach and usually harassment of a predator—being one of the most commonly observed. Here, we investigate patterns of temporal variation in the mobbing response exhibited by a precocial species, the northern lapwing (Vanellus vanellus). We test whether brood age and self-reliance, or the perceived risk posed by various predators, affect mobbing response of lapwings. We quantified aggressive interactions between lapwings and their natural avian predators and used generalized additive models to test how timing and predator species identity are related to the mobbing response of lapwings. Lapwings diversified mobbing response within the breeding season and depending on predator species. Raven Corvus corax, hooded crow Corvus cornix and harriers evoked the strongest response, while common buzzard Buteo buteo, white stork Ciconia ciconia, black-headed gull Chroicocephalus ridibundus and rook Corvus frugilegus were less frequently attacked. Lapwings increased their mobbing response against raven, common buzzard, white stork and rook throughout the breeding season, while defence against hooded crow, harriers and black-headed gull did not exhibit clear temporal patterns. Mobbing behaviour of lapwings apparently constitutes a flexible anti-predator strategy. The anti-predator response depends on predator species, which may suggest that lapwings distinguish between predator types and match mobbing response to the perceived hazard at different stages of the breeding cycle. We conclude that a single species may exhibit various patterns of temporal variation in anti-predator defence, which may correspond with various hypotheses derived from parental investment theory.     

Larval life history and anti-predator strategies are affected by breeding phenology in an amphibian

Seasonal time constraints can pose strong selection on life histories. Time-constrained animals should prioritise fast development over predation risk to avoid unfavourable growing conditions. However, changes in phenology could alter the balance between anti-predator and developmental needs. We studied variation of anti-predator strategies in common frog (Rana temporaria) tadpoles in four populations from the two extremes of a latitudinal gradient across Sweden. We examined, under common conditions in the laboratory, the anti-predator responses and life histories of tadpoles raised with predatory Aeshna dragonfly larvae in two consecutive years with a difference of 20 days in breeding time in the north, but no difference in breeding time in the nouth. In a year with late breeding, northern tadpoles did not modify their behaviour and morphology in the presence of predators, and metamorphosed faster and smaller than tadpoles born in a year with early breeding. In the year with early breeding, northern tadpoles showed a completely different anti-predator strategy by reducing activity and developing morphological defences in the presence of predators. We discuss the possible mechanisms that could activate these responses (likely a form of environmentally-mediated parental effect). To our knowledge, this is the first study to show that a vertebrate modifies the anti-predator strategy of its offspring in response to natural variation in reproductive phenology, which highlights the need to consider phenology in studies of life-history evolution.     

Species interactions in a mixed colony of common terns (Sterna hirundo) and black skimmers (Rynchops niger)

Black skimmers (Rynchops niger) frequently nest among common terns (Sterna hirundo) on coastal islands from North Carolina to New York. Behavioural interactions among terns, skimmers, and several potential avian predator species were quantified during the 1973 breeding season at Fisherman Island, Virginia. Common terns were dominant to skimmers, chasing and dive-attacking neighbouring pairs, while skimmers rarely chased terns. Despite agonistic interactions, nesting success of both species was no different whether nearest neighbours were conspecifics or heterospecifics. Common terns chased away potential avian predators much more frequently than did skimmers. Black skimmers, then, appear to derive a ‘protective cover’ benefit from nesting among the terns. Neither nest-site limitation nor food ‘information-sharing’ appear to account for the species associations.     

Functional responses of the rough-legged buzzard in a multi-prey system

The functional response is a key element of predator–prey interactions. Basic functional response theory explains foraging behavior of individual predators, but many empirical studies of free-ranging predators have estimated functional responses by using population-averaged data. We used a novel approach to investigate functional responses of an avian predator (the rough legged-buzzard Buteo lagopus Pontoppidan, 1763) to intra-annual spatial variation in rodent density in subarctic Sweden, using breeding pairs as the sampling unit. The rough-legged buzzards responded functionally to Norwegian lemmings (Lemmus lemmus L. 1758), grey-sided voles (Myodes rufocanus Sundevall, 1846) and field voles (Microtus agrestis L. 1761), but different rodent prey were not utilised according to relative abundance. The functional response to Norwegian lemmings was a steep type II curve and a more shallow type III response to grey-sided voles. The different shapes of these two functional responses were likely due to combined effects of differences between lemmings and grey-sided voles in habitat utilisation, anti-predator behaviour and size-dependent vulnerability to predation. Diet composition changed less than changes in relative prey abundance, indicating negative switching, with high disproportional use of especially lemmings at low relative densities. Our results suggest that lemmings and voles should be treated separately in future empirical and theoretical studies in order to better understand the role of predation in this study system.     

The exclusion of avian predators from aggregations of nesting lapwings (Vanellus vanellus)

Territorial lapwings in Aberdeenshire, Scotland, largely prevented carrion crows, (Corvus corone), their main egg predators, from approaching close to nest-sites and from entering the area within an aggregation of 11 nests. Predation rates of artifical nests were significantly lower when placed within 10 m of active lapwing nests than when>200 m from nest-sites, indicating that the presence of lapwings afforded protection from crow predation. Further artificial nest experiments showed that the amount of protection declined linearly with increasing distance from the nest-site, and some degree of protection occurred to at least 30–50 m from it. The size of this protected zone was apparently related to the number of lapwing nests present, and the survival time of eggs in artificial nests reflected the lapwings' nesting density. The defended zones around each lapwing nest overlapped appreciably in dense or large nesting aggregations, leading to the observed protection by generally excluding crows, from the nesting area.     

Competition, predation and nest niche shifts among tropical cavity nesters: phylogeny and natural history evolution of parrots (Psittaciformes) and trogons (Trogoniformes)

Nest site selection by birds is a critically important life history trait as competition for suitable sites can be intense, and because birds are at their most vulnerable to predators during nesting. Previous studies show that the clutch size and nestling period evolve in response to competition for nest sites and nest predation, respectively. This provides the opportunity to study the relative contribution of competition and predation to the evolution of nesting niche. Using previously published phylogenies for parrots and trogons, I found evidence for at least 13 independent evolutionary transitions from tree cavities to alternative nesting niches (including termitaria, cliffs, and burrows). I analyzed variations in clutch size, incubation period and nestling period for 16 phylogenetically controlled pairs of species to test the relative roles of competition for tree cavities and nest predation, in favoring evolutionary switches to alternative nest sites. Tree cavity nesting species did not have larger clutch sizes as predicted if competition for tree cavities leads birds to invest heavily in nesting once they obtain a nest site (the limited breeding opportunities hypothesis). Instead I found that shifts to alternative nesting niches were accompanied by an increase in nestling period. As nestling period is a surrogate measure for long-term nest predation rates, this finding suggests that nest predation has been more important than competition in niche diversification among cavity nesting parrots and trogons. The timing of events in South America suggests that the explosive radiation of mammalian nest predators during the Upper-Oligocene, Lower-Miocene (20–30 million years ago) corresponded with the radiation of parrot and trogon taxa that exploit novel nesting niches.     

Aggression of breeding eastern bluebirds (Sialia sialis) toward their mates and models of intra-and interspecific intruders

Hypotheses regarding the evolution of aggressive patterns during the breeding season in eastern bluebirds (Sialia sialis) were tested by placing models of eastern bluebirds and house sparrows (Passer domesticus) into territories of experimentally naive breeding pairs of eastern bluebirds during three stages of the nesting cycle. The responses of the resident males to models of male bluebirds decreased from the egg-laying to the nestling stage, and this response pattern can be interpreted as support for the protection-of-parentage hypothesis or the nest-site-defence hypothesis. The aggression of resident males to resident females during trials with models of male bluebirds weakly conforms to the pattern predicted by the protection-of-parentage hypothesis. The aggressive response to models of house sparrows also decreased from the egg-laying to the nestling phase in a manner consistent with the nest-site-defence explanation. No aggression toward mates by males or females was observed during trials with house sparrow models, an observation consistent with the predictions of the protection-of-parentage hypothesis.     

Island accessibility and distance from beach influence nesting success of Sooty Falcons Falco concolor in Oman

Colonial island‐breeding birds can be particularly vulnerable to anthropogenic disturbance, which can adversely affect their nesting success. We studied Sooty Falcons Falco concolor breeding on 10 ground‐predator‐free islands in the Sea of Oman during 2007–2014 and evaluated spatio‐temporal trends in the number of breeding pairs occurring on the islands and the factors influencing nesting success. The number of breeding pairs on the islands declined during the study, due mostly to the decline on accessible islands; the rate of decline on islands accessible to humans was double that on inaccessible ones. The number of nests with one or more eggs declined during the study period, and the percentage of nests with eggs that produced one or more chicks showed an increasing trend over time. Sooty Falcon nests located farther away from beaches experienced a significantly higher probability of nesting success than those located closer to beaches. Our results suggest that the number of breeding Sooty Falcons on the islands of northern Oman is declining and that human disturbance may be a contributing factor; this probably mirrors the situation in other parts of the breeding range of this species.     

Predation risk and inter-population variation in antipredator behaviour in the three-spined stickleback, Gasterosteus aculeatus L.

Predation risk from fish and bird predators was assessed at seven Scottish Gasterosteus aculeatus L. sites. Samples of adult male and female sticklebacks and fry from each site were tested with either a model heron or a live pike to measure anti-predator responses. Principal Component Analyses were then used to describe the responses, the first factor to emerge from each multivariate analysis providing an index of overall level of response toward the predator. Significant sexual and inter-population differences in behaviour are described. Sticklebacks from populations at high risk from either predatory fish or birds showed higher fright response scores than fish from low risk sites. In general, adult male sticklebacks are bolder than adult females during interactions with predators.     

Sex differences in nest defence by the red-backed shrike<Emphasis Type="Italic"> Lanius collurio</Emphasis>: effects of offspring age,brood size,and stage of breeding season

The red-backed shrike Lanius collurio is an open-cup nesting passerine bird, which protects its nest aggressively attacking predators near the nest. The response of parents to a human observer was investigated during the nesting period 1999–2001 in Poland. The intensity of nest defence increased as the breeding cycle progressed in accordance to the offspring value hypothesis. However, we did not find an increase in the level of aggression according to the number of offspring and the time of season. Contrary to previous predictions, we did not find gender differences in nest defence. More aggressive parents, both females and males, have significantly better breeding success than quieter individuals.     

Behavioral responses of native prey to disparate predators: naiveté and predator recognition

It is widely accepted that predator recognition and avoidance are important behaviors in allowing prey to mitigate the impacts of their predators. However, while prey species generally develop anti-predator behaviors through coevolution with predators, they sometimes show accelerated adoption of these behaviors under strong selection pressure from novel species. We used a field manipulation experiment to gauge the ability of the common ringtail possum (Pseudocheirus peregrinus), a semi-arboreal Australian marsupial, to recognize and respond to olfactory cues of different predator archetypes. We predicted that ringtails would display stronger anti-predator behaviors to cues of the invasive European red fox (Vulpes vulpes) in areas where fox impacts had been greatest, and to cues of the native lace monitor (Varanus varius) in areas of sympatry compared with allopatry. We found that ringtails fled quickly and were more alert when exposed to the fecal odors of both predators compared to neutral and pungent control odors, confirming that predator odors are recognized and avoided. However, these aversive responses were similar irrespective of predator presence or level of impact. These results suggest that selection pressure from the fox has been sufficient for ringtails to develop anti-predator behaviors over the few generations since foxes have become established. In contrast, we speculate that aversive responses by ringtails to the lace monitor in areas where this predator is absent reflect recent coexistence of the two species. We conclude that rapid evolution of anti-predator behaviors may occur when selection is strong. The maintenance of these behaviors should allow re-establishment of predator–prey relationships if the interactants regain sympatry via range shifts or management actions to reintroduce them to their former ranges.     

Enemy Recognition of Reed Warblers (Acrocephalus scirpaceus): Threats and Reproductive Value Act Independently in Nest Defence Modulation

Organisms should respond more aggressively towards species perceived as a danger to their offspring, but intensity of defence may be gauged by the value of current offspring weighed against the value of future reproductive opportunities. We tested whether defensive responses of nesting reed warblers (Acrocephalus scirpaceus) are the result of an interaction effect between the type of stimulus confronted and the value of the warbler’s nesting attempt. We quantified the ability of reed warblers to discriminate among brood parasites, nestling predators and non‐threatening species at different stages of the breeding cycle. We also determined whether variables that influence the value of offspring, such as time of season, size and age of clutch or brood, and time of day and number of visits to the nest, explain variation in the intensity of defence recorded during the egg and nestling stages. Responses to the three stimuli differed significantly, as reed warblers consistently directed their mobbing calls and attacks towards parasites, whereas they were less conspicuous when confronted with models of predators. Reed warblers modulated their responses towards each stimulus in accordance with the threat each posed at a specific nesting stage, whereas they were not affected by other variables relative to their reproductive potential. The churr call, however, was uttered independently of the stimulus, as it was triggered by the mere presence of nestlings in the nest.     

Activity budgets of a marked common loon (Gavia immer) nesting population

A newly devised nightlighting technique was used to capture breeding adult common loons (Gavia immer) at the Seney National Wildlife Refuge in northern Michigan in 1989. The behaviors of 6 pairs of known-sex, color-marked common loons were subsequently quantified during the breeding cycle in 1990. Collected observational data indicate that foraging, resting, locomotion, and preening were frequent throughout the breeding cycle. Time spent foraging was greatest during the pre-nesting period (53 to 57%), but declined significantly during the nesting and post-nesting periods (p<0.05). Time spent foraging during the pre-nesting period was similar to that of fall and winter studies. During the pre-nesting period adult loons spent about 15% of the time in locomotion; this was significantly greater than the other time periods (p<0.05) and is attributed to selecting a nest site. During the nesting cycle, almost half of each bird's time spent was nest-sitting. Sexual differences were negligible during nest-sitting. Resting and chick-rearing were the predominant behaviors during the post-nesting period and were responsible for the biggest difference in parental duties. Time spent preening declined from 8% during the pre-nesting period to 4 to 5% during the post-nesting period. Time spent by nesting pairs to produce chicks is approximately 10% during pre-nesting, 48 to 49% during nesting, and between 38 to 44% during post-nesting. By quantifying and establishing behavioral standards, subtle abnormalities or changes can be detected to better manage for viable common loon populations.     

Effect of predator exposure on the performance and survival of red drum (<Emphasis Type="Italic">Sciaenops ocellatus</Emphasis>)

The influence of predator exposure on the survival of hatchery red drum, Sciaenops ocellatus was investigated under laboratory conditions. Several prey-capture (attack distance, mean attack velocity, capture time, and gape cycle duration) and anti-predator (reaction distance, response distance, mean velocity, and maximum velocity) performance variables were quantified using high-speed video for juvenile red drum (25–30 mm standard length, L _S ) reared with and without predators (pinfish, Lagodon rhomboides). Univariate contrasts of prey-capture events demonstrated that attack distance (mean ± s.e.) was significantly greater in red drum reared with (1.20 ± 0.16 mm) versus without (0.65 ± 0.09 mm) pinfish predators. During anti-predator events, red drum reared with predators had approximately 300% greater reaction distance and 20%–30% greater response distance, mean velocity and maximum velocity versus fish reared without predators; however, these differences were not statistically significant. Following video assessments, a series of mortality experiments were conducted using free-ranging pinfish predators. Mortality rates (Z) ranged from 0.047 to 0.060 (Z/hr/predator) and did not differ significantly among treatments.     

Alarm calls of the Southern House Wren Troglodytes musculus: variation with nesting stage and predator model

Alarm calls given by parents when risk is detected during nesting may be considered a form of parental defense. We analyzed variations in callings of breeding pairs of the Southern House Wren Troglodytes musculus during the nesting cycle and when faced with different predator models. Nesting birds were exposed to stuffed models at different nesting stages (early and late during incubation, and nests with younger and older nestlings). Nests were also exposed to different predator models where the calling response of breeding adults and acoustic structure variations of the calls were analyzed. The presence of a predator model increased the parents’ alarm calls along the nesting stage. This result supports the hypothesis that the higher the nest reproductive value, the higher the nest defense performed by the Southern House Wren. However, it also supports the notion that alarm calls could be used by parents to silence nestlings and reduce their detectability. Alarm calls also varied according to the predator model presented. We suggest that alarm calls variations of Southern House Wrens could encode information about the kind of predator and the risk envisaged through variations of call rates.     

Habitat preference, breeding success and the effect of primary productivity on Tawny Eagles Aquila rapax in the tropics

Breeding success of 176 Tawny Eagle pairs nesting on basalt (4724 km²– 92 pairs) and kalahari sand (9876 km²– 84 pairs) was monitored from 1973 to 1984 in Hwange National Park, Zimbabwe. Flat-topped trees were preferred for nesting but kalahari sand pairs used round-crowned trees more often, probably due to tree availability. Mean inter-nest distances on both biomes were significantly different (3–5 km on basalt, 59 km on kalahari sand). Clutch size and frequency of one-egg clutches in both biomes were similar and many more one-egg clutches were laid in the drier period of the study (1979–1984). There was no correlation between laying date and breeding success. More basalt pairs laid synchronously, probably due to the smaller inter-nest distances. Breeding success in both biomes was similar and significantly more chicks were reared during 1973–1978 than during 1979–1984 (wet and dry periods, respectively). Some districts produced more chicks than others, suggesting good and marginal areas within both biomes. The greater inter-nest distances and ability to breed successfully in round-crowned trees by kalahari sand Tawny Eagles resulted in a similar number of chicks reared per breeding attempt as basalt pairs, in spite of the lower primary production of the sand area.     

Gonadal and hormonal characteristics of captive adult mourning doves during the nesting cycle

Paired mourning doves ($\text{Zenaida}$$\text{macroura}$) of a captive breeding colony were selected and sacrificed on specific day intervals postlaying and posthatching during 1976–1977. Blood plasma samples were analyzed for androgens, estradiol, and progestins using radioimmunoassays. Testes, ovary and oviduct weights, spermatozoan numbers, diameter of the three largest follicles in the ovary, the presence or absence of eggs in the reproductive tract, and crop gland characteristics also were recorded. Differences (P<0.05) existed between nonbreeding and breeding pairs in testes weights, spermatozoan numbers, and ovary and oviduct weights. No distinct cyclic pattern was observed in gonadal variables among breeding males and females during the nesting cycle, although trends were observed. No differences (P > 0.05) in androgen and progestin concentrations in males, or estradiol and progestin concentrations in females, between nonbreeding and breeding pairs, or among breeding pairs during different stages of the nesting cycle, were observed. Analysis of reproductive variables by stage of crop gland activity tended to indicate an inverse relationship between gonadal and crop gland activity in both male and female doves during the nesting cycle.     

Numerical responses of an avian predator to prey fluctuations in a temperate latitude: breeders vs. entire population

Distinct numerical responses of predators to fluctuations in the abundance of their prey are often observed in northern regions but occur more rarely in temperate latitudes. This statement is, however, mostly based on observations of breeding populations, while in some predators, for example in raptors, numerous non-breeding floaters can occur. I estimated the breeding density and reproductive performance (nest survey) as well as the density of entire population (transects with distance sampling) of the common buzzard Buteo buteo in western Poland (52°N) in the years 2005–2014 to test the hypotheses that in temperate latitudes: (1) the breeding population of these birds does not show any numerical response, understood as annual changes in their abundance; (2) its reproductive success, however, changes with the abundance of main prey, the common vole Microtus arvalis; and (3) hence the entire buzzard population (including potential immature floaters) present in a given area during the nesting period responds numerically with some time delay. The reproductive success of buzzards was positively correlated with their prey abundance. Contrary to my predictions, however, the breeding population of buzzards showed a slight numerical response with a 3-year lag and the entire population tracked vole fluctuations without any time delay. The immediate numerical response of the entire buzzard population was probably caused by large-scale movements of floaters. Such rapid numerical responses of some predators may contribute to the relative stability of prey populations in temperate latitudes compared to northern regions.     

Reduction in predator defense in the presence of neighbors in a colonial fish

Predation pressure has long been considered a leading explanation of colonies, where close neighbors may reduce predation via dilution, alarming or group predator attacks. Attacking predators may be costly in terms of energy and survival, leading to the question of how neighbors contribute to predator deterrence in relationship to each other. Two hypotheses explaining the relative efforts made by neighbors are byproduct-mutualism, which occurs when breeders inadvertently attack predators by defending their nests, and reciprocity, which occurs when breeders deliberately exchange predator defense efforts with neighbors. Most studies investigating group nest defense have been performed with birds. However, colonial fish may constitute a more practical model system for an experimental approach because of the greater ability of researchers to manipulate their environment. We investigated in the colonial fish, Neolamprologus caudopunctatus, whether prospecting pairs preferred to breed near conspecifics or solitarily, and how breeders invested in anti-predator defense in relation to neighbors. In a simple choice test, prospecting pairs selected breeding sites close to neighbors versus a solitary site. Predators were then sequentially presented to the newly established test pairs, the previously established stimulus pairs or in between the two pairs. Test pairs attacked the predator eight times more frequently when they were presented on their non-neighbor side compared to between the two breeding sites, where stimulus pairs maintained high attack rates. Thus, by joining an established pair, test pairs were able to reduce their anti-predator efforts near neighbors, at no apparent cost to the stimulus pairs. These findings are unlikely to be explained by reciprocity or byproduct-mutualism. Our results instead suggest a commensal relationship in which new pairs exploit the high anti-predator efforts of established pairs, which invest similarly with or without neighbors. Further studies are needed to determine the scope of commensalism as an anti-predator strategy in colonial animals.     

Abundance and spatial distribution of sympatrically breeding Catharacta spp. (skuas) in Admiralty Bay, King George Island, Antarctica

We examined the abundance and spatial distribution of sympatrically breeding skuas (Catharacta spp.) within Admiralty Bay, King George Island, Antarctica, during the austral summer of 2004/2005 in relation to spatial variables, which correspond to access to resources and nesting site safety and quality. We also compared the distribution and abundance of skua pairs observed in 2004/2005 to published skua census data from 1978/1979. Similar to previous studies, we found that brown skua (C. antarctica lonnbergi) pairs often nested in close proximity to penguin colonies and actively excluded other pair types from having direct access to penguin resources. In areas directly around penguin colonies, brown skua displace south polar skua (C. maccormicki) and other pair types, indirectly forcing them to nest in possibly lower quality territories, which are farther away from the coastline and in areas with lower incident solar radiation. When examining skua population trends, we discovered that the total number of breeding skuas in Admiralty Bay had increased by 293%, from 128 to 468 pairs, since 1978/1979. This dramatic increase was driven primarily by a tenfold increase in south polar skua pairs, as well as smaller increases in mixed and hybrid pairs. In contrast, there has been an overall decline (by 40%) in brown skua pairs during this same time, driven primarily by a large decrease in the breeding density of brown skua pairs in areas without penguin colonies.