Early is better: seasonal egg fitness and timing of reproduction in a zooplankton life-history model

Timing of reproduction influences future prospects of offspring and therefore the reproductive value of parents. Early offspring are often more valuable than later ones when food availability and predation risk fluctuate seasonally. Marine zooplankton have evolved a diversity of life history strategies in response to seasonality. We present a state-dependent life history model for the annual and herbivorous high-latitude copepod Calanoides acutus . Individuals are characterised by four states; developmental stage, structural size, energy reserves and vertical location. There are two habitats, a surface habitat with seasonal predation risk and food availability, and a safer deep habitat with no food and low metabolism (diapause). Optimal life histories (diapause and energy allocation strategies) are found by dynamic programming. Seasonal egg fitness (reproductive value) emerges from the model and peak values are typically before the feeding season. Disentangling the fitness components, we conclude that seasonality in egg fitness is caused both by environmental seasonality in food and predation risk and by time-constraints on development and diapause preparation. Realised egg production, as predicted from population simulations, does not match the seasonal peak in offspring fitness but is delayed relative to peak egg fitness. We term this an 'internal life history mismatch' as constraints and tradeoffs cause sub-optimal birth dates for most eggs whereas mothers maximise their reproductive value by high number of eggs rather than few and optimally timed eggs. The earliest eggs have a disproportionately high contribution to population recruitment, emphasising the importance of early eggs and the need to understand seasonal patterns in offspring fitness.     

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.     

Assessment of the consumption of walleye pollock eggs by nekton and jellyfish in the northern Sea of Okhotsk in the spring

In the northern Sea of Okhotsk, nekton and jellyfish consumed as many as 831 × 10⁹ walleye pollock eggs per day in 2011. The nekton exerted the highest pressure, viz., 98.3% of the overall predation on pollock egg by aquatic animals. Of the entire quantity of consumed eggs, 55.9% were eaten by herring, 35.9% by walleye pollock, 6.5% by Sakhalin sole, and 1.7% by jellyfish. Among jellyfish, scyphomedusae Cyanea capillata and Chrysaora melonaster, as well as the hydromedusa Tima sachalinensis consumed the largest quantities of eggs. The total consumption of pollock egg by aquatic animals in 2011 was estimated at 42.4 × 10¹², or 11.4% of the entire quantity of eggs that were spawned by walleye pollock in the waters of the northern part of the sea. The total amount of pollock eggs that were eaten by herring and pollock together for 51 days in 2011 amounted to 38.9 × 10¹², which was 5.7 times as much as that in 2002. Thus, a significant growth of predation on pollock eggs by their main consumers, viz., herring and walleye pollock, was observed in 2011. This was caused by an increase in the populations of both species during the recent years and also by a higher concentration of pollock eggs.     

Parental care of a cowbird host: caught between the costs of egg-removal and nest predation

Avian brood parasites reduce host fitness through the addition of parasitic eggs and the removal of host eggs. Both parasitic egg-addition and host egg-removal may be important sources of selection on host behaviour, creating fitness trade-offs with selection imposed by nest predation. However, the relative costs hosts suffer from egg-addition and host egg-removal and the responses to these costs are largely unstudied. Through experimental manipulations and observations, we demonstrate that increased nest attentiveness by female yellow warblers (Dendroica petechia) reduces the cost of brood parasitism by reducing egg-removal by brown-headed cowbirds (Molothrus ater). However, female attentiveness does not reduce the addition of parasitic eggs. Experimentally parasitized females respond to the threat of egg-removal by increasing nest attentiveness. Increased attentiveness, however, reduces time for females to gather food and requires males to visit the nest more often to feed incubating females. This increased activity in turn increases the risk of nest predation. Thus, brood parasitism (specifically egg-removal) and nest predation produce conflicting selection on incubation strategies, as parasitized hosts are caught between the costs of egg-removal by brood parasites, and the costs of increased nest predation if the female spends more time on the nest to reduce egg-removal.     

Do shark declines create fear‐released systems?

We illustrate the theoretical plausibility that fishery removal of sharks can indirectly alter predation pressure on different fish species via the behavioural responses of mesoconsumers released from predator intimidation. Our dynamic state variable model of foraging decisions by harbour seals, a mesopredator, predicted indirect effects of the removal of Pacific sleeper sharks on two species consumed by seals, Pacific herring and walleye pollock, as mediated by seal behaviour. Herring, a fatty fish, form near‐surface aggregations that often are ephemeral and widely dispersed. Pollock are found in the deeper strata preferred by sharks and have lower energy density than herring, but also are larger and their more continuous distribution potentially makes them the more predictable resource for seals. During simulations, predation risk from sharks produced an asymmetric trophic cascade mediated by the seal's underutilisation of deeper, riskier strata. Risk management by seals reduced mortality on pollock, which required riskier access in deep strata, while increasing mortality on herring, which could be accessed with less risk in shallow strata. This effect, however, attenuated if herring were scarcer and seal energy state was poor. During shark removal scenarios, seals shifted to deeper strata, increasing pollock consumption and substantially decreasing use of herring; the proportional change was greatest if seals were in good energy state. Prior studies have addressed how shark declines might affect community structure through density responses by species consumed by sharks; earlier models incorporating antipredator behaviour of the mesopredator (i.e. Ecosim/Ecospace) allow for activity reduction but not for the spatial shifts that altered the asymmetric trophic cascade in our simulations. Our results suggest that shark declines might have stronger ecological consequences than previously recognised if we account for spatial and diet shifts by mesoconsumers released from shark intimidation.     

The power of the seasons: rainfall triggers parental care in poison frogs

The quality of breeding sites is of great importance for the reproductive success and accordingly the fitness of many animal species. Hence, individuals should decide carefully where to rear their offspring. Often parents have to account for multiple characteristics of habitat quality at once, which in turn might change over time. Specimens confronted with such variability may evolve the ability to display context-dependant decision plasticity. Anuran amphibians breeding in ephemeral pools largely face two risks for their offspring: desiccation and predation. The Neotropical poison frog Ranitomeya variabilis deposits both eggs and tadpoles in phytotelmata. These small tadpole nurseries lower the risk of offspring predation. However, because most poison frog tadpoles are cannibalistic, even these pools need to be surveyed for predators, and parents tend to avoid deposition with conspecifics. We tested if this avoidance behaviour does change in parental R. variabilis depending on seasonal circumstances. Over several months we provided the frogs the option to deposit their eggs or tadpoles in pools that did and did not contain chemical cues of cannibalistic conspecifics, respectively. During the rainy season, frogs strongly avoided conspecific cues for both eggs and tadpoles. Anyway, with the change to the dry season, parental preferences changed such that parent frogs were more likely to deposit tadpoles (but not eggs) in pools containing cues of conspecific tadpoles. We suggest that R. variabilis, a species that typically isolates its cannibalistic offspring, has evolved a plastic feeding behaviour with regard to the risk of phytotelmata desiccation. We interpret that parents provide older tadpoles with younger ‘trophic’ tadpoles in order to accelerate their development and save them from impending desiccation.     

Time-dependent reproductive decisions in the blue tit

Many breeding attempts in birds do not result in any fledged young due to predation on eggs or young. Consequently, the influence of time constraints on reproductive decisions are integrated parts of the reproductive behaviour of birds breeding within short, seasonal climate zones. In this study, I mimicked nest predation by removing blue tit (Parus caeruleus) clutches shortly after completion. Around 75% of the removed clutches were followed by a repeat clutch. Females producing their first clutch early in the season and females with an early onset of incubation in the laying sequence (an indication of high parental or territory quality) were most likely to initiate a repeat clutch. A trade‐off between the benefits of a repeat clutch and survival likely stopped late females in bad condition from investing more in the current reproductive season. Females producing a repeat clutch laid fewer eggs, had an earlier onset of incubation in the laying sequence and produced larger eggs than they did when producing their original first clutch. Eggs produced after the onset of incubation were especially large in the repeat clutches. Since food availability was presumably higher when the female produced her repeat clutch compared with her first clutch, females made a strategical decision when reducing clutch size, whereas onset of incubation and egg size may have been energetically constrained when producing the first clutch. Females that produced a relatively large clutch, had a relatively early onset of incubation, and laid relatively large eggs in their first clutch also did so when producing a repeat clutch, indicating that some of the variation in breeding parameters are due to differences in parental or territory quality. Differences between years in the temperature‐dependent development rate of caterpillars seem to affect the time constraints on breeding. A year with a predicted early seasonal decline in caterpillars resulted in short intervals between removal and relaying, small clutches and an early onset of incubation.     

Nest desertion: a trade-off between current and future reproduction

Life history theory predicts that parents should desert a reproductive attempt if the costs of rearing this brood exceed the expected benefits. Thus, if the value of the current breeding attempt is reduced, for example through an unexpected reduction in size, parents are expected to reconsider whether it is worth continuing investing in their brood. With regard to nest desertion behaviour two predictions can be made: individuals are (1) more likely to desert if the reduction in clutch size is large and (2) less likely to desert if the reduction is at a late stage of breeding. We investigated the threshold at which nest desertion takes place by experimentally reducing great tit, Parus major, clutches to different sizes and at different stages of the incubation period. The results were in accordance with the predictions: clutch desertion rates were negatively related to the number of eggs that remained in the nest, and nest desertion was less likely nearer the end of the incubation period. In addition, we estimated the fitness consequences of nest desertion behaviour. For this purpose we made one group of birds desert in favour of a replacement clutch and another group rear a reduced brood. The latter were more likely to produce a second clutch after the first-brood fledglings had left the nest. As a consequence, the number of fledglings produced over the entire breeding season did not differ between the two experimental groups. We also counted the number of recruits and breeding adults in the following breeding season and found that the experimental groups did not differ in local recruitment and adult survival. Therefore, the results did not indicate that parents improved their fitness by deserting a reduced clutch.Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved .     

Assessing predation risk: optimal behaviour and rules of thumb

We look at a simple model in which an animal makes behavioural decisions over time in an environment in which all parameters are known to the animal except predation risk. In the model there is a trade-off between gaining information about predation risk and anti-predator behaviour. All predator attacks lead to death for the prey, so that the prey learns about predation risk by virtue of the fact that it is still alive. We show that it is not usually optimal to behave as if the current unbiased estimate of the predation risk is its true value. We consider two different ways to model reproduction; in the first scenario the animal reproduces throughout its life until it dies, and in the second scenario expected reproductive success depends on the level of energy reserves the animal has gained by some point in time. For both of these scenarios we find results on the form of the optimal strategy and give numerical examples which compare optimal behaviour with behaviour under simple rules of thumb. The numerical examples suggest that the value of the optimal strategy over the rules of thumb is greatest when there is little current information about predation risk, learning is not too costly in terms of predation, and it is energetically advantageous to learn about predation. We find that for the model and parameters investigated, a very simple rule of thumb such as 'use the best constant control' performs well.     

Does the cost of incubation set limits to clutch size in common eiders Somateria mollissima?

We examined the effect of natural clutch size on the cost of incubation in a population of common eiders Somateria mollissima nesting in Tromsø, northern Norway. The body condition of females at day 5 in the incubation period was not related to clutch size (3–6 eggs), but females incubating large clutches lost more mass and had a lower body condition at day 20 in the incubation period than females incubating small clutches. Females incubating large clutches had a slightly shorter incubation period and a lower egg predation rate. The results do not support the hypothesis that the female's ability to produce eggs is the only ultimate control of clutch size in eider. Instead the results suggest that there may be an interaction between the allocation of body reserves to eggs and incubation, and that females producing large clutches allocate more of their body reserves to incubation than females producing small clutches, in order to shorten the incubation period and to minimise the risk of predation on eggs.     

Size and visibility of Baltic cod eggs with reference to size-selective and stage dependent predation mortality

Size of Baltic cod eggs from incubation experiments and from field samples was determined by microscopic analysis. Results from plankton samples were compared with corresponding size distributions of cod eggs found in herring stomachs. The influence of fixation on size of different developmental stages was studied. Live eggs from incubation experiments were also sized repeatedly throughout the developmental period with an optical plankton counter (OPC) based on light attenuance measurements as this was assumed to be more closely related to the visibility of the eggs for potential predators than egg diameter as obtained by microscopic analysis.
Preservation in formaldehyde solution caused a small reduction in egg diameter (2.2%) whereby no differences between the developmental stages were detected. Egg size decreased slightly during incubation (6.9%) while the OPC measurements revealed a substantial increase in light attenuance during egg development (42.2%). In the field, a general decrease in egg size with increasing depth was observed while no change between the developmental stages was detectable. The mean size of eggs ingested by herring was slightly lower than in the water column which was most pronounced for the late stages containing a well-developed embryo. The frequency of eggs in an advanced stage of development was considerably higher in the stomachs than in corresponding plankton samples. Therefore, it is suggested that the selection of further developed egg stages by predatory fish in the central Baltic Sea, i.e. herring and sprat, is due to an increase of visibility during egg development in relation to growth and pigmentation of the embryo. Thus it is likely that egg mortality due to predation is stage-dependent rather than strictly size-selective.     

Vulnerability of late larval and early juvenile Atlantic herring, Clupea harengus, to predation by whiting,Merlangius merlangus

Laboratory experiments investigated changes in the vulnerability of herring to predation by whiting during growth from <34 to >80mm in length, Premetamorphic herring (<50mm) failed to react to c . 50% of attacks by the predator, but this was reduced to c . 20% in postmetamorphic fish. Premetamorphic herring failing to react had c. 30% probability of survival due to unforced errors by the predator (unsuccessful attacks that did not elicit a reaction by the prey), but these errors did not occur after herring metamorphosis. The loss of the advantage of lower conspicuousness following metamorphosis was counterbalanced by increased reactivity. Antipredator benefits should increase with schooling behaviour (first evident in experiments at c . 50 mm), so predation mortality should decrease in postmetamorphic herring in the sea. Metamorphosis itself would be associated with high predation risk since conspicuousness is enhanced but reactivity and schooling behaviour are still not fully developed.     

Defending the Weak: Parental Defense Peaks When Chick Vulnerability is Greatest in the Herring Gull (Larus argentatus)

Successful reproduction often depends upon parents providing offspring with resources and protection. In birds, reproductive success can often be enhanced by parents engaging in antipredator behaviors, but these behaviors can be costly. Theoretically, individuals should temporally modify the intensity of nest defense behavior to balance the costs and benefits of current and future reproductive success. More specifically, nest defense should vary throughout a nesting attempt to maximize fitness of the adults. Here, we consider the relationship between nest defense behavior and chick vulnerability in the herring gull (Larus argentatus), where chicks are under high predation risk. We estimated chick vulnerability by quantifying survival probabilities at different periods of the nestling stage. Simultaneously, we quantified changes in parental aggression throughout the nesting cycle by simulating predation attempts using a human predator model. We found that chick survival probabilities were lowest (i.e., vulnerability was highest) and parental aggression in nest defense was greatest during the first 10 days after hatching. Thus, we show that parents are most defensive when chicks are most vulnerable and that adults optimize nest defense behaviors in a way that maximizes their fitness.     

Determination of clutch-size in the Kentish Plover Charadrius alexandrinus

We studied the clutch-size distribution of Kentish Plover Charadrius alexandrinus in 1991 and 1992 and tested three hypotheses for determination of clutch-size: egg-formation ability of females, incubation ability of parents and nest predation. Variation in clutch-size was small: 71 out of 74 clutches had three eggs (coeff. var. = 7%). Females spent more time foraging (51 ± 6%) pre-laying and during egg-laying than their mates (39 ± 5%). However, we concluded that egg-formation was not constrained by food availability because 77% and 100% of clutches were initiated before the peaks of prey density and of prey mass, respectively. Furthermore, the number of clutches initiated and completed over time was unrelated to prey density. By experimentally reducing and enlarging clutches, we found that enlarged clutches of four eggs took longer to hatch (24.8 ± 0.9 days) than control clutches of three (21.6 ± 0.7 days). Eggs of enlarged clutches also lost weight more slowly during incubation in both years compared with control clutches. No difference was found in the incubation behaviour or weight loss of parents between reduced, control and enlarged clutches. We found no evidence to support the nest predation hypothesis, since neither the proportion of nests predated nor the number of chicks hatched was different between reduced, control and enlarged clutches. The results of this study are most consistent with the incubation ability hypothesis, although the parental ability hypothesis has remained untested.     

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

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

Potential predation pressure of littoral mysids on herring (Clupea harengus membras L.) eggs and yolk-sac larvae

The predation potential of littoral mysid shrimps (Mysidacea) on Baltic herring (Clupea harengus membras L.) eggs and yolk-sac larvae was studied experimentally. The results showed that littoral mysids feed actively on both eggs and yolk-sac larvae. It was shown that Neomysis integer preys on eggs, which are not attached to the substrate. Alternative food (yolk-sac larvae or zooplankton) did not decrease feeding rate on eggs. Only gravel as a bottom material lowered the ingestion rate to nearly zero. The largest of the mysid species Praunus flexuosus ate yolk-sac larvae more than other mysids and most efficiently. Mysids switched to feed on eggs when larvae and eggs were offered simultaneously, thus predation focused on eggs. It is possible that hydrodynamic signals of moving larvae induced mysids to prey and eggs were easier prey to catch as well as more numerous. In addition egg size is optimal and the nutritive value (measured as C:N ratio) is better compared with larvae. The results indicate that mysids may have local effects on populations of Baltic herring by eating the early life stages, mostly eggs. Especially when large swarms of N. integer shoal in the spawning areas. However, the effect on recruitment of herring is still hard to evaluate.     

Costs of parental care on hunting behaviour of Helobdella papillornata (Euhirudinea: Glossiphoniidae)

Helobdella papillornata, an Australian freshwater leech, feeds primarily on snails and has a high level of parental care involving brooding eggs and young, with direct feeding of young. Parental costs and offspring benefits from these behaviours are poorly understood. A potential cost of parental care may be a change in the time taken to hunt prey. To test this hypothesis, the hunting behaviour of adults without progeny, parents with eggs, and parents with young were compared. We found that parents brooding eggs had a significantly (P = 0.029) longer lag time to begin hunting than parents brooding young, and spent significantly (P = 0.018) less time actively hunting than non-brooding adults. These costs, which may represent lost potential for the parent’s future reproductive success, should be outweighed by the fitness benefits of improved growth and survival of offspring, if parental care is favoured by selection. The hunting costs of care in Helobdella and other benthic, dorsoventrally flattened leeches in the family Glossiphoniidae may be smaller than the costs of brood tending that would be imposed on other freshwater leeches, and this difference may help explain the restriction of care to a single clade of the Euhirudinea.     

Facultative nest-parasitism among Iberian shrubsteppe passerines

This paper analyses intra- and interspecific facultative nest parasitism of shrubsteppe passerines on the Iberian peninsula through 4 seasons, from 1991–94. Daily frequency of nest parasitism was very low (f = 3.9 × 10^-3 during laying and f = 2.8 × 10^-4 during incubation). Observation of the 3 examples found in the field of 2 intraspecific and one interspecific instances revealed that the eggs were accepted and, in the latter case, that the parasitic chick was viable. The experimental tests in the field of placing 1 additional egg that was similar to (n = 57) or clearly different from the host (n = 50), in order to simulate both intra and interspecific parasitism revealed: (i) higher rejection rates in areas with higher risk of predation and (ii) similar rejection rates for both types of eggs. The experimental addition of nestlings of other species (n = 6) revealed that in 67% of the cases studied, the parasitic chick had a normal growth rate. It is suggested that nest parasitism among these species is an occasional behaviour pattern, caused by high nest predation rates in Iberian shrubsteppe habitats.     

Predators and the breeding bird: behavioral and reproductive flexibility under the risk of predation

A growing body of work suggests that breeding birds have a significant capacity to assess and respond, over ecological time, to changes in the risk of predation to both themselves and their eggs or nestlings. This review investigates the nature of this flexibility in the face of predation from both behavioural and reproductive perspectives, and also explores several directions for future research. Most available work addresses different aspects of nest predation. A substantial change in breeding location is perhaps the best documented response to nest predation, but such changes are not always observed and not necessarily the best strategy. Changes in nesting microhabitat (to more concealed locations) following predation are known to occur. Surprisingly little work addresses the proactive avoidance of areas with many nest predators, but such avoidance is probably widespread. Individual birds could conceivably adopt anti‐predator strategies based on the nest predators actually present in an area, but such effects have yet to be demonstrated. In fact, the ways in which birds assess the risk of nest predation is unclear. Nest defence in birds has historically received much attention, but little is known about how it interacts with other aspects of decision‐making by parents. Other studies concentrate on predation risk to adults. Some findings suggest that risk to adults themselves influences territory location, especially relative to raptor nests. An almost completely unexplored area concerns the sorts of social protection from predators that might exist during the breeding season. Flocking typical of the non‐breeding season appears unusual while breeding, but a mated pair may sometimes act as a “flock of two”. Opportunistic heterospecific sociality may exist, with heterospecific protector species associations more prevalent than currently appreciated. The dynamics of singing during the breeding season may also respond to variation in predation risk, but empirical research on this subject is limited. Furthermore, a few theoretical and empirical studies suggest that changes in predation risk also influence the behaviour of lekking males. The major influence of predators on avian life histories is undoubtedly expressed at a broad phylogenetic scale, but several studies hint at much flexibility on an ecological time scale. Some species may forgo breeding completely if the risk of nest predation is too high, and a few studies document smaller clutch sizes in response to an increase in nest predation. Recent evidence suggests that a female may produce smaller eggs rather than smaller clutches following an increase in nest predation risk. Such an increase may also influence decisions about intraspecific brood parasitism. There are no clear examples of changes in clutch/egg size with changes in risk experienced by adults, but parental responses to predators have clear consequences for offspring fitness. Changes in risk to adults may also influence body mass changes across the breeding season, although research here is sparse. The topics highlighted herein are all in need more empirical attention, and more experimental field work whenever feasible.     

Factors related to aggressive nest protection behaviour: a comparative study of Holarctic Waders

Data on 12 factors presumed to influence the distribution of aggressive nest defence in 111 species of waders (incubation-sharing by the parents, number of parents present near the nest, incubation time, nest habitat, breeding latitude, body mass, wing loading, wing structure, detectability on the nest, predator regime, coloniality and alternative prey) were collected from literature and field researchers. Body mass and number of parents present on the nest territory (within response range when avian predators appear) explain 50% of the variation in aggressive defence behaviour. The results support the notion that ecological conditions like predation pressure are important in shaping wader parental care systems, with implications for mating systems. Altogether, the investigated factors explain around 70% of the variation in the samples. Future research on the level of individuals is suggested in order to explain the remaining variation.