Predator population dynamics under a cyclic prey regime: numerical responses,demographic parameters and growth rates

The consequences of cyclic fluctuations in abundance of prey species on predator continue to improve our understanding of the mechanisms behind population regulation. Among predators, vole‐eating raptors usually respond to changes in prey abundance with no apparent time‐lag and therefore contradict predictions from the predator–prey theory. In such systems, the interplay between demographic traits and population growth rate in relation to prey abundance remains poorly studied, yet it is crucial to characterize the link between ecological processes and population changes. Using a mechanistic approach, we assessed the demographic rates associated to the direct and indirect numerical responses of a specialist raptor (Montagu's harrier) to its cyclic prey (common vole), using long term data from two adjacent study sites in France. First‐year survival rates were weakly affected by vole abundance, probably due to the fact that Montagu's harriers are trans‐Saharan migrants and thus escape the vole collapse occurring in autumn–winter. Recruitment of yearling as well as breeding propensity of experienced adult females were strongly affected by vole abundance and at least partially shaped the trajectory of the breeding population. We argued that the strong density dependent signal detected in predator time series was mostly the phenomenological consequence of the positive direct numerical response of harriers to vole abundance. Accounting for this, we proposed a method to assess density dependence in predator relying on a cyclic prey. Finally, the variation in Montagu's harrier population growth rates was best explained by overwinter growth rates of the prey population and to a lesser extent by previous residual predator density.     

Brood sex ratio varies with diet composition in a generalist raptor

While sex allocation has been investigated productively at both population and family levels, as yet no general theory has been developed that is capable of linking processes at these two ecological scales, and very few empirical studies have examined cross‐scale patterns. In Finnish northern goshawks (Accipiter gentilis), nestling sex ratio of local subpopulations is related to the spatial and temporal variation in the abundance of their principal avian prey, woodland grouse. Using data from an urban breeding population in Hamburg, Germany, I investigated: (1) whether brood sex ratio of goshawks varies with diet composition at the family level; (2) whether such variation could reflect adaptive adjustment; and (3) how family‐level allocation can drive population‐level patterns, such as those observed in Finland. Feral pigeons (Columba livia) were the most important prey species, with a pooled contribution to total diet of 36%. Brood sex ratio varied significantly with the proportion of pigeons in the breeding‐season diet of pairs (increasing male bias). However, there was no evidence for sex‐differential effects of diet composition, so it remains unclear whether the observed sex‐ratio variation was an adaptive response. As all study pairs inhabited an (urban) environment where pigeons were unusually abundant, family‐level sex‐ratio adjustment caused a marked male bias in offspring sex ratio at the population level (male‐biased nestling sex ratio in four of five years; pooled data: 60% males). This suggests that the large‐scale variation observed in Finnish goshawk populations mirrors sex‐ratio adjustment shown by individual families in response to small‐scale environmental conditions. Apart from linking patterns empirically across ecological scales, this study is, to my knowledge, the first to demonstrate that family‐level brood sex ratio varies with realized resource use (diet composition) in a raptor species. Previous studies either failed to find significant associations or, more commonly, violated theoretical assumptions by measuring environmental prey abundance (often integrated over large areas) rather than realized prey use of individual breeding pairs. I conducted a meta‐analysis of offspring sex‐ratio data from 17 goshawk populations across Europe to put my results into perspective. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 937–951.     

Maternal correlates of brood sex ratio variation in the lekking lance-tailed manakin Chiroxiphia lanceolata

Theory predicts that overall population sex ratios should be around parity. But when individual females can receive higher fitness from offspring of one sex, they may benefit by biasing their brood sex ratios accordingly. In lekking species, higher variance in male reproductive success relative to that of females predicts that male offspring gain disproportionately from favorable rearing conditions. Females should therefore produce male-biased broods when they are in a position to raise higher quality offspring: i.e., in better body condition or when they reproduce earlier in the breeding season. To investigate these hypotheses, we studied brood sex ratios of lance-tailed manakins Chiroxiphia lanceolata . We found that overall sex ratios and mean brood sex ratios were not different from random expectation. Brood sex ratios were not related to laying date or female body condition. However, we detected a quadratic relationship between brood sex ratios and maternal age: both young (1–2 years) and old (8+ years) females produced female-biased brood sex ratios. This relationship was most clear in a year also distinguished by early rainy and breeding seasons. We suggest that breeding inexperience in young females and senescence in older females is the most plausible explanation for these results, and that the relationship between female age and brood sex ratio is mediated by environmental conditions.     

Tawny Owls Strix aluco with reliable food supply produce male-biased broods

Tawny Owls Strix aluco have been reported to skew the sex ratio of their offspring towards males when facing food shortage during the nestling period (and vice versa), because female fitness is more compromised by food shortage during development than male fitness. To test the generality of these results we used a DNA marker technique to determine the sex ratio in broods of Tawny Owls in Danish deciduous woodland during two years of ample food supply (rodent population outbreak) and two years of poor food supply. Of 268 nestlings, 59% were males (95% CI: 53–65%). This proportion was higher than previously reported for the species (49% in Northumberland, UK, and 52% in Hungary), but consistent with Fisherian sex allocation, which predicts a male bias of c . 57% based on inferred differences in energy requirements of male and female chicks. Contrary to previous results, brood sex ratios were not correlated with the resource abundance during the breeding seasons, despite considerable variation in breeding frequency, brood size or hatching date across years. Brood sex ratios were unaffected by brood reduction prior to DNA sampling, and nestling mortality rates after DNA sampling were not related to gender. The inconsistency between the sex ratio allocation patterns in our study and previous investigations suggests that adaptive sex allocation strategies differ across populations. These differences may relate to reproductive constraints in our population, where reproductive decisions seem primarily to concern whether to lay eggs at all, rather than adjust the sex ratio to differences in starvation risk of nestlings.     

Tawny owl reproduction and offspring sex ratios under variable food conditions

Tawny owl reproduction and offspring sex ratios have been considered to depend on the abundance of small voles. We studied reproductive performance (laying date, clutch and brood size) during 1995–2003 and offspring sex ratios from 1999 to 2003 in relation to the abundance of small voles and food delivered to the nest in a tawny owl population in southern Finland. Abundance of small voles (field and bank voles) was based on trappings in the field, and estimates of food delivery was based on diet analysis of food remains in the nest boxes. In this population, reproductive output was not related to the abundance of small voles. Analysis of food delivered to the nest showed that the prey weight per offspring varied more than twofold between years and revealed that this difference was mainly related to the proportion of water voles in the diet. Only the number of water voles correlated with laying dates. Offspring sex ratios were weakly male biased (55%) but did not differ from parity. Sex ratios were not related to the abundance of small voles, and we found no evidence that parents delivered more food to nests with proportionally more offspring of the larger (female) sex. Our results underline the notion that populations may differ in their sex allocation pattern, and suggest such differences may be due to diet.     

Do predators limit the abundance of alternative prey? Experiments with vole‐eating avian and mammalian predators

Predation has been invoked as a factor synchronizing the population oscillations of sympatric prey species, either because predators kill prey unselectively (the Shared Predation Hypothesis; hereafter SPH), or because predators switch to alternative prey after a density decline in their main prey (the Alternative Prey Hypothesis; APH). A basic assumption of the APH is that the impact of predators on alternative prey depends more on the density of main prey than on the predator/alternative prey ratio. Both SPH and APH assume that the impact of predators on alternative prey is at least periodically strong enough to depress prey populations. To examine these assumptions, we utilized data from replicated field experiments in large areas where we reduced the breeding densities of avian predators during three years and the numbers of least weasels (Mustela nivalis) in two years when vole populations declined. In addition, we reduced the breeding densities of avian predators in two years when vole populations were high. The reduction of least weasels increased the abundance of their alternative prey, small birds breeding on the ground, but did not affect the abundance of common shrews (Sorex araneus). In years when vole populations declined, the reduction of avian predators increased the abundance of their alternative prey, common shrews and small birds. Therefore, vole‐eating predators do at least periodically depress the abundance of their alternative prey. At high vole densities, the reduction of avian predators did not increase the abundance of common shrews, although the ratio of avian predators to alternative prey was similar to years when vole populations declined, which supported APH. In contrast, the abundance of small birds increased after the reduction of avian predators also at high vole densities, which supported SPH. The manipulations had no obvious effect on the number of game birds, which are only occasionally killed by these small‐sized predators. We conclude that in communities where most predators are small or specialize on a single prey type, the synchronizing impact of predation is restricted to a few similar‐sized species.     

Variation in offspring sex ratio of a long‐lived sexually dimorphic raptor,the Eastern Imperial Eagle Aquila heliaca

Sex ratio theory attempts to explain observed variation in offspring sex ratio at both the population and the brood levels. In the context of low‐fecundity organisms producing high‐investment offspring, the drivers of adaptive variation in sex ratio are incompletely understood. For raptors that display reverse sexual dimorphism (RSD), preferential allocation of resources to the putatively cheaper sex (male) may be a response to environmental, social or demographic stressors. To assess the extent of skew in offspring sex ratios and to evaluate possible dietary, environmental and demographic correlates of such skew to long‐lived RSD avian species, we evaluated the offspring sex ratio of 219 chicks from 119 broods in 30 territories of Eastern Imperial Eagles Aquila heliaca across 7 years and four regions at a nature reserve in Kazakhstan. Only in one region in 1 year of our study did the offspring sex ratio differ significantly from parity (10 males : 1 female in 11 territories). Whereas offspring sex ratios were independent of dietary diversity, precipitation, temperature and productivity, we found that year had a moderate effect on brood sex ratio within territories. Our results provide limited evidence of brood sex manipulation in these populations of Eastern Imperial Eagles, and no mechanistic insight into predictions associated with it. Stochastic variation is likely to explain much of the fluctuation we observed in sex ratios, but our observations are also consistent with the hypothesis that sex‐ratio manipulation may occur irregularly, in concurrence with atypical environmental or demographic conditions that fluctuate at a time scale longer than that of our 7‐year study.     

Population dynamics in a cyclic environment: consequences of cyclic food abundance on tawny owl reproduction and survival

1. Understanding which factors regulate population dynamics may help us to understand how a population would respond to environmental change, and why some populations are declining.
2. In southern Finland, vole abundance shows a three-phased cycle of low, increase and decrease phases, but these have been fading out in recent years. During five such cycles (1981–1995), all tawny owls Strix aluco were censused in a 250-km² study area, and their reproduction and survival were monitored.
3. Males and females showed similar dynamics, but experienced breeders recruited more offspring and had higher survival than first breeders. Offspring recruitment, but not survival of breeding individuals varied in accordance with vole abundance.
4. The population's numerical response to prey abundance was primarily due to first-breeding individuals entering the population in the increase phase when immigration was the highest. First-breeding birds were younger, but experienced breeders were older in more favourable vole years.
5. A stage-specific matrix population model integrating survival and fecundity showed that, despite obvious variation in fecundity between vole cycle phases, this variation had limited importance for overall tawny owl population dynamics, but that the survival of experienced breeders during the low phase is most important for population growth.
6. Model and data agreed that the vole cycle drives the dynamics of this avian predator by limiting the recruitment of new breeders during the low phase. Population dynamics hence differ not only from the classic example of the species in a more temperate region in the UK where the number of territories is stable across years, but also from the dynamics of other avian vole predators in Fennoscandia where the recurring crash in vole abundance drastically lowers adult survival thereby creating vacancies.     

Numerical response of predators to large variations of grassland vole abundance and long‐term community changes

Voles can reach high densities with multiannual population fluctuations of large amplitude, and they are at the base of predator communities in Northern Eurasia and Northern America. This status places them at the heart of management conflicts wherein crop protection and health concerns are often raised against conservation issues. Here, a 20‐year survey describes the effects of large variations in grassland vole populations on the densities and the daily theoretical food intakes (TFI) of vole predators based on roadside counts. Our results show how the predator community responded to prey variations of large amplitude and how it reorganized with the increase in a dominant predator, here the red fox, which likely negatively impacted hare, European wildcat, and domestic cat populations. This population increase did not lead to an increase in the average number of predators present in the study area, suggesting compensations among resident species due to intraguild predation or competition. Large variations in vole predator number could be clearly attributed to the temporary increase in the populations of mobile birds of prey in response to grassland vole outbreaks. Our study provides empirical support for more timely and better focused actions in wildlife management and vole population control, and it supports an evidence‐based and constructive dialogue about management targets and options between all stakeholders of such socio‐ecosystems.

A 20‐year survey describes the effects of large variations in grassland vole populations on the densities and the daily theoretical food intakes of a vole predator community based on roadside counts. Our results show how the predator community responds to prey variations of large amplitude and how it reorganized with the increase of the red fox, which likely negatively impacted hare, European wildcat, and domestic cat populations.     

Carotenoids in nestling Montagu’s harriers: variations according to age, sex, body condition and evidence for diet-related limitations

Carotenoids are colored pigments forming the basis of many avian social traits. Before their utilization carotenoids must be acquired through diet and mobilized for specific uses. The relationships between carotenoid-based coloration, circulating carotenoids and body condition have been well studied in adult birds, but little is known in nestlings. Here, we investigated variations in carotenoid-based coloration in a raptor nestling, the Montagu’s harrier (Circus pygargus), both in captivity and in natural conditions, and within a vole (poor-carotenoid source and cyclic prey) specialist population. We studied these variations according to nestling age and sex, and possible limitations in carotenoid availability by comparing years of contrasted prey abundance and using carotenoid supplementation experiments. Captive nestlings, fed only with mice, were strongly carotenoid limited. Wild nestlings were also carotenoid limited, especially in a year of high vole abundance. Nestlings were in better condition but less colored during a peak vole abundance year than during a low vole abundance year, when harriers targeted more alternative preys (birds, insects). Thus, variation in vole abundance resulted in a de-coupling of body condition and carotenoid-based coloration in this population. This suggested that the positive relation between the body condition and carotenoid-based traits, typically found in adult birds, could be restricted to adults or nestlings of species that feed on carotenoid-rich food. Our results should stimulate more work on the functions and mechanisms of carotenoid-based traits in nestlings, which deserve more attention and most likely differ from those of adult birds.     

Spatial dynamics in breeding performance of a predator: the connection to prey availability

Using nationwide long-term data on goshawk and grouse populations in Finland we study the spatial dynamics of the numbers of breeding northern goshawk ( Accipiter gentilis ) pairs, goshawk brood size and offspring sex ratio and their connection to the abundance of grouse. Our first large-scale data comprise of observations on goshawk nests during 1986–2001 pooled to 21 different regions. The second set are annual (1989–1998) observations of brood size and offspring sex ratio (females over the sum of females and males) in goshawk nests all over the country, aggregated to 50 km grid level (n=28 grid units). The third set comprises counts (1989–2001) of four species of woodland grouse, split to adults and juveniles, also given in the same 50 km grid units. Using these data, we show that the annual numbers of northern goshawk nests in the different regions fluctuate in synchrony. Synchrony is also found in long-term fluctuations of northern goshawk brood size and offspring sex ratio. Moreover, synchrony is found in annual numbers of grouse juveniles and adults, the main prey for the northern goshawk. In the brood size and offspring sex ratio of the goshawk, as well as in the annual numbers of grouse juveniles and adults the degree of synchrony falls off with increasing distance. However, only in sex ratios and in grouse dynamics are the slopes of synchrony vs distance roughly matching. We also found that sex ratio either vs grouse juveniles or grouse adults has a more matching spatial dimension (50 km radius) that sex ratio vs brood size. These observation lend support to the hypothesis that goshawk offspring sex ratio and grouse abundance are interconnected. Despite the reason, consequences of spatial coupling in sex ratio could have repercussions on other life history events.     

Increased autumn rainfall disrupts predator–prey interactions in fragmented boreal forests

There is a pressing need to understand how changing climate interacts with land‐use change to affect predator–prey interactions in fragmented landscapes. This is particularly true in boreal ecosystems facing fast climate change and intensification in forestry practices. Here, we investigated the relative influence of autumn climate and habitat quality on the food‐storing behaviour of a generalist predator, the pygmy owl, using a unique data set of 15 850 prey items recorded in western Finland over 12 years. Our results highlighted strong effects of autumn climate (number of days with rainfall and with temperature <0 °C) on food‐store composition. Increasing frequency of days with precipitation in autumn triggered a decrease in (i) total prey biomass stored, (ii) the number of bank voles (main prey) stored, and (iii) the scaled mass index of pygmy owls. Increasing proportions of old spruce forests strengthened the functional response of owls to variations in vole abundance and were more prone to switch from main prey to alternative prey (passerine birds) depending on local climate conditions. High‐quality habitat may allow pygmy owls to buffer negative effects of inclement weather and cyclic variation in vole abundance. Additionally, our results evidenced sex‐specific trends in body condition, as the scaled mass index of smaller males increased while the scaled mass index of larger females decreased over the study period, probably due to sex‐specific foraging strategies and energy requirements. Long‐term temporal stability in local vole abundance refutes the hypothesis of climate‐driven change in vole abundance and suggests that rainier autumns could reduce the vulnerability of small mammals to predation by pygmy owls. As small rodents are key prey species for many predators in northern ecosystems, our findings raise concern about the impact of global change on boreal food webs through changes in main prey vulnerability.     

Effects of vole fluctuations on the population dynamics of the barn owl <Emphasis Type="Italic">Tyto alba</Emphasis>

Many predator species feed on prey that fluctuates in abundance from year to year. Birds of prey can face large fluctuations in food abundance i.e. small mammals, especially voles. These annual changes in prey abundance strongly affect the reproductive success and mortality of the individual predators and thus can be expected to influence their population dynamics and persistence. The barn owl, for example, shows large fluctuations in breeding success that correlate with the dynamics in voles, their main prey species. Analysis of the impact of fluctuations in vole abundance (their amplitude, peaks and lows, cycle length and regularity) with a simple predator prey model parameterized with literature data indicates population persistence is especially affected by years with low vole abundance. In these years the population can decline to low owl numbers such that the ensuing peak vole years cannot be exploited. This result is independent of the length and regularity of vole fluctuations. The relevance of this result for conservation of the barn owl and other birds of prey that show a numerical response to fluctuating prey species is discussed.     

Sex allocation in the sexually monomorphic fairy martin

Offspring sex ratios were examined at the population and family level in the sexually monomorphic, socially monogamous fairy martin Petrochelidon ariel at five colony sites over a 4-year period (1993–1996). The sex of 465 nestlings from 169 broods was determined using sex-specific PCR at the CHD locus. In accordance with predicted sex allocation patterns, population sex ratios at hatching and fledging did not differ from parity in any year and the variance in brood sex ratios did not deviate from the binomial distribution. Further, brood sex ratio did not vary with hatching date during the season, brood number, brood size or colony size. The sex ratio of broods with extra-pair young did not differ from those without, while the sex ratio of broods fathered by males that gained extra-pair fertilizations did not differ from broods fathered by other males. Extra-pair chicks were as likely to be male as female. Neither the total number of feeding visits to the brood nor the relative feeding contribution by the sexes varied significantly with brood sex ratio. Brood sex ratios were also unrelated to paternal size, condition and breeding experience or maternal condition and breeding experience. However, contrary to our prediction, brood sex ratio was negatively correlated with maternal size. Generally, these results were consistent with our expectations that brood sex ratios would not vary with environmental factors or parental characteristics, and would not influence the level of parental provisioning. However, the finding that females with longer tarsi produced an excess of daughters is difficult to reconcile with our current understanding of fairy martin life history and breeding ecology.     

Spatial dynamics of Microtus vole populations in continuous and fragmented agricultural landscapes

Small mammal populations often exhibit large-scale spatial synchrony, which is purportedly caused by stochastic weather-related environmental perturbations, predation or dispersal. To elucidate the relative synchronizing effects of environmental perturbations from those of dispersal movements of small mammalian prey or their predators, we investigated the spatial dynamics of Microtus vole populations in two differently structured landscapes which experience similar patterns of weather and climatic conditions. Vole and predator abundances were monitored for three years on 28 agricultural field sites arranged into two 120-km-long transect lines in western Finland. Sites on one transect were interconnected by continuous agricultural farmland (continuous landscape), while sites on the other were isolated from one another to a varying degree by mainly forests (fragmented landscape). Vole populations exhibited large-scale (>120 km) spatial synchrony in fluctuations, which did not differ in degree between the landscapes or decline with increasing distance between trapping sites. However, spatial variation in vole population growth rates was higher in the fragmented than in the continuous landscape. Although vole-eating predators were more numerous in the continuous agricultural landscape than in the fragmented, our results suggest that predators do not exert a great influence on the degree of spatial synchrony of vole population fluctuations, but they may contribute to bringing out-of-phase prey patches towards a regional density level. The spatial dynamics of vole populations were similar in both fragmented and continuous landscapes despite inter-landscape differences in both predator abundance and possibilities of vole dispersal. This implies that the primary source of synchronization lies in a common weather-related environment.     

The impact of raptors on the abundance of upland passerines and waders

The issue of predator limitation of vertebrate prey populations is contentious, particularly when it involves species of economic or conservation value. In this paper, we examine the case of raptor predation on upland passerines and waders in Scotland. We analysed the abundance of five wader and passerine species on an upland sporting estate in southern Scotland during an eight-year period when hen harrier, peregrine and merlin numbers increased due to strict law enforcement. The abundance of meadow pipit and skylark declined significantly during this time. Golden plover also showed a declining trend, whereas curlew increased significantly and there was a near significant increase in lapwings. Contrasting the local population trends of these species with trends on nearby areas revealed higher rates of decline for meadow pipit and skylark at the site where raptors increased, but no differences in trends for any of the three wader species. There was a negative relationship between the number of breeding harriers and meadow pipit abundance the same year and between total annual raptor numbers and meadow pipit abundance. Predation rates of meadow pipit and skylark determined from observations at harrier nests suggested that predation in June was sufficient to remove up to 40% of the June meadow pipit population and up to 34% of the June skylark population. This 'quasi-natural' experiment suggests that harrier predation limited the abundance of their main prey, meadow pipit, and possibly the abundance of skylark. Thus, high densities of harriers may in theory reduce the abundance of the prey species which determine their breeding densities, potentially leading to lower harrier breeding densities in subsequent years. We found no evidence to suggest that raptor predation limited the populations of any of the three wader species. We infer that concerns over the impact of natural densities of hen harriers on vulnerable upland waders are unjustified.     

No evidence of skewed secondary sex ratios in nestlings of the Common Raven (Corvus corax)

Many bird species adjust their offspring sex ratio as a response to environmental conditions or sexual dimorphism in size and dispersal. Offspring sex ratios may therefore vary among populations depending on the different demographic and ecological trajectories. We sampled Common Raven Corvus corax nestlings close to the fledging stage from three Central European regions to test for skewed secondary sex ratios and to investigate differences in sex ratios between populations that differ in recent recolonization history and breeding densities. Between 2005 and 2007, a total of 108 broods with 335 nestlings were sampled and their sex determined using molecular methods. We observed a mean of 3.1 (±1.2) nestlings per brood with no differences among nesting sites, years or regions. Nestling sex ratios were independent of the number of siblings. The overall secondary sex ratio was close to parity and did not differ between the variably structured populations.     

Factors Affecting Growth of Tengmalm’s Owl (Aegolius funereus) Nestlings: Prey Abundance,Sex and Hatching Order

In altricial birds, energy supply during growth is a major predictor of the physical condition and survival prospects of fledglings. A number of experimental studies have shown that nestling body mass and wing length can vary with particular extrinsic factors, but between-year observational data on this topic are scarce. Based on a seven-year observational study in a central European Tengmalm’s owl population we examine the effect of year, brood size, hatching order, and sex on nestling body mass and wing length, as well as the effect of prey abundance on parameters of growth curve. We found that nestling body mass varied among years, and parameters of growth curve, i.e. growth rate and inflection point in particular, increased with increasing abundance of the owl’s main prey (Apodemus mice, Microtus voles), and pooled prey abundance (Apodemus mice, Microtus voles, and Sorex shrews). Furthermore, nestling body mass varied with hatching order and between sexes being larger for females and for the first-hatched brood mates. Brood size had no effect on nestling body mass. Simultaneously, we found no effect of year, brood size, hatching order, or sex on the wing length of nestlings. Our findings suggest that in this temperate owl population, nestling body mass is more sensitive to prey abundance than is wing length. The latter is probably more limited by the physiology of the species.     

Family planning in a stemborer parasitoid: sex ratio, brood size and size-fitness relationships in Parallorhogas pyralophagus (Hymenoptera: Braconidae), and implications for biological control

Various aspects were studied of the brood size and sex allocation strategies, and of size-fitness relationships in Parallorhogas pyralophagus (Marsh), a gregarious ectoparasitoid of Eoreuma loftini Dyar. Brood size was significantly correlated with host size; larger hosts were allocated larger broods. Brood sex ratios were fixed precisely at 1 male per 4 females, and eggs were likely to be deposited in that order; differential mortality did not contribute to this precise sex ratio. The sex allocation strategy of P. pyralophagus is likely to conform to strict, i.e. single foundress, local mate competition. Adoption of this strategy is probably influenced by a limited insemination capacity of males; a smaller proportion of females (0.09 vs. 0.21) remained virgin in broods with precise or higher sex ratios (> or = 0.20 males) relative to broods with lower than precise sex ratios (< 0.20 males). Moreover, all females were inseminated in most broods (60%) with precise or higher sex ratios, whereas this did not occur in broods with lower than precise sex ratios. The hypothesized occurrence of strict local mate competition in P. pyralophagus was supported also by observations that: (i) offspring brood sex ratios were independent of maternal brood sex ratios and number of parental females concurrently allocating offspring to a group of hosts, and; (ii) the rate of superparasitism under no-choice conditions was low (approximately 20%), suggesting that rates of outbreeding in the field are low. Other results suggested that fitness in P. pyralophagus was correlated with adult size; longevity and reproductive capacity both increased with adult size in males and females. However, adult size may be more important for females than for males because the differences in reproductive capacity between the largest and smallest individuals was up to 7.3 times greater in females versus < 2 times in males.     

Of mice and mallards: positive indirect effects of coexisting prey on waterfowl nest success

Coexisting prey species interact indirectly via their shared predators when one prey type influences predation rates of the second prey type. In a temperate system where the predominant shared predator is a generalist, I studied the indirect effects of rodent populations on waterfowl nest success, both within the nesting season among sites and among years. Among six to ten upland fields (14 to 27 ha), mallard ( Anas platyrhynchos ) nest success was positively correlated with rodent abundance in all three years of the study. After removing year effects, mallard nest success remained positively correlated with the relative abundance of rodents. Of the rodent species present, California voles ( Microtus californicus ) were the most important coexisting prey type influencing nest success. Among years, mallard nest success was positively correlated with vole abundance; the asymptotic relationship suggests a threshold response to vole abundance, beyond which predators become satiated and additional voles do little to affect nest success. I tested and rejected three alternative explanations for the observed positive correlation between mallard nest success and rodent abundance that do not involve an indirect effect of coexisting prey populations. The influences of dense nesting cover, nesting density, and predator activity did not explain the observed patterns of nest success. These results suggest that rodent populations buffer predation on waterfowl nests, both within and among years, via the behavioral responses of shared predators to coexisting prey.