Suggestions of collapse in small herbivore cycles since the 1980s have raised concerns about the loss of essential ecosystem functions. Whether such phenomena are general and result from extrinsic environmental changes or from intrinsic process stochasticity is currently unknown. Using a large compilation of time series of vole abundances, we demonstrate consistent cycle amplitude dampening associated with a reduction in winter population growth, although regulatory processes responsible for cyclicity have not been lost. The underlying syndrome of change throughout Europe and grass-eating vole species suggests a common climatic driver. Increasing intervals of low-amplitude small herbivore population fluctuations are expected in the future, and these may have cascading impacts on trophic webs across ecosystems.
Summary 1.A geographical gradient in the relative impact of generalist and specialist predators on small rodent populations has been hypothesized to be responsible for the gradient in cyclicity found in Fennoscandia. Population oscillations resulting from weasel±vole interactions are said to be dampened by the increasing stabilizing impact of generalist predators in southern Fennoscandia resulting from: (i) a greater abundance and diversity of predators sustained by alternative prey; (ii) the absence of signi®cant snow cover leading to constant exposure of voles to generalist predators; and (iii) a heterogeneous habitat that makes dispersing voles more vulnerable to predators. 2. Changes in the abundance of ®eld voles (Microtus agrestis L.) in a man-made spruce forest in northern England were recorded during 1984±98 using sign indices at 14±18 sites calibrated with capture±recapture estimates of vole density. 3. Field vole populations exhibited cyclic dynamics which were in many ways similar to those reported from Fennoscandia, including population declines taking place during the breeding season and long periods with no recovery in numbers following population crashes. 4. The density dependence structure of the time series was explored by means of partial autocorrelation functions, which suggested second-order density dependence. Analyses based on two density estimates per year (spring and autumn) reveal signi®cant negative values for lags of 1, 1´5 and 2 years, suggesting that the time-lag might be somewhat shorter than 2 years. 5. Estimates of predation on ®eld voles by red foxes and tawny owls at high vole density were above the value predicted for this site and for the whole generalist predator community by a published model assuming that predation by generalist predators stabilizes vole populations. However, empirical estimates of the parameter used both for designing and testing the model are inherently imprecise. 6. A qualitative evaluation of the three variables (see 1) correlated to the Fennoscandian gradient and assumed to contribute to variations in generalist predation pressure did not support the hypothesis that low predation rates by generalist predators are necessary for vole dynamics to be dominated by the destabilizing impact of weasel±vole interactions. The specialist/generalist predation hypothesis must therefore be modi®ed to account for the regular population cycles occurring in northern Britain.
We demonstrate evidence for the presence of travelling waves in a cyclic population of field voles in northern Britain by fitting simple, empirical models to spatially referenced time series data. Population cycles were broadly synchronous at all sites, but use of Mantel correlations suggested a strong spatial pattern along one axis at a projection line 72 degrees from North. We then fitted a generalized additive model to log population density assuming a fixed-form travelling wave in one spatial dimension for which the density at each site was offset in time by a constant amount from a standard density-time curve. We assumed that the magnitude of this offset would be proportional to the spatial separation between any given site and the centroid of the sampling sites, where separation is the distance between sites in a fixed direction. After fitting this model, we estimated that the wave moved at an average speed of 19 km yr-1, heading from West to East at an angle of 78 degrees from North. Nomadic avian predators which could synchronize populations over large areas are scarce and the travelling wave may be caused by density-dependent dispersal by field voles and/or predation by weasels, both of which act at a suitably small spatial scale.
Tawny owls, Strix aluco, laid female-biased clutches on territories with more abundant prey (field voles) in June, the month that chicks fledge. This appeared to enhance the subsequent reproductive success of fledglings, as in 1995 there was a significant correlation between the number of chicks fledged by adult females and the June vole abundance in the territory on which they were reared as chicks. This relationship did not hold for males. Since tawny owls lay eggs in March, these results indicate that owls are able to predict the June vole numbers on their territory, and respond by producing more of the sex most likely to gain a long-term benefit when resources are good.
We have previously documented the decline of the Common Kestrel Falco tinnunculus over a 23‐year period in a large coniferous forest in northern England. Kestrels fed predominantly on Field Voles Microtus agrestis, which were most abundant in young plantations (1–11 years old). Over the 23 years, voles remained abundant in the study area, but their numbers fluctuated cyclically. Here we consider whether the decline of Kestrels was linked to predation by Northern Goshawk Accipiter gentilis. Goshawks first bred in the study area in 1973 and increased until 1989, after which numbers stabilized. We use a number of approaches to explore the role of Goshawk predation, all of which are correlative, but independent. First, there was a significant negative relationship between Kestrel and Goshawk numbers after controlling for a decline in vole habitat. Short‐eared Owls Asio flammeus, which also hunt by day, declined over the same period as Kestrels. Second, numbers of Tawny Owl Strix aluco and Long‐eared Owl Asio otus did not decline as Goshawk numbers increased. These two species are also vole‐dependent, but active by night, and less vulnerable to Goshawk attack. Third, six species of raptor comprised 4.5% of 5445 Goshawk prey items during the breeding season, but more Kestrels were killed than the combined total of all other raptors. Goshawks not only killed many adult Kestrels in early spring, prior to breeding, when it would have most impact on population levels, but there was also a temporal trend for predation on Kestrels to be inversely density‐dependent. Finally, we estimated that Goshawks removed more Kestrels than were recorded each spring in the study area. We interpreted this as indicating that immigrant Kestrels were being removed continually, mostly before they could breed. We conclude that the decline of Kestrels (and possibly Short‐eared Owls) was mainly due to predation by Goshawks. This study provides some of the strongest evidence yet of the role of predation in the hierarchical structuring of raptor communities.
5. While we still do not know what causes the local oscillations, our models question the need to invoke additional factors to explain large-scale synchrony and travelling waves beyond small-scale dispersal and local density-dependent feedback. Our work also suggests that the higher degrees of synchrony observed in Fennoscandian habitats compared with Kielder may be due in part to the relative ease of movement of voles in these former habitats. As our work con®rms that the rates of exchange among local populations will have a strong in¯uence on synchrony, then we anticipate that the spatiotemporal distribution of clear-cuts will also have an important in¯uence on the dynamics of predators of voles.
Summary1. Natal conditions and senescence are two major factors shaping life-history traits of wild animals. However, such factors have rarely been investigated together, and it remains largely unknown whether they interact to affect age-specific performance. 2. We used 27 years of longitudinal data collected on tawny owls with estimates of prey density (field voles) from Kielder Forest (UK) to investigate how prey density at birth affects ageing patterns in reproduction and survival. 3. Natal conditions experienced by tawny owls, measured in terms of vole density, dramatically varied among cohorts and explained 87% of the deviance in first-year apparent survival (annual estimates ranging from 0AE07 to 0AE33). 4. We found evidence for senescence in survival for females as well as for males. Model-averaged estimates showed that adult survival probability declined linearly with age for females from age 1. In contrast, male survival probability, lower on average than for female, declined after a plateau at age 1-3. 5. We also found evidence for reproductive senescence (number of offspring). For females, reproductive performance increased until age 9 then declined. Males showed an earlier decline in reproductive performance with an onset of senescence at age 3. 6. Long-lasting effects of natal environmental conditions were sex specific. Female reproductive performance was substantially related to natal conditions (difference of 0AE24 fledgling per breeding event between females born in the first or third quartile of vole density), whereas male performance was not. We found no evidence for tawny owls born in years with low prey density having accelerated rates of senescence. 7. Our results, combined with previous findings, suggest the way natal environmental conditions affect senescence varies not only across species but also within species according to gender and the demographic trait considered.
The goshawk Accipiter gentilis has recently been reintroduced into parts of Great Britain. During the course of a study of one population, lesions of stomatitis were observed in 14 young from five broods and all the affected birds died. Postmortem examination of three birds revealed live Trichomonas gallinae in exudate from one, and histological findings consistent with a diagnosis of trichomoniasis were made in this and one other bird. It is suggested that trichomoniasis may be a significant mortality factor in goshawks from Britain.
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