Emil Tkadlec

Europe-Wide Dampening of Population Cycles in Keystone Herbivores

Cycling in Unison Many small mammals, especially voles, display semi-regular cycles of population boom and bust. Given the fundamental importance of small mammals as basal consumers and prey, such cycles can have cascading effects in trophic food webs. Cornulier et al. (p. 63) collated raw data from vole populations across Europe collected over the past 18 years. Reduction in winter growth rate was common across a wide variety of habitats with very different local climates, suggesting the presence of a continental-scale climatic driver of vole populations. Synchronicity in vole population fluctuation across Europe suggests a common climatic driver. 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.

A new geographical gradient in vole population dynamics.

A new geographical gradient in the dynamics of small rodents is demonstrated by analysing 29 time series of density indices of the common vole (Microtus arvalis) from Poland, the Czech Republic and the Slovak Republic. This gradient extends from more stable northerly populations in coastal Poland to more variable and cyclic populations in the southernmost parts of the Slovak Republic, and is hence a reversal of the Fennoscandian gradient. All studied variables (such as mean density, cycle amplitude, density variability and the coefficients in a second–order autoregressive model) exhibit consistent latitudinal variation. Possible underlying factors are discussed. In particular, we suggest that seasonality may be a key element in explaining the observed new gradient.

Small rodent population fluctuations: The effects of age structure and seasonality

Despite more than 50 years of effort, the causes and mechanisms of small rodent population fluctuations remain unknown. The two major questions are as follows: (1) what is the cause of population decline and (2) what is the cause of cyclicity and its geographical variation? At present, no hypothesis can provide answers to both these questions. Recently, progress has been made by Boonstra (1994), who proposed the senescence hypothesis to explain the cause of cyclic decline in population numbers. Here, we tested the main prediction that voles in decline are older than in other phases of the cycle, by analysing changes in age structure in a fluctuating population of the bank vole (Clethrionomys glareolus). The results generally support this prediction; however, the differences in absolute age seem to be too small to explain the occurrence of senescent animals exclusively in declines. We propose a new model to explain changes in age structure and the mechanisms behind the decline and geographic variation in cyclicity. It is based on the idea that voles are oldest in declines, developed independently of Boonstra. However, it differs in three respects: (1) it is more general and thereby applicable to the whole cycle; (2) density-dependent changes in age structure are based on the bimodality in a females age at first reproduction; and (3) it stresses developmental rather than physiological changes in the quality of decline of animals as being relevant to the rate of senescence. We propose that seasonality of the environment is a principal candidate to explain geographical variation in cyclicity. We present substantial theoretical and empirical evidence to indicate that in more seasonal environments with shortened vegetation periods, population dynamics is inevitably less stable due to increased variation in two critical parameters – age at first reproduction and the length of the breeding season – which determine population growth rates. Any external perturbation may then easily destabilize population numbers. The general applicability of the seasonality-senescence hypothesis to other mammalian species decreases with declining r and increasing life span. The hypothesis is falsifiable, and testable predictions are provided.

Increasing Incidence of Geomyces destructans Fungus in Bats from the Czech Republic and Slovakia

Background White-nose syndrome is a disease of hibernating insectivorous bats associated with the fungus Geomyces destructans. It first appeared in North America in 2006, where over a million bats died since then. In Europe, G. destructans was first identified in France in 2009. Its distribution, infection dynamics, and effects on hibernating bats in Europe are largely unknown. Methodology/Principal Findings We screened hibernacula in the Czech Republic and Slovakia for the presence of the fungus during the winter seasons of 2008/2009 and 2009/2010. In winter 2009/2010, we found infected bats in 76 out of 98 surveyed sites, in which the majority had been previously negative. A photographic record of over 6000 hibernating bats, taken since 1994, revealed bats with fungal growths since 1995; however, the incidence of such bats increased in Myotis myotis from 2% in 2007 to 14% by 2010. Microscopic, cultivation and molecular genetic evaluations confirmed the identity of the recently sampled fungus as G. destructans, and demonstrated its continuous distribution in the studied area. At the end of the hibernation season we recorded pathologic changes in the skin of the affected bats, from which the fungus was isolated. We registered no mass mortality caused by the fungus, and the recorded population decline in the last two years of the most affected species, M. myotis, is within the population trend prediction interval. Conclusions/Significance G. destructans was found to be widespread in the Czech Republic and Slovakia, with an epizootic incidence in bats during the most recent years. Further development of the situation urgently requires a detailed pan-European monitoring scheme.

Density‐dependent life histories in female bank voles from fluctuating populations

Using long-term data sets from two lowland floodplain forest bank vole populations in central Europe, we tested two predictions that with increasing densities: (i) proportions of nonparous females in winter populations would increase; and (ii) age of both nonparous and parous females would increase. These two predictions follow from the assumption that changes in age structure are driven by density-dependent shifts in age at first reproduction. Both populations were sampled by snap trapping between 1956 and 1976. For each year, we examined samples of animals collected between November and April for proportions of females with and without breeding experience and analysed their variation in age relative to the population density. The presence or absence of placental scars was used to discriminate between the parous and nonparous animals. Age was determined by measuring the length of the first mandibular molar (M1 ) roots. Both populations exhibited multi-annual fluctuations in numbers closely resembling those in northern Fennoscandia. The proportion of nonparous females in our total sample was 0.73, suggesting that it is uncommon for parous female bank voles to breed in two successive years and that their life histories are largely designed for breeding in one season only as a major reproductive strategy. Using a logistic-binomial regression model, we found that the probability of females being nonparous or parous at capture varied significantly with time, space and population density. The final model producing the best fit to data predicted that the proportion of nonparous females would be slightly larger over winter and substantially larger after high-density breeding seasons, which is consistent with the tested prediction. With increasing densities during the breeding seasons, both the nonparous and parous females became older at the onset of winter. Again, the field evidence was consistent with the predicted pattern. The age of parous females in the autumn at high densities was older, mainly because of an increased proportion of females entering their second winter. We propose a new intrinsic mechanism by which a delayed density dependence in age-specific fitness components can arise in microtines. This mechanism, based on an increased bimodality in age at first reproduction towards northern latitudes, has a potential to explain the south-north gradient in cyclicity observed in microtine populations in Fennoscandia.

The distribution and spreading pattern of Dermacentor reticulatus over its threshold area in the Czech Republic—How much is range of this vector expanding?

Host-seeking Dermacentor reticulatus ticks were detected by flagging method at 46 localities at south-east part of the Czech Republic, in the basins of rivers Morava and Dyje. Exact north-west distribution limits of D. reticulatus were defined in this area for the first time. Detailed prediction map of probabilities of D. reticulatus occurrence was obtained using GIS analysis. Spatial model delimited a south-north gradient in probability across the studied area, with highest probabilities above 0.8 in its southernmost part. Abundance of D. reticulatus varied markedly between localities in interval 0.33-222 of ticks per flag per hour. The highest abundances were in flooded areas at lower streams, towards upper streams abundance and density of these ticks decreased. Females prevailed in samples with population sex ratio of 0.413, significantly deviating from parity. Larvae and nymphs of this species were not detected by flagging. Although D. reticulatus range expansion probably did not reach such a degree as reported in other countries, these ticks became very abundant in some parts of studied area. Since spreading of vector-borne diseases became a problem in Europe, the knowledge of their exact recent geographic ranges is important for future modelling of their shift predictability.

West-to-east differences of Babesia canis canis prevalence in Dermacentor reticulatus ticks in Slovakia

Babesia canis canis is the most frequent causative agent of canine babesiosis in Central Europe, frequently causing severe disease. Recently, many new endemic foci of this disease have been reported from European countries. Growing incidence of canine babesiosis was recorded also in Slovakia during the last decade, from first cases in eastern Slovakia ten years ago to recent cases all over the south of the country. We have used nested PCR-RFLP method to study prevalence of B. c. canis in its natural tick vector Dermacentor reticulatus, collected at three geographically isolated lowland areas of southern Slovakia situated in the southeast, southwest, and west of Slovakia, respectively. The highest prevalence of B. c. canis was observed in D. reticulatus from eastern Slovakia (14.7%; n=327), whereas the prevalence in southwest was significantly lower (2.3%; n=1205). Notably, all 874 D. reticulatus ticks collected at Záhorská nížina lowland (W Slovakia) were B. c. canis-negative. Recorded differences in Babesia prevalence concurs well with the shift in incidence of clinical cases of canine babesiosis as observed by vet practitioners. Presented results revealed that eastern Slovakia represents an area of high risk of B. c. canis infection, whereas western areas of the country still remain Babesia canis-free.

Precocious breeding in female common voles and its relevance to rodent fluctuations

Age at first reproduction has been recognized as the most significant life-history parameter in determining population growth rates. Females in fluctuating rodent populations show great phenotypic variation in this parameter with precocious breeding, defined as pre-weaning fertile mating, being among the most extreme cases of early reproduction in mammals. Here we present the results of experiments in which 14-d-old female common voles (Microtus arvalis) were paired with adult males. Precocious breeding was common and coupled with unexpectedly large litters. We consider this reproductive feature to be adaptive in strictly seasonal environments with greater variation in length of the breeding season, with high resource availability, and a low level of intraspecific competition. Under these conditions, linkage between precocious breeding and high fertility has evolved as part of the most effective life-history strategy which produces the high intrinsic rates of population increase typical of small rodent populations. Seasonality of the environment was suggested to be the main factor behind the geographic pattern in small rodent cyclicity

Age variation in a fluctuating population of the common vole

We analysed variation in age in a fluctuating population of the common vole (Microtus arvalis) in southern Moravia, Czech Republic, to test the assumption of the senescence hypothesis that the age of voles increases with increasing population density. Between 1996 and 1998, we monitored the demographic changes by snap-trapping and live-trapping in a field population passing through the increase, peak and decline phase of the population cycle. We used the eye lens mass method to determine the age of snap-trapped animals and those that died in live-traps. The average age of winter males was clearly higher after the peak phase breeding season than before it. No such phase-dependent shift in age, however, was observed in the female component. Male age continued to increase from autumn to spring over the pre-peak winter, and the highest age was in spring of the peak phase year. However, after the peak phase breeding season the highest age was achieved in winter, with the decline phase males during the next spring tending to be younger. The average age of females in spring populations was always lower than in winter populations. The average age of voles from live-traps was always higher than voles from snap-traps, particularly in winter and spring populations, suggesting the presence of senescent animals. Although the density-dependent changes in age are consistent with those observed for other voles, they provide only weak evidence that population cycles in the common vole are accompanied by pronounced shifts in individual age, particularly in female voles.

Distribution of the common hamster in the Czech Republic after 2000: retreating to optimum lowland habitats

Abstract. Abundances of the common hamster in western Europe declined dramatically over the last 30 years. Recently, severe restrictions in distribution range have also been reported from central European countries. Here we update knowledge of the hamster distribution range in the Czech Republic based on information from six independent sources: (1) monitoring programme for the common vole carried out by State Phytosanitary Administration, (2) monitoring carried out directly by us, (3) questionnaire data from farming companies, (4) questionnaire data from district museums, (5) data from a public server BioLib for mapping species distribution, and (6) observations from nonrelated research activities and persons taking interest in hamsters. The comparison of locations detected after 2000 with those from the last survey > 30 years ago suggests that the range is severely reduced with hamsters retreating to the optimum lowland habitats along large rivers. These results suggest that the demographic mechanisms causing population decline in western populations operate in central Europe as well.