Studies on forest health and vegetation changes in Greece under the effects of climate changes

Greece, as part of the Mediterranean Basin, is projected to be among the most vulnerable countries to climate change. It is therefore quite urgent to adapt forest management to the changing climate in order to enhance biodiversity and to enable the conservation of healthy and productive forests. In the framework of the project LIFE+ AdaptFor (www.life-adaptfor.gr), an effort was made to understand the ecological responses and the vulnerability of forest ecosystems in the face of climate change; the overall aim of the project is the development and implementation of appropriate adaptation strategies. Four study areas were selected where changes in vegetation, quite likely attributed to climate change, have already been observed (dieback of Scots pine and Greek fir, intrusion of conifers in broadleaved forests). To investigate the synergism of climatic parameters’ alterations in the development of the occurring phenomena, time series of temperature and precipitation for the period 1950–2009 were produced and parameters of forest status were investigated, including mapping of vegetation changes through remote sensing. The findings support the hypothesis that climate change has an impact on forest health; the dieback of tree species can be attributed to outbreaks of pathogens (fungi and insects) which are associated with climatic parameters. However, the intrusion of conifer species into broadleaved forests showed no direct connection to climatic parameters, something that needs to be further investigated. In all cases, insufficient or inappropriate management practices applied in the areas exacerbated the occurring phenomena.     

Current nest box designs may not be optimal for the larger forest dormice: Pre‐hibernation increase in body mass might lead to sampling bias in ecological data

1. Biologists commonly use nest boxes to study small arboreal mammals, including the forest dormouse (Dryomys nitedula). Hibernating dormouse species often experience pronounced seasonal variations in body mass, which might lead to sampling biases if it is not taken into account when designing nest boxes. In my study of the forest dormouse, I noticed that the entrance hole of nest boxes had been gnawed. I hypothesized that this behavior was exhibited by the individual dormice of higher body mass, who were unable to pass through the entrance holes.
2. To test my hypothesis, I categorized the individual dormice present inside nest boxes based on their body mass and then compared the seasonal body mass dynamics with the timing of the gnawing behavior. I also compared nest box occupancy by the forest dormouse before and after the gnawing behavior.
3. Interestingly, I found that the gnawing behavior was displayed exclusively when part of the dormouse population increased considerably in body mass, which supports my hypothesis. Additionally, nest box occupancy decreased significantly from 20% before to 4.6% after the gnawing behavior.
4. I suggest that researchers include nest boxes with entrance holes larger than 4 cm in future studies of the forest dormouse to prevent the possible exclusion of the conspecifics that have higher body mass before hibernation. This type of sampling bias might also concern studies of other species, such as the fat dormouse, that similarly show pronounced seasonal variations in body mass. I recommend that biologists consider the seasonal body mass dynamics of the target species when designing nest boxes to minimize bias in ecological data and improve management actions.

     

Nest losses of cavity nesting birds caused by dormice (Gliridae, Rodentia)

Mammals are common predators on bird nests. However, their species identity frequently remains unknown. Here we present long-term data (1975–2005) from a central European woodland on the predatory effect of three dormice species (Rodentia, Gliridae) on cavity-nesting birds. Dormice are mostly frugivorous during the active late-summer season, but shortly after they terminate hibernation, they frequently depredate cavity-nesting-bird nests. The seven bird species studied, lost on average between 2.9 to 18.4% of their broods. MigratoryFicedula flycatchers suffered the highest brood losses, while the residentParus titmice and the nuthatchSitta europaea had much lower brood losses. The three dormice species differed significantly in their predatory effect during different avian breeding stages. The edible dormouseGlis glis (Linnaeus, 1766) depredated both eggs and nestlings equally, while the common dormouseMuscardinus avellanarius (Linnaeus, 1758) and the forest dormouseDryomys nitedula (Pallas, 1778) destroyed more nests during egg laying and the incubation period. Among adult birds, females were taken more frequently by dormice than males. Among avian species, adultFicedula flycatchers were more often depredated than the titmice and nuthatch. Our study provided further evidence, that among the traditional studies on the costs of reproduction, parental mortality at the nests needs to be considered and that incubating or brooding females might be under higher predation risk than the males.     

Influences of the feeding ecology on body mass and possible implications for reproduction in the edible dormouse (<Emphasis Type="Italic">Glis glis</Emphasis>)

The edible dormouse (Glis glis) is a small rodent and an obligate hibernator. Dormice undergo strong fluctuations of reproductive output during years that seem to be timed to coincide with future food supply. This behaviour enables them to avoid producing young that will starve with a high probability due to food shortage, and to increase their lifetime reproductive success. Aims of this study were to elucidate the extent to which feeding ecology in the edible dormouse has an impact on body mass and the fatty acid (FA) pattern of the white adipose tissue (WAT) before and after hibernation, which in turn might influence reproductive status in spring. Dormice show strong seasonal fluctuations of the body mass, which is reduced by one third during hibernation. Body mass and its changes depend on autumnal food availability as well as on the dietary FA pattern. During the pre-hibernation fattening period, dormice eat lipid rich food with a high content of linoleic acid. During hibernation, linoleic acid content is slightly but significantly reduced and body mass loss during winter is negatively correlated with the pre-hibernation linoleic acid content in the WAT. No relation between reproductive status and body mass, body condition or the FAs pattern of the WAT could be detected. However, in a year of high reproduction, dormice commence the shift to seed eating earlier than in a year of low reproduction. These seeds could be either a predictor for future food supply in autumn, or represent a high-energy food compensating high energetic costs of sexual activity in male edible dormice.     

Quantitative ultrastructural changes of hepatocyte constituents in euthermic,hibernating and arousing dormice (Muscardinus avellanarius)

Hibernating animals represent a suitable model for investigating the structural effects of drastic changes in cell activity under physiological conditions. In this study we investigated by means of electron microscopy and morphometrical analysis the fine structural counterpart of functional rest in hepatocytes of the hibernating dormouse, Muscardinus avellanarius, in comparison with arousing and euthermic dormice. Our observations demonstrate that during hibernation several structural constituents of the hepatocyte undergo modifications. In particular, during deep hibernation, the total cell and cytoplasm area significantly reduced, as well as the total and percent glycogen and residual body area, and the Golgi apparatus almost disappeared. Upon arousal, the amount of glycogen was minimal, whereas total cell and cytoplasm area significantly increased towards the euthermic value as well as total and percent residual body area. In comparison with the euthermic condition, the total and percent cell lipid area significantly increased in early hibernation, reduced in deep hibernation and almost disappeared during arousal. Taken together, our findings give quantitative ultrastructural support to the marked reduction found in hepatocyte functional activities during hibernation. Such a reduced activity involves profound rearrangement of the euthermic cell structure, which is rapidly resumed upon arousal.     

Stranded on an island: consequences of forest fragmentation for body size variations in an arboreal mammal, the edible dormouse (Glis glis)

The island rule states that small mammals isolated on islands have the evolutionary tendency to become larger, while large mammals tend to become smaller. However, the underlying mechanisms and life history consequences of these insular shifts in body size still remain speculative. The aim of this study was to investigate whether an arboreal mammal, the edible dormouse (Glis glis), showed shifts in body size when inhabiting isolated forest fragments. We analysed a data set of 541 individuals captured between 2005 and 2010 in four different forest fragments and one continuous forest, which served as a reference area. Sex, age, body mass, and size of all individuals were known. We used linear mixed-effect models to investigate whether individuals differed in their body size and mass between forest fragments and continuous forest. Our study revealed that edible dormice inhabiting forest fragments were significantly larger and heavier than individuals in the continuous forest, in accordance with patterns described by the island rule for small mammals. Because edible dormice frequently use nest boxes to rest during the day and to rear offspring, the life history strategies of this rodent can be easily investigated under evolutionary relevant conditions in the field. Thus the edible dormouse represents an excellent model organism for studying the mechanisms underlying shifts in body size as a response to habitat fragmentation and to investigate the consequences of these shifts on their life history strategies.     

Summer mortality in the hazel dormouse (Muscardinus avellanarius) and its effect on population dynamics

In the period 2000–2012, 38.3 % of 950 marked overwintered hazel dormice (Muscardinus avellanarius) were not recaptured at a study site in Lithuania in autumn. As adult dormice are sedentary, it is presumed that those dormice not recaptured died between late April and August. The highest total number of dormice captured for the last time was recorded in May and the lowest in August. The total summer mortality was significantly higher in females (42.5 %) than in males (34.6 %), but it did not depend on dormouse age or body weight. Tawny owl (Strix aluco) is the main known dormouse predator in Lithuania, and likely, it has the highest impact on summer mortality of M. avellanarius. Over the years, the total summer mortality of adult dormice ranged from 27 % to 52 %. The increased summer mortality resulted in decreased total dormouse population density and particularly decreased density of adult females in summer. Decreased densities led to more intensive breeding in the dormouse population, specifically breeding by young-of-the-year females, a pattern that is not common for this species. The number of breeding cases by young-of-the-year females was inversely related to the density of adult overwintered females in summer and to the number of breeding cases of these females. Breeding by young-of-the-year females was the main factor in the restoration of decreased population density in summer. Lithuanian populations of M. avellanarius are unique in their high proportion of breeding cases by young-of-the-year females amongst all populations investigated in the entire species distributional range.     

The influence of high nestbox density on the common dormouseMuscardinus avellanarius population

The response of common dormouseMuscardinus avellanarius Linnaeus, 1758 population to availability of nest sites was studied by manipulating the nestbox grid and ring-marking dormice. Abundance of adult dormice more than doubled in the 25 × 25 m nestbox grid in comparison to the 50 × 50 m grid, as a result of increased nestbox density from four to 16 boxes/ha. This effect already became apparent in the first year after additional nestboxes were made available and resulted from dormouse immigration, mostly from adjacent areas without nestboxes. In the second and third years, the number of two-year-old and older resident dormice, which had their home ranges in this plot, increased considerably. The average size of dormouse home range decreased by approximately half both in males and females in the 25 × 25 m grid compared to the 50 × 50 m grid. The proportion of breeding adult females did not differ between the two grids in spite of different adult dormouse density. Shortage of secure nest sites was a limiting factor for the common dormouse population abundance in the forest where natural tree hollows were absent, and high nestbox density increased environmental carrying capacity.     

Characteristics of Garden Dormice That Contribute to Their Capacity as Reservoirs for Lyme Disease Spirochetes

To describe the contribution of garden dormice to the epizootiology of Lyme disease, we compared their reservoir capacity for these pathogens to that of other sympatric hosts. Garden dormice are trapped most abundantly during early spring and again during midsummer, when their offspring forage. They are closely associated with moist forests. Garden dormice serve as hosts to nymphal ticks far more frequently than do other small mammals. Spirochetal infection is most prevalent in dormice, and many more larval ticks acquire infection in the course of feeding on these than on other rodents in the study site. Mature dormice appear to contribute more infections to the vector population than juveniles do. Replete larval ticks generally detach while their dormouse hosts remain within their nests. The population of garden dormice contributes five- to sevenfold more infections to the vector population than the mouse population does. Their competence, nymphal feeding density, and preference for a tick-permissive habitat combine to favor garden dormice over other putative reservoir hosts of Lyme disease spirochetes.     

Flea burden and its influence on nest selection and use in Dryomys nitedula Pallas, 1778

Since dormice usually use natural tree hollows or artificial nest boxes as nest sites and these nests are reused from 1 year to another, the present study investigated how the parasite loads from old nests affected the behaviour of the forest dormouse Dryomys nitedula Pallas, 1778. Dormice were subjected to three different experiments: 1. They were given the choice of a nest box infected with ectoparasites and a new unused nest box; 2. Dormice were given the choice of a treated (uninfected) previously used nest boxes and previously used nest box that was infected; and 3. Using infrared motion sensor cameras, we tested weather they preferred to avoid nest boxes when only infected nests were available. The results revealed that the females with pups are pickier when choosing the nursery site. These preferred empty nest boxes or nest boxes with treated nests instead of the infested ones. However, non-breeding adults and sub-adults showed indifference to the flea infestation of the nest.     

Diet of the fat dormouse (<Emphasis Type="Italic">Glis glis</Emphasis>) on the northern periphery of its distributional range

We studied the seasonal and annual variation in diet composition of the fat dormouse (Glis glis) in Lithuania, a locality situated on the northern periphery of the dormouse range and outside of the range of the European beech (Fagus sylvatica). After emergence from hibernation, dormice fed on oak acorns (from the previous year), inflorescences of various trees, vegetative parts of plants and food of animal origin (birds, their eggs and insects). In June, soft mast and seeds of birches supplemented the dormouse diet, and diet composition was the most diverse during this period. In July, raspberries and fruits of glossy buckthorn constituted the bulk of dormouse diet, but seeds of birches dominated in a specific year. Hard mast (mainly acorns) dominated the diet of G. glis from August until the beginning of hibernation in October. A high prevalence of acorns, comparatively high proportion of birch seeds and low proportion of food of animal origin in the diet, as well as feeding on fruits of glossy buckthorn, are specific features of feeding by G. glis in Lithuania. The diet of G. glis on the northern periphery of its range resembles its diet on the eastern periphery of the range where beech trees are also absent. According to the composition of G. glis diet, feeding conditions in both of these peripheral regions are poorer in comparison to central or southern regions.     

Climate- and resource-driven long-term changes in dormice populations negatively affect hole-nesting songbirds

Global climate change has been shown to cause variable shifts in phenology in a variety of animals and unexpected outcomes across food chains are to be found. Here we examined how rising annual spring temperatures affected the interactions between seed masting, cavity nesting birds and dormice using long-term data from Eastern Czech Republic. We have shown that climate change was associated with unequal shifts in the phenology of two cavity-breeding groups: dormice and birds. Rising spring temperatures have progressively advanced the termination of hibernation for the edible dormouse Glis glis , a common bird predator, leading to an increasing overlap in the use of nesting boxes between dormice and birds. In contrast, only the collared flycatcher Ficedula albicollis , of the four cavity-nesting bird species, advanced its breeding dates in response to rising temperatures. At the same time, favourable weather conditions, coupled with good seed masting years, have been associated with a substantial rise in dormice numbers. Concurrent with the increasing dormice abundance, the number of bird nests destroyed significantly increased in three out of four bird species. We showed that while there was a significant change in the date that the dormice emerged from hibernation during the course of the study, it did not significantly contribute to predation levels when controlling for their abundance and timing of breeding in birds. We found that the increasing dormice abundance was the main factor causing high brood losses in birds, while the timing of breeding in birds had a variable effect between bird species. This study illustrates how changes in climate might affect organisms at various trophic levels with often unexpected outcomes. Limited evidence from other study organisms suggests that species most at risk are those at different trophic levels that do not shift at the same rate or in the same direction as their food resources, predators or competitors.     

人工林生态系统持续发展的调控对策研究

基于田间试验和室内分析数据,分析了不同管理方式对人工林生态系统地力和持续发展的影响．结果表明,幼林期间种豆科牧草等农作物、营造混交林、针叶纯林间伐后引进阔叶树等生物途径以及改变整地方式、合理施肥等工程措施都有利于地力的恢复和林分生产力的提高．     

Survival rates in a small hibernator,the edible dormouse: a comparison across Europe

Understanding how local environmental factors lead to temporal variability of vital rates and to plasticity of life history tactics is one of the central questions in population ecology. We used long‐term capture‐recapture data from five populations of a small hibernating rodent, the edible dormouse Glis glis, collected over a large geographical range across Europe, to determine and analyze both seasonal patterns of local survival and their relation to reproductive activity. In all populations studied, survival was lowest in early summer, higher in late summer and highest during hibernation in winter. In reproductive years survival was always lower than in non‐reproductive years, and females had higher survival rates than males. Very high survival rates during winter indicate that edible dormice rarely die from starvation due to insufficient energy reserves during the hibernation period. Increased mortality in early summer was most likely caused by high predation risk and unmet energy demands. Those effects have probably an even stronger impact in reproductive years, in which dormice were more active. Although these patterns could be found in all areas, there were also considerable differences in average survival rates, with resulting differences in mean lifetime reproductive success between populations. Our results suggest that edible dormice have adapted their life history strategies to maximize lifetime reproductive success depending on the area specific frequency of seeding events of trees producing energy‐rich seeds.     

Mammal Society dormouse survey, January 1975-April 1979

The dormouse survey was initiated by the Mammal Society and ran from January 1975 to April 1979. The dormouse is very much a southern mammal in Britain with a reduced range compared to that found 100 years ago. There seems to have been a decline in population and possible reasons for this are discussed. The habitat favoured by dormice is considered; a thick secondary layer is very important with bramble forming a major component. Dormouse nests are most often found in bramble at an average height of just over a metre from the ground. Dormouse requirements, particularly for scrub, should be considered when areas for wildlife conservation are being managed.     

Seasonal Prevalence of Lyme Disease Spirochetes in a Heterothermic Mammal,the Edible Dormouse (Glis glis)

In Europe, dormice serve as competent reservoir hosts for particular genospecies of the tick-borne agent of Lyme disease (LD) and seem to support them more efficiently than do mice or voles. The longevity of edible dormice (Glis glis) and their attractiveness for ticks may result in a predominance of LD spirochetes in ticks questing in dormouse habitats. To investigate the role of edible dormice in the transmission cycle of LD spirochetes, we sampled skin tissue from the ear pinnae of dormice inhabiting five different study sites in south western Germany. Of 501 edible dormice, 12.6% harbored DNA of LD spirochetes. Edible dormice were infected most frequently with the pathogenic LD spirochete Borrelia afzelii. The DNA of B. garinii and B. bavariensis was detected in ca. 0.5% of the examined individuals. No spirochetal DNA was detectable in the skin of edible dormice until July, 6 weeks after they generally start to emerge from their obligate hibernation. Thereafter, the prevalence of spirochetal DNA in edible dormice increased during the remaining period of their 4 to 5 months of activity, reaching nearly 40% in September. Males were more than four times more likely to harbor LD spirochetes than females, and yearlings were almost twice more likely to be infected than adults. The seasonality of the prevalence of LD spirochetes in edible dormice was pronounced and may affect their role as a reservoir host in respect to other hosts.     

Fungal disease incidence along tree diversity gradients depends on latitude in European forests

European forests host a diversity of tree species that are increasingly threatened by fungal pathogens, which may have cascading consequences for forest ecosystems and their functioning. Previous experimental studies suggest that foliar and root pathogen abundance and disease severity decrease with increasing tree species diversity, but evidences from natural forests are rare. Here, we tested whether foliar fungal disease incidence was negatively affected by tree species diversity in different forest types across Europe. We measured the foliar fungal disease incidence on 16 different tree species in 209 plots in six European countries, representing a forest‐type gradient from the Mediterranean to boreal forests. Forest plots of single species (monoculture plots) and those with different combinations of two to five tree species (mixed species plots) were compared. Specifically, we analyzed the influence of tree species richness, functional type (conifer vs. broadleaved) and phylogenetic diversity on overall fungal disease incidence. The effect of tree species richness on disease incidence varied with latitude and functional type. Disease incidence tended to increase with tree diversity, in particular in northern latitudes. Disease incidence decreased with tree species richness in conifers, but not in broadleaved trees. However, for specific damage symptoms, no tree species richness effects were observed. Although the patterns were weak, susceptibility of forests to disease appears to depend on the forest site and tree type.     

Nest site attributes and temporal patterns of northern flicker nest loss: effects of predation and competition

To date, most studies of nest site selection have failed to take into account more than one source of nest loss (or have combined all sources in one analysis) when examining nest site characteristics, leaving us with an incomplete understanding of the potential trade-offs that individuals may face when selecting a nest site. Our objectives were to determine whether northern flickers (Colaptes auratus) may experience a trade-off in nest site selection in response to mammalian nest predation and nest loss to a cavity nest competitor (European starling, Sturnus vulgaris). We also document within-season temporal patterns of these two sources of nest loss with the hypothesis that flickers may also be constrained in the timing of reproduction under both predatory and competitive influence. Mammalian predators frequently depredated flicker nests that were: lower to the ground, less concealed by vegetation around the cavity entrance and at the base of the nest tree, closer to coniferous forest edges and in forest clumps with a high percentage of conifer content. Proximity to coniferous edges or coniferous trees increased the probability of nest predation, but nests near conifers were less likely to be lost to starlings. Flickers may thus face a trade-off in nest site selection with respect to safety from predators or competitors. Models suggested that peaks of nest predation and nest loss to eviction occurred at the same time, although a competing model suggested that the peak of nest loss to starlings occurred 5 days earlier than the peak of mammalian predation. Differences in peaks of mammalian predation and loss to starlings may constrain any adjustment in clutch initiation date by flickers to avoid one source of nest loss.     

Effects of food availability and ambient temperature on hibernation in the Japanese dormouse,Glirulus japonicus

The effects of food availability and ambient temperature (Ta) on hibernation in the Japanese dormouse,Glirulus japonicus, were examined. When the dormice were deprived of food under natural environmental conditions in midwinter, they showed typical hibernation patterns; in contrast, the animals given ad lib food did not show continuous long-term torpor. Furthermore, exposure to constant low temperature did not affect the torpor patterns in the animals receiving adequate food. These results suggest that food availability plays a decisive role in the onset and maintenance of hibernation in this species, and that low Ta does not have a great influence on the torpor patterns in animals with access to food. This differs from the previous study that emphasized the importance of Ta in the occurrence of torpor and discounted the effects of food shortage.     

Polyandry by wood mice in natural populations

The microhabitat used by the garden dormouse Eliomys quercinus during its nocturnal activity was examined. Twenty-three variables describing habitat structure were measured at 100 trapping points. Logistic regression models were utilized to select the variables that discriminate between used and avoided trapping points and, in a series of pairwise contrasts, between trap sites used by different sexes, age classes and animals in different reproductive periods. Used sites were characterized by a higher (>40%) rock cover, a thicker shrub layer and a younger understory with trees of smaller trunk diameter. These variables describe areas with rockfalls from the upper slope, where trees were younger and smaller, the canopy closure reduced and the shrub layer more developed than in other parts of the forest. Dense shrubby vegetation provided protection from aerial predators and, in conjunction with rocks and stones, might have made hunting by birds and mammals more difficult. Garden dormice may also have been attracted by rocky areas acting as a heat source during the night. The selection of rocky areas was more important in the first months of the active period, when animals emerged from hibernation and started the mating season. At the end of summer, the animals used areas richer in herbs, where insects and other food resources were probably more abundant.