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.     

Frequency and demographic parameters of white-tipped common dormiceMuscardinus avellanarius

Out of 1154 marked common dormiceMuscardinus avellanarius (Linnaeus, 1758), 219 animals (19.0%) had white tail tips at Vilkaraistis forest (Moletai district, east Lithuania) in 1984–1993. The extent of white tail-tip varied from a slight tuft to a notable 22 mm length of white hair in the tail tip. In this population white-tipped individuals represented, on average, 25.7% among juvenile females, 24.9% among adult females, 13.4% among juvenile males, and 15.7% among adult males. White-tipped females were significantly more frequent than males, both in juveniles and in adults. The share of white-tipped dormice fluctuated in different years from 12.5% to 25.6% among adult dormice, and from 9.2% to 28.3% among juveniles. The percentage of white-tipped dormice among juveniles in different years was directly proportional to population density in autumn. There was no clear difference or trend in the mortality rate in white-tipped versus normal dormice. The contribution of white-tipped females to population reproduction was relatively lower than that of normal females: only 18.4% out of all registered breeding cases (n=212) were due to white-tipped females.     

Daily torpor in free-ranging common dormice (Muscardinus avellanarius) in Lithuania

Torpid common dormice (Muscardinus avellanarius) were found in nestboxes during all the activity period from early April until late October. Prevalence of torpor among dormice was highest in spring, decreased considerably in summer and increased again in October. The proportion of adult dormice that were torpid was inversely related to mean monthly air temperature, except in September, when dormice prepare for hibernation by accumulating fat reserves. In spring and summer, torpid dormice were found when ambient temperature was up to 14–15 °C and in the arousal phase of torpor – up to 19–20 °C. In autumn, dormice were active at much lower ambient temperatures compared to spring and summer, and only dormice that were fat enough and ready for hibernation were torpid. Torpor bouts usually lasted only until noon, but also in the afternoon during persistently low ambient temperatures in April and October. In April–July, torpor was more frequent among adult males than females (75% vs. 46%), and pregnant females were never found in deep torpor. Prevalence of torpor among young-of-the-year was lower compared to adult dormice at the same time. In October, the average weight of torpid young-of-the-year dormice was significantly higher compared to the weight of active dormice (22.3 vs. 17.5 g).

Zusammenfassung

Tagestorpor bei frei lebenden Haselmäusen (Muscardinus avellanarius) in LitauenHaselmäuse (Muscardinus avellanarius) im Torpor wurden während der gesamten aktiven Saison zwischen zeitigem April und Ende Oktober in Nistkästen gefunden. Im Frühjahr war die Torporfrequenz am höchsten, während des Sommers ging sie beträchtlich zurück, und im Oktober stieg sie wieder deutlich. Der Anteil an Haselmäusen im Torpor war umgekehrt abhängig von der mittleren Monatslufttemperatur, mit Ausnahme des Septembers, wo sich die Haselmäuse Fettreserven in Vorbereitung des Winterschlafes anlegen. Während des Frühlings und des Sommers konnten Haselmäuse im Torpor gefunden werden, wenn die Umgebungstemperatur bis 14–15 °C war, und in der Aufwachphase bei Temperaturen bis 19–20 °C. Im Herbst waren die Haselmäuse bei viel tieferen Temperaturen als im Sommer oder Frühjahr aktiv und nur die Haselmäuse die fett genug waren, um in den Winterschlaf gehen zu können, waren lethargisch. Die Torporphasen dauerten in der Regel nur bis zum Mittag, aber im April und Oktober während lang anhaltenden tiefen Umgebungstemperaturen auch bis in den Nachmittag hinein. Zwischen April und Juli konnten mehr adulte Männchen als Weibchen im Torpor nachgewiesen werden (75% vs 46%). Tragende Weibchen waren niemals im tiefen Torpor zu finden. Die Torporfrequenz war bei Haselmäusen in ihrem ersten Lebensjahr geringer als bei adulten im selben Zeitraum. Im Oktober war bei den lethargischen Jungtieren des Jahres das Durchschnittsgewicht höher als bei den aktiven Haselmäusen (22.3 vs. 17.5 g).     

Protecting Free-Living Dormice: Molecular Identification of Cestode Parasites in Captive Dormice (<Emphasis Type="Italic">Muscardinus avellanarius</Emphasis>) Destined for Reintroduction

The success of any population translocation programme relies heavily on the measures implemented to control and monitor the spread of disease. Without these measures, programmes run the risk of releasing immunologically naïve species or, more dangerously, introducing novel infectious agents to native populations. As a precaution, a reintroduction programme for the common or hazel dormouse, Muscardinus avellanarius, in England screens dormice before release following captive breeding. Using PCR sequencing of a range of genes, we tested whether the same species of tapeworm(s) were present in captive and free-living dormice. Whilst only Rodentolepis straminea were identified in free-living dormice, cestode ova found in a captive individual produced a molecular match closely related to Hymenolepis microstoma and a previously unrecorded Rodentolepis species. To prevent putting at risk the free-living population, we recommended the continued treatment of dormice showing tapeworm infection before release. Our work demonstrates how molecular techniques can be used to inform reintroduction programmes, reduce risk from disease and increase chances of reintroduction success.     

Using Long-Term Volunteer Records to Examine Dormouse ( Muscardinus avellanarius ) Nestbox Selection

Within ecology, there are unanswered questions about species-habitat interactions, which could potentially be resolved by a pragmatic analysis of a long-term volunteer-collected dataset. Here, we analysed 18 years of volunteer-collected data from a UK dormouse nestbox monitoring programme to determine the influence of habitat variables on nestbox choice by common dormice ( Muscardinus avellanarius ). We measured a range of habitat variables in a coppiced woodland in Gloucestershire, UK, and analysed these in relation to dormouse nestbox occupancy records (by dormice, other small mammals, and birds) collected by volunteers. While some characteristics of the woodland had changed over 18 years, simple transformation of the data and interpretation of the results indicated that the dataset was informative. Using stepwise regressions, multiple environmental and ecological factors were found to determine nestbox selection. Distance from the edge of the wood was the most influential (this did not change over 18 years), with boxes in the woodland interior being selected preferentially. There was a significant negative relationship with the presence of ferns (indicative of damp shady conditions). The presence of oak (a long-lived species), and the clumped structural complexity of the canopy were also important factors in the final model. There was no evidence of competition between dormice and birds or other mammals. The results provide greater understanding of artificial dormouse nest-site requirements and indicate that, in terms of habitat selection, long-term volunteer-collected datasets contribute usefully to understanding the requirements of species with an important conservation status.     

Prevalence of multiple mating by female common dormice, <Emphasis Type="Italic">Muscardinus avellanarius</Emphasis>

Mating behaviour is an important component of species’ life histories. Knowledge of natural patterns of mating can lead also to more effective management strategies for populations of conservation concern. Despite a high conservation profile many aspects of the biology of the common dormouse (Muscardinus avellanarius) remain unknown, potentially limiting present conservation efforts. We determine the mating behaviour of M. avellanarius at two woodland sites in the UK: (1) Bontuchel (a natural population in Wales) and (2) Wych (a population in England that was established by reintroducing captive-bred animals) by genotyping mothers and litters at a panel of 10 microsatellite loci. Adult female body weight positively correlates with litter size and no apparent reproductive skew was evident. We found that multiple mating by female dormice is prevalent at both sites, with litters containing three or more offspring sired by multiple fathers; moreover, multiple mating is adopted by released animals even after a period of captive breeding where females are mated singly or as a breeding pair. We also present evidence for low proportion of fathers identified in our samples that probably related to unsampled individuals and/or larger than anticipated population sizes. This first report of mating behaviour in M. avellanarius highlights the role of genetic studies to uncover species’ reproductive behaviours and include these data for conservation management.     

Protecting Free-Living Dormice: Molecular Identification of Cestode Parasites in Captive Dormice (Muscardinus avellanarius) Destined for Reintroduction

The success of any population translocation programme relies heavily on the measures implemented to control and monitor the spread of disease. Without these measures, programmes run the risk of releasing immunologically naïve species or, more dangerously, introducing novel infectious agents to native populations. As a precaution, a reintroduction programme for the common or hazel dormouse, Muscardinus avellanarius, in England screens dormice before release following captive breeding. Using PCR sequencing of a range of genes, we tested whether the same species of tapeworm(s) were present in captive and free-living dormice. Whilst only Rodentolepis straminea were identified in free-living dormice, cestode ova found in a captive individual produced a molecular match closely related to Hymenolepis microstoma and a previously unrecorded Rodentolepis species. To prevent putting at risk the free-living population, we recommended the continued treatment of dormice showing tapeworm infection before release. Our work demonstrates how molecular techniques can be used to inform reintroduction programmes, reduce risk from disease and increase chances of reintroduction success.

     

Take Only Photographs,Leave Only Footprints: Novel Applications of Non-Invasive Survey Methods for Rapid Detection of Small,Arboreal Animals

The development of appropriate wildlife survey techniques is essential to promote effective and efficient monitoring of species of conservation concern. Here, we demonstrate the utility of two rapid-assessment, non-invasive methods to detect the presence of elusive, small, arboreal animals. We use the hazel dormouse, Muscardinus avellanarius, a rodent of conservation concern, as our focal species. Prevailing hazel dormouse survey methods are prolonged (often taking months to years to detect dormice), dependent on season and habitat, and/or have low detection rates. Alternatives would be of great use to ecologists who undertake dormouse surveys, especially those assessing the need for mitigation measures, as legally required for building development projects. Camera traps and footprint tracking are well-established tools for monitoring elusive large terrestrial mammals, but are rarely used for small species such as rodents, or in arboreal habitats. In trials of these adapted methods, hazel dormice visited bait stations and were successfully detected by both camera traps and tracking equipment at each of two woodland study sites, within days to weeks of installation. Camera trap images and footprints were of adequate quality to allow discrimination between two sympatric small mammal species (hazel dormouse and wood mouse, Apodemus sylvaticus). We discuss the relative merits of these methods with respect to research aims, funds, time available and habitat.     

Long-term common dormouse monitoring: effects of forest management on abundance

The common dormouse (Muscardinus avellanarius) is a species included in Annex IV of the Habitat Directive and red-listed in many European countries. Monitoring of M. avellanarius is implemented in several countries using different methods. In Lithuania, regular control of standard bird nestboxes and marking of all dormice caught were used during 1984–1990 and 1999–2007. Nestboxes were spaced in a grid system at 50 m intervals in an area of 60 ha. Monitoring of the M. avellanarius population was carried out in a managed forest with varying levels of human activity occurring in most parts of the forest (felling of all understorey, selective felling of trees, clear felling, etc.). During the entire study period, the average density of the population was comparatively low (about 1 ind./ha in spring and 3 ind./ha in autumn), but stable and without substantial fluctuations in spite of considerable human activity. The forestry management operations that were used in different parts of the forest over different years had only temporary and localised negative effects on the abundance of M. avellanarius, and did not influence the whole population substantially. The spacing of nestboxes in a grid system in a large forest area increased the level of accuracy with which the monitoring scheme reflected the population status. This enabled both the state of the entire population of M. avellanarius and temporary changes in abundance in selected smaller plots to be tracked. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Surviving on the edge: a conservation-oriented habitat analysis and forest edge manipulation for the hazel dormouse in the Netherlands

We studied the effect of a forest edge management quasi-experiment on hazel dormouse (Muscardinus avellanarius) nest numbers in the reproductive season in the Netherlands. We found that the number of nests increased significantly from 2009 to 2013 in managed edges after clearing 75–100 % of mature trees, with a peak in the second year, whereas those in unmanaged edges decreased. In addition, we studied habitat characteristics in forest edges and hedgerows, and found a positive effect of a high upper shrub layer and a high abundance of blackberry (Rubus fruticosus agg. and Rubus spp.), rowan (Sorbus aucuparia) and common hawthorn (Crataegus monogyna) on nest densities. Based on these results, we propose a potential dormouse management strategy that extends conventionally advocated coppice management for regions with well-developed forest edges by (i) postulating a blackberry-oriented landscape restoration approach and (ii) using clearing of mature trees as a tool to boost local dormouse populations in forest edges that have become unsuitable as dormouse habitat due to vegetational succession.     

Long distance field crossings by hazel dormice (Muscardinus avellanarius) in fragmented landscapes

The hazel dormouse (Muscardinus avellanarius) is an arboreal species with suspected limited capabilities of moving over open ground. To date, however, only one study has investigated gap crossing capabilities in an experimental manner. We here report the results of an empirical assessment of hazel dormice gap crossing capabilities by means of a translocation study. We translocated 12 dormice, 10 in completely isolated patches and 2 in a hedgerow. Our results show how, at least under conditions of an experimental homing, hazel dormice may abandon forest areas and cross open fields (with grass or mowed) travelling up to 106 m. In most cases the gap crossing was relatively quick (concluded within one night) but in one case it lasted several days. The results of our experiment suggest that a stepping stone approach to connectivity may be a possible management strategy where it is not possible to implement a continuous network of hedgerows.     

Energy expenditure increases during the active season in the small,free-living hibernator Muscardinus avellanarius

Little is known about strategies employed by small mammals to reduce energy expenditure during the summer. To understand whether ambient conditions impact euthermic energy demands in a small free-living hibernator, we measured metabolic rate of hazel dormice (Muscardinus avellanarius) in the field. Furthermore, we aimed to reveal which variables influence torpor use. Our results show that hazel dormice altered euthermic energy expenditure during summer but not as expected as a response to environmental conditions. Euthermic resting metabolic rate was lowest directly after emergence from hibernation and increased by about 95% until the end of August. A considerable part of this increase was presumably caused by the changing influence of gender and rain on energy demands during different months, variation in food quality and quantity, and reversible size changes of organs that had been atrophied during hibernation. Torpor use in hazel dormice occurred more frequently when it was colder, earlier during the day, and in lighter individuals. Torpor was used routinely in males and non-reproductive females. We show that torpor is used more frequently than previously suggested by studies that only used visual proof of torpor use by surveying nest boxes.     

Nest box use by woodland dormice (Graphiurus murinus): the influence of life cycle and nest box placement

The use of nest boxes by the woodland dormouse, Graphiurus murinus, was investigated over a 13-month period in a riverine forest of the Great Fish River Reserve, South Africa. We predicted that some characteristics of nest box placement would affect nest box use and that the seasonal pattern of nest box use would be linked to the species' life cycle and physiological and socioecological characteristics. Generalized linear models indicated that the time since nest box installation and nest box height above ground positively affected the frequency and intensity of nest box use. Male and female dormice, as well as adults and juveniles, did not differ in the number of nest boxes used and equally occupied individual nest boxes. The percentage of nest boxes used peaked during spring and summer (breeding period) and dropped during winter (hibernation). However, whereas significantly more males were caught during the mating season (spring), the number of females occupying nest boxes was constant during the year. As female dormice successfully bred in the nest boxes, the observed sexual patterns suggest that (artificial) nest sites represent an important resource for females, whereas females seem to constitute the main resource for males, as predicted by the socioecological model.     

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.     

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.     

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.     

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.     

Footprint tunnels are effective for detecting dormouse species

Glirids are arboreal rodents that are difficult to monitor due to their nocturnal activity and their relatively low density. We compared results from footprint tunnels with those from two other monitoring methods (nest boxes and nesting tubes) for three dormouse species in an occupancy framework. Footprint tunnels performed better than the other two methods for the hazel dormouse Muscardinus avellanarius and the edible dormouse Glis glis, and were the only method which detected the garden dormouse Eliomys quercinus. Our findings provide evidence to support the effectiveness of footprint tunnels in monitoring the occurrence of dormouse species in comparison with other methods.     

Evidence of a complex phylogeographic structure in the common dormouse,Muscardinus avellanarius (Rodentia: Gliridae)

This is the first mitochondrial phylogeography of the common dormouse, Muscardinus avellanarius (Linnaeus, 1758), a hibernating rodent strictly protected in Europe (Habitat Directive, annex IV; Bern Convention, annex III). The 84 individuals of M. avellanarius, sampled throughout the distributional range of the species, have been sequenced at the mitochondrial DNA gene (cytochrome b, 704 base pairs). The results revealed two highly divergent lineages, with an ancient separation around 7.7 Mya and a genetic divergence of 7.7%. Lineage 1 occurs in Western Europe (France, Belgium, and Switzerland) and Italy, and lineage 2 occurs in Central–Northern Europe (Poland, Germany, Latvia, and Lithuania), on the Balkan Peninsula, and in Turkey. Furthermore, these two lineages are subdivided into five sublineages genetically isolated with a strong geographical association. Therefore, lineage 1 branches into two further sublineages (Western European and Italian), whereas lineage 2 contained three sublineages (Central–Northern European, Turkish, and Balkan). We observed low genetic diversity within the sublineages, in contrast to the significant level of genetic differentiation between them. The understanding of genetic population structure is essential for identifying units to be conserved. Therefore, these results may have important implications for M. avellanarius conservation. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 648–664.     

Spatial variation in population dynamics of the sand-burrowing amphipod Haustorioides japonicus

Densities of sandy beach organisms along coastlines often vary considerably even within small local scales. In order to understand the demographic basis of density variations in shore animals, we examined the population dynamics of the amphipod Haustorioides japonicus Kamihira, which is an ideal subject because it lacks a planktonic dispersal stage in its life history. Quantitative sampling at three adjacent sandy beaches along the Sea of Japan coast revealed that the spring density, i.e., the density of overwintered individuals, was similar among the three beaches. However, the amount of recruitment of newly hatched juveniles from late spring to summer greatly differed among the beaches, resulting in large spatial variations in the summer density. Cohort analyses revealed that at all beaches, young-of-the-year (YOY) individuals reproduced and then died, and that individuals which recruited in late summer overwintered. Moreover, growth, fecundity, and survival schedules of overwintered and YOY individuals were very similar among beaches. These observations suggest that the difference in recruitment success, i.e., the survival of newly hatched juveniles, is the primary cause of the inter-population variation in the summer density. It is likely that variations in salinity and sandy sediment affect the recruitment success of newly hatched individuals. This study indicates that biological productivity may differ substantially even among adjacent sandy beaches, with important implications for beach management.