Long-term scatterhoarding by Eurasian red squirrels (Sciurus vulgaris)

Hoarding behavious, seed species hoarded and hoarding intensity by red squirrels ( Sclurus vulgaris L.) are described for two population; one in a coniferous woodland with few deciduous trees, the other in a mixed deciduous woodland with some conifers. Seasonal variation in the recovery of hoards and the importance of hoarded food with regard to a squirrel's annual energy demands are discussed. The importance of having hoards is compared among tree squirrel species.     

Dispersal and habitat cuing of Eurasian red squirrels in fragmented habitats

Animal dispersal and subsequent settlement is a key process in the life history of many organisms, when individuals use demographic and environmental cues to target post-dispersal habitats where fitness will be highest. To investigate the hypothesis that environmental disturbance (habitat fragmentation) may alter these cues, we compared dispersal patterns of 60 red squirrels (Sciurus vulgaris) in three study sites that differ in habitat composition and fragmentation. We determined dispersal distances, pre- and post-dispersal habitat types and survival using a combination of capture–mark–recapture, radio-tracking and genetic parentage assignment. Most (75%) squirrels emigrated from the natal home range with mean dispersal distance of 1,014 ± 925 m (range 51–4,118 m). There were no sex-related differences in dispersal patterns and no differences in average dispersal distance, and the proportion of dispersers did not differ between sites. In one of the sites, dispersers settled in patches where density was lower than in the natal patch. In the least fragmented site, 90% of animals settled in the natal habitat type (habitat cuing) against 44–54% in the more strongly fragmented sites. Overall, more squirrels settled in the natal habitat type than expected based on habitat availability, but this was mainly due to individuals remaining within the natal wood. In the highly fragmented landscape, habitat cuing among emigrants did not occur more frequently than expected. We concluded that increased habitat fragmentation seemed to reduce reliable cues for habitat choice, but that dispersing squirrels settled in patches with lower densities of same-sex animals than at the natal home range or patch, independent of degree of fragmentation.     

Food choice of Eurasian red squirrels and concentrations of anti-predatory secondary compounds

Secondary metabolites (SMs) are organic compounds of low molecular mass that represent a vast chemical diversity. In plants, one of their preeminent roles is their repellent activity against predators. The Eurasian red squirrels (Sciurus vulgaris) is one of the most important predators of conifer tree seeds in boreal and alpine forests in Europe. Its population dynamics and space use are strongly affected by the size of Norway spruce (Picea abies) seed-crops, but not by silver fir (Abies alba) seed-crops. Moreover, squirrel heavily feed on spruce seeds but tend to avoid fir seeds, although the latter has a higher seed-energy content per cone. We tested the hypothesis that a higher concentration of some SMs in fir than in spruce seeds and/or cone scales, the protective tissue of seeds, was related with squirrel feeding preferences. We determined terpene concentrations in the cyclohexane extract of seeds and scales by gas-chromatography–electron ionisation mass-spectrometry (GC/EI-MS), and measured the protein precipitation activity of tannin contained in spruce and fir seeds and scales. Of the nearly 300 separated chemical entities, only limonene, α- and β-pinene and myrcene occurred in all samples and their levels accounted for 80% or more of all cyclohexane-extractable materials of fir and for less than 50% of that of spruce. In fir, limonene was by far the most abundant compound, and fir scales had a 13 times higher concentration of α-pinene than spruce scales. Fir seeds had much higher limonene concentrations than fir scales and than seeds or scales of spruce. Tannin concentrations were higher in cone scales than in seeds, with no differences between the two tree species. This study suggests that high concentrations of limonene might reduce the feeding activity of red squirrels on fir seeds, and that the preferred spruce seeds had much lower monoterpene concentrations. Feeding trials using food items treated with different concentrations of limonene will be carried out to confirm this hypothesis.     

凉水自然保护区松鼠巢址选择的特征

2006 年6 月11 日至9 月27 日和2007 年8 月14 日至9 月21 日采用样线法,并结合定位观察的方法在凉水国家级自然保护区对松鼠的巢址选择进行了定量研究,共获取107 个巢样方和230 个对照样方。对每个样方分别测定了12 个巢树参数和7 个环境参数。统计分析结果表明,松鼠仅在针叶树上营巢,59.8 ± 4. 7% 的巢建于红松上,但对枝叶比红松更为浓密的云杉和臭冷杉有显著的选择性,而且显著倾向于在以云杉、臭冷杉为优势树种的生境中营巢。松鼠选择比较高大(树高21.5 ± 0.4 m)、活动通道比较多(3.6 ±0.1)的树营巢,巢树周围的树木密度(169 ± 4 株/ hm² )和多样性(辛普森指数0.51 ± 0.01)显著高于对照样方。巢址生境多位于南向的中、下坡位。巢址高12.4 ± 0.4 m,紧贴树干,多朝南。红松树上的巢多建于树冠的中下部,云杉和臭冷杉树上的巢多建于树冠的上部。松鼠巢对包括道路在内的林隙没有显著的回避。     

Effects of spatial variation in food availability on spacing behaviour and demography of Eurasian red squirrels

In heterogeneous habitats with limited resources, spacing behaviour will affect individual variation in breeding success and density of populations, and is thus of general interest to ecologists. We investigated how red squirrels Sciurus vulgaris adapt their social organisation to tine‐grained heterogeneity in habitat quality, studying spacing behaviour, habitat use and population dynamics in a forest in north Italy, characterised by a mosaic of high‐quality (chestnut‐pine) and poor‐quality patches. We compared the data with those from more homogeneous broadleaf and mixed woodlands with similar overall tree seed abundance (“stable” habitats). Squirrels lived at lower densities (pre‐breeding density 0.39‐0.58 ha^‐1) than in “stable” habitats, although breeding rate was not reduced. Female breeding success was related to being primiparous as yearlings, and increased with body mass and proportion high‐quality habitat in the home range. Persistency rate of females was as in stable habitats, It was higher than male persistency, but immigration and recruitment rates were male biased, resulting in even sex‐ratio. All residents occupied high‐quality patches, and no subadults established a permanent home range in poor‐quality habitat. Home range and core‐area size was typically larger in males than in females and a male's core‐area overlapped those of other males and of females. Female core‐areas were overlapped by males but not. or very little, by other females (intrasexual territoriality). Home ranges, or core‐areas, were not smaller than in “stable” habitats, nor did we find a higher degree of core‐area overlap. We conclude that in patchy habitats dominant, resident red squirrels exclude dispersing animals from preferred, high‐quality habitat, producing a spacing pattern referred to as ideal despotic distribution, and that poor patches were only used temporarily by transient individuals, resulting in a reduction of density in comparison to populations in “stable” habitats.     

Competition between Eurasian red and introduced Eastern grey squirrels: the energetic significance of body-mass differences

Daily energy expenditure (DEE) was measured in sympatric populations of red and grey squirrels using the doubly labelled water technique. Grey squirrels had significantly higher DEEs than red squirrels. However, the difference between the species was not separable from the effects of body mass on DEE. The DEEs of both species were in accordance with published allometric predictions incorporating body mass and ambient temperature. The differences in energetic requirements and social dominance, both consequences of body size, may represent means by which grey squirrels exert more interspecific competition on red squirrels than do conspecifics, potentially driving populations below viable levels in some sites.     

Interspecific effects of grey squirrels (Sciurus carolinensis) on the space use and population demography of red squirrels (Sciurus vulgaris) in conifer plantations

Interspecific competition between red squirrels and grey squirrels was investigated by comparing the population demography, spacing behavior and habitat use of red squirrels in two large conifer plantations in northern England: one site had only red squirrels (the red-only site), in the other both red and grey squirrels occurred (the red–grey site). Despite more abundant food at the red–grey site, red squirrel densities (0.26 ha^–1 at the red–grey site, 0.29 ha^–1 at the red-only site), adult survival rates and the breeding rates of females were similar at both study sites. Grey squirrels at the red–grey site occurred at higher densities (0.92–1.1 ha^–1) than did the reds and tended to have higher breeding rates. In the presence of grey squirrels, the recruitment pattern of red squirrels changed and there was little recruitment of subadults. The juvenile recruitment rate in the red–grey site (13%) was much lower than in the red-only site (50%). Grey squirrels, in contrast, had higher juvenile recruitment rates at the red–grey site (41%). The core areas of the home ranges of red squirrels in the red–grey site were more strongly overlapped by grey squirrels than by conspecifics. Red squirrels did not select the habitat with the best tree seed crop (Scots pine) but preferred dense Sitka spruce plantations; they appeared to avoid the Scots pine area with its high grey squirrel density. Data on foot length and body condition indicated decreased body growth in young red squirrels when grey squirrels were present. Our data suggest that adult red squirrels suffered little from interspecific competition with grey squirrels and that the key factor is decreased juvenile recruitment in red squirrels.     

Acute fatal toxoplasmosis in three Eurasian red squirrels (Sciurus vulgaris) caused by genotype II of Toxoplasma gondii

Toxoplasma gondii parasites belonging to endemic genotype II caused fatal infection in three (16%) of 19 Eurasian red squirrels (Sciurus vulgaris) sent for necropsy in Finland between May 2006 and April 2009. The liver, spleen, and lungs were the organs most affected in all three cases, and high numbers of T. gondii parasites were visualized immunohistochemically in all the tissue samples available from them. The genotyping of the parasite strains was based on the results of analysis of length polymorphism at six microsatellite markers (B18, TUB2, TgM-A, W35, B17, and M33). The length of the PCR product at the additional seventh marker (M48) was 233 base pairs from the first two cases that were found dead together, suggesting a common infection source, and 215 base pairs from the third. Eurasian red squirrels may be exceptionally susceptible to T. gondii infection.     

Contrasting stress response of male arctic ground squirrels and red squirrels

A hormonal-challenge protocol was used to compare the stress response of males of Arctic ground squirrels and red squirrels during the breeding season (May). These squirrels live in the same boreal forest of the Yukon, but have very different life histories and utilize the forest in markedly different ways. Red squirrels had levels of total cortisol, maximum corticosteroid-binding capacity, and free cortisol that were 5, 7, and 2 times, respectively, those of Arctic ground squirrels. Red squirrels were resistant to suppression by an artificial glucocorticoid, dexamethasone (DEX); Arctic ground squirrels were not. Cortisol levels in red squirrels responded slowly but continuously to the ACTH injection; Arctic ground squirrels responded rapidly and then stabilized. Testosterone levels in red squirrels were extremely sensitive to the challenge, being suppressed by both DEX and ACTH; levels in Arctic ground squirrels were resistant to the challenge, being modestly suppressed by DEX and stimulated by ACTH. Energy mobilization, as measured by glucose and free fatty acid responses, was not affected. Red squirrels had four times the levels of white blood cells and higher proportions of lymphocytes and lower proportions of eosinophils than Arctic ground squirrels, indicating that the latter were in worse condition immunologically. Our evidence suggests that the functions associated with the hypothalamic-pituitary-adrenal axis are compromised in breeding male Arctic ground squirrels, but not in red squirrels. We propose that in male red squirrels this axis has evolved in the context of a stable social system based on long-lived animals with individual territories which are needed to deal with unpredictable winter food supplies. In contrast, Arctic ground squirrels escape the rigors of winter by hibernation and this hormonal axis has evolved in short-lived males in the context of intense intra-sexual competition in a social system based on female kin groups and regular male dispersal to avoid inbreeding. J. Exp. Zool. 286:390-404, 2000.     

The effects of hoarding habitat selection of Eurasian red squirrels (Sciurus vulgaris) on natural regeneration of the Korean pines

Cone-cores discarded by Eurasian red squirrels were used to study the habitat selection of Korean pine-seeds hoarding, in forest patch Nos. 16 and 19 in Liangshui Nature Reserve, China. Ten transects with a total length of 15 km were uniformly set, and data from 343 valid samples were collected in a 369 hm² area. One hundred and eighty four were hoarding samples which were determined according to the cluster analysis based on the number of the cone-cores, while the other 159 were control samples. The principal component analysis, using 11 habitat factors, suggested that the distance from Korean pine forest, forest type, number of Korean pine seedlings, density and type of bush significantly influenced the habitat selection of hoarding by Eurasian red squirrels. The results of Bailey’s method indicated that the squirrels showed (1) preference for natural coniferous forest, natural fir and spruce forest and planted spruce forest; (2) avoidance of planted Korean pine forest and planted larch forest; and (3) random use of natural Korean pine forest. Moreover the distance from the Korean pines in the range of 150–600 m showed no effect on the habitat selection of hoarding by the Eurasian red squirrels. More than 50% of the cone-cores were discarded in either fringe or gap of the Korean pine forest with more cone-cores found at <300 m than at 300 m away (One-Way ANOVA; df = 3, 183, F = 5.76, p = 0.0009). This demonstrated that the Eurasian red squirrels could take the cone-cores out of the Korean pine forest. The density of bushes in samples of hoarding area was significantly lower than that in control samples (Kruskal–Wallis test; df = 1, χ² = 83.99, p < 0.0001). The number of the Korean pine seedlings in samples of hoarding area was significantly higher than that in the control samples (Kruskal–Wallis test; df = 1, χ² = 104.13, p < 0.0001). This illustrated that the hoarding habitat favoured the germination of the Korean pine seedlings. In conclusion the behavior of hoarding Korean pine seeds by the Eurasian red squirrels can promote the regeneration and dispersal of the Korean pines.     

Geographical distance and physical barriers shape the genetic structure of Eurasian red squirrels (Sciurus vulgaris) in the Italian Alps

Red squirrels (Sciurus vulgaris) are widely distributed throughout Eurasia, occurring in many types of coniferous and mixed-deciduous forests. In fragmented landscapes, small and partly isolated populations with low immigration rates show reduced genetic diversity, but reforestation can increase gene flow and restore levels of genetic variation in a few decades. No studies have so far investigated the genetic structure of red squirrel in large, continuous forests. The Italian Alps are presently characterized by almost continuous, recently reconnected forest habitats, that were affected by deep landscape changes during last glaciations but remained mostly unchanged between 10 000 and 200 years bp, when forest cover was heavily reduced. In this study we analyse patterns of genetic variability of red squirrels in and between seven sites distributed over 250 km of Alpine habitat, using mitochondrial DNA (mtDNA) and microsatellites. We use isolation-by-distance (IBD) models to investigate the relative importance that past (Pleistocene glaciations) and recent (fragmentation, bottlenecks) events had on the present genetic situation. Both nuclear and mtDNA data indicate a significant differentiation among study sites and a significant correlation between genetic and geographical distance only over a large scale. No recent bottlenecks are recorded through microsatellites and demographic models strongly support equilibrium between gene flow and drift; however, mtDNA suggests that there may have been local demographic crashes, probably in correspondence with the 19th-century forest fragmentation. These findings indicate that local landscape factors other than geographical distance per se, such as barriers of unsuitable habitat, affect gene flow and determine differentiation.     

Making red squirrels more visible: the use of baited visual counts to monitor populations

Understanding changes in the distribution and abundance of the red squirrel Sciurus vulgaris in Europe, as a result of the spread of the introduced North American grey squirrel Sciurus carolinensis, is vital to planning and implementing red squirrel conservation measures. We studied the effectiveness of a variation on a common method of monitoring squirrels, standardised visual counts that made use of a diffuse baiting approach to increase their detectability. Significantly more sightings of squirrels occurred on baited than on unbaited visual transect lines in trials with the assistance of volunteers, and we recommend the use of baits in future studies.     

What future for British red squirrels?

The geographic range of red squirrels contracted sharply in Britain during the 1940s arid 1950s, as increasingly large areas were colonized by the congeneric North American grey squirrel. Red squirrels remain common only on offshore islands, and in the large conifer forests of northern England and Scotland. The initial replacement of red squirrels was in arras dominated by oak woodland, probably because acorn crops are exploited less efficiently by red squirrels than by grey squirrels. Dirt studies have shown that acorns are digested less efficiently by the red squirrel, which occurs in conifers through most of its Eurasian range, than by the introduced grey squirrel, which is primarily a native of deciduous woodland. The red squirrel will probably be replaced in deciduous and mixed woodland throughout mainland Britain, and may eventually persist only in large areas of conifers which arc far from oak trees. The conservation of red squirrels on islands is therefore particularly important for their survival, perhaps making it worthwhile to create new island populations where they do not at present exist.     

The effects of hoarding habitat selection of Eurasian red squirrels (Sciurus vulgaris) on natural regeneration of the Korean pines

Cone-cores discarded by Eurasian red squirrels were used to study the habitat selection of Korean pine-seeds hoarding, in forest patch Nos. 16 and 19 in Liangshui Nature Reserve, China. Ten transects with a total length of 15 km were uniformly set, and data from 343 valid samples were collected in a 369 hm² area. One hundred and eighty four were hoarding samples which were determined according to the cluster analysis based on the number of the cone-cores, while the other 159 were control samples. The principal component analysis, using 11 habitat factors, suggested that the distance from Korean pine forest, forest type, number of Korean pine seedlings, density and type of bush significantly influenced the habitat selection of hoarding by Eurasian red squirrels. The results of Bailey’s method indicated that the squirrels showed (1) preference for natural coniferous forest, natural fir and spruce forest and planted spruce forest; (2) avoidance of planted Korean pine forest and planted larch forest; and (3) random use of natural Korean pine forest. Moreover the distance from the Korean pines in the range of 150–600 m showed no effect on the habitat selection of hoarding by the Eurasian red squirrels. More than 50% of the cone-cores were discarded in either fringe or gap of the Korean pine forest with more cone-cores found at <300 m than at 300 m away (One-Way ANOVA; df = 3, 183, F = 5.76, p = 0.0009). This demonstrated that the Eurasian red squirrels could take the cone-cores out of the Korean pine forest. The density of bushes in samples of hoarding area was significantly lower than that in control samples (Kruskal–Wallis test; df = 1, χ² = 83.99, p < 0.0001). The number of the Korean pine seedlings in samples of hoarding area was significantly higher than that in the control samples (Kruskal–Wallis test; df = 1, χ² = 104.13, p < 0.0001). This illustrated that the hoarding habitat favoured the germination of the Korean pine seedlings. In conclusion the behavior of hoarding Korean pine seeds by the Eurasian red squirrels can promote the regeneration and dispersal of the Korean pines.