Selective stripping of rowan (Sorbus aucuparia L.) bark by cattle in North-east Scotland

Summary

During winter 1973–4 beef cattle removed bark selectively and extensively from rowan growing in two Aberdeenshire woods. Some bark was also taken from Norway spruce and willow, but damage to all other species, including ash, beech, birch, gean, larch, Scots pine and sycamore, was negligible. In other woodlands utilised by cattle in this area bark-stripping was either less severe or absent. Possible causes for the bark-stripping are discussed. The rowan bark was not distinctive in cover, species of epiphytic lichen or inorganic composition, except perhaps for magnesium.     

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.     

Subspecific selectivity by a mammalian herbivore: Geographic differentiation of interactions between two taxa of Sciurus aberti and Pinus ponderosa

We compared host selection by two subspecies of Abert's squirrel, a highly specialized and selective herbivore. Both subspecies are restricted to stands of ponderosa pine, upon which they are ecologically dependent, but the two subspecies are geographically disjunct and each is closely associated with stands of trees that represent strongly differentiated chemical races. The criteria by which trees were selected as sources of phloem by each subspecies of squirrel included heritable features of xylem oleoresin and phloem. Trees that were potentially available to the subspecies of squirrel in Colorado differed substantially in biochemical features from trees that were available to the subspecies in Arizona; as a result, chemically mediated feeding patterns were distinct between the two squirrel subspecies. Based on multivariate analysis of chemical characteristics, trees utilized by the squirrels for feeding (target trees) differed significantly from control trees (non-target tre es) for each subspecies. However, the discriminant functions that separated target trees from non-target trees within a site generated different patterns of predicted classification when applied to trees at the other site. Vertebrate herbivores that are feeding specialists can exert selection pressures in populations of their host plants, and results suggest that geographically differentiated herbivore–host interactions can produce different sets of selection pressures, which may result in different evolutionary outcomes. Such geographic differentiation is a potentially important evolutionary aspect of chemically mediated mammal–plant interactions.     

A grey future for Europe: Sciurus carolinensis is replacing native red squirrels in Italy

Introduced mammals can cause extinction of native species due to replacement competition, disease, predation or hybridization. We studied the colonization of Piedmont (NW-Italy) by American grey squirrel (Sciurus carolinensis) and its effect on the native red squirrel (Sciurus vulgaris). Presence/absence data (2 × 2 km²), of both species were (re)constructed using questionnaires, literature, existing databases, unpublished information, and direct monitoring with hair-tubes. In 1970 red squirrels were still widespread and greys were restricted to forests near the introduction site. By 1990, grey squirrels had increased their range to 220 km², which coincided with the disappearance of native squirrels from 33 squares inside this range. The invasive species continued its spread occupying an area of 2,016 km² in 2010; within this area red squirrels went extinct in 88 squares. Overall, from 1970 to 2010 red squirrel went extinct in 62 % of 2 × 2 km² (ca. 1,689 km²), and were replaced by grey squirrels. The spread of the alien species was slow in the first 20 years, but doubled in the successive two decades. Nevertheless spread was slower than in Ireland and England. Grey squirrel adapt to climate and habitats in both North and South Europe, causing extinction of the native red squirrel. A EU LIFE co-funded project with the aim to control the grey squirrel in North Italy and recent trade-restrictions and trade-ban are a first step in reducing the risk of grey squirrels invading other countries, but their effectiveness will have to be evaluated.     

Landscape scale impacts of culling upon a European grey squirrel population: can trapping reduce population size and decrease the threat of squirrelpox virus infection for the native red squirrel?

The control of grey squirrels (Sciurus carolinensis) is widely undertaken as a conservation measure to protect red squirrel (Sciurus vulgaris) populations in the UK. However, inconsistencies and omissions in data collection, as well as fluctuating financial resourcing of control efforts, have meant that it has to date proved difficult to quantify the impact of any regional control initiative upon populations. Here we have scrutinized a 13 years period (1998–2010) within an ongoing grey squirrel control project that reflects the resource challenges typically faced by red squirrel conservation programmes. We present evidence that despite variation in grey squirrel control intensity, the abundance of grey squirrels ultimately decreased significantly. Trapping success was significantly higher in spring and summer months and a greater abundance of grey squirrels was found in deciduous woodland and hazel dominated scrub relative to other habitats; two findings that reinforce existing guidance within national control best practice. Grey squirrels carry an infection that causes epidemic pathogenic disease if spread to the native red squirrel. We observed that the proportion of seropositive grey squirrels decreased constantly from 2003 to 2010 when only 4 % of sampled animals were seropositive. This discovery indicates that culling can in parallel remove both the competitive and disease threat posed to red by grey squirrels. The historical paucity of scientific data on the effectiveness of grey squirrel control as a tactic in UK red squirrel conservation means that the findings of this study will significantly advance conservation best practice and inform the development of future national strategy.     

Habitat Use of Fox Squirrels in an Urban Environment

Abstract: Tree squirrels are one of the most familiar mammals found in urban areas and are considered both desirable around homes and, conversely, a pest. We examined fox squirrel (Sciurus niger) habitat use in inner city and suburban areas using radiotelemetry. We estimated habitat selection ratios at differing scales by season and fox squirrel activity. Telemetry data suggests that during periods of inactivity radiocollared fox squirrels (n = 82) selected 1) areas with greater tree canopy, 2) live oaks (Quercus fusiromis and Q. virginiana), and 3) trees with larger diameters and canopies. When inactive during the winter and spring, fox squirrels also preferred, within their core areas, to use the inside of buildings, and during periods of activity in the autumn and spring, fox squirrels preferred grassy areas. During periods of activity, fox squirrels avoided using pavement but did not exclude it from their core-area movements. Fox squirrels' ability to use buildings and to tolerate pavement in core-area movements make vast areas of the urban environment available to fox squirrels. In evaluating habitat variables that increased fox squirrel activity in urban areas, we found the number of large and medium trees, amount of pavement and grassy areas, canopy cover, number of oaks, and the area covered by buildings were all important factors in predicting fox squirrel activity in an urban environment. Our data suggests urban planners, animal damage control officials, wildlife managers, and landscapers who want to control urban fox squirrel populations through habitat manipulation should consider the reduction of oaks trees, a reduction of the canopy cover, and restricting the access of fox squirrels to buildings. Alternatively, home owners and squirrel enthusiasts hoping to bolster fox squirrel populations in urban areas should consider increasing the number of large mast—bearing trees and canopy cover and providing nest boxes.     

A predator's perspective of nest predation: predation by red squirrels is learned,not incidental

Nest predation has been used to explain aspects of avian ecology ranging from nest site selection to population declines. Many arguments rely on specific assumptions regarding how predators find nests, yet these predatory mechanisms remain largely untested. Here we combine artificial nest experiments with behavioural observations of individual red squirrels Tamiasciurus hudsonicus to differentiate between two common hypotheses: predation is incidental versus learned. Specifically, we tested: 1) whether nest survival could be explained solely by a squirrel's activity patterns or habitat use, as predicted if predation was incidental; or 2) if predation increased as a squirrel gained experience preying on a nest, as predicted if predation was learned. We also monitored squirrel activity after predation to test for evidence of two search mechanisms: area‐restricted searching and use of microhabitat search images. Contrary to incidental predation and in support of learning, squirrels did not find nests faster in areas with high use (e.g. forest edges). Instead, survival of artificial nests was strongly related to a squirrel's prior experience preying on artificial nests. Experience reduced nest survival times by over half and increased predation rates by 150–200%. Squirrels returned to and doubled their activity at the site of a previously preyed on nest. However, neither area‐restricted searching nor microhabitat search images can explain how squirrels located artificial nests more readily with experience. Instead, squirrels likely used cues associated with the nests or eggs themselves. Learning implies that squirrels could be increasingly effective predators as the density or profitability of nests increases. Our results add support to the view that nest predation is complex and broadly influenced (e.g. by predator experience, motivation), and is unlikely to be predicted consistently by simple relationships with predator activity, abundance or habitat.     

Forces structuring tree squirrel communities in landscapes fragmented by agriculture: species differences in perceptions of forest connectivity and carrying capacity

The North American red squirrel ( Tamiasciurus hudsonicus ) has expanded its range into the central hardwoods of the United States in conjunction with increasing forest fragmentation and declining gray squirrel ( Sciurus carolinensis ) populations. We used translocation experiments and patch occupancy data to test for interspecific differences in mobility and sensitivity to habitat loss and modification by agriculture. We released squirrels in fencerows to test the hypothesis that gray squirrels display inferior mobility relative to red and fox ( S. niger ) squirrels. Elapsed time to movement from fencerows for 76 individuals increased with distance to forest patches and harvesting of crops. Gray and red squirrels took longer to move from fencerows than fox squirrels, and gray squirrels were less successful at moving from fencerows than red and fox squirrels. Ecologically scaled landscape indices revealed the degree to which interspecific differences in mobility and individual area requirements accounted for the occurrence of these species across landscapes. Gray squirrel distribution was constrained both by individual area requirements and dispersal ability. Occurrence of red and fox squirrels was related to patch size but was unaffected by landscape connectivity.     

The Mechanism of Replacement of Red Squirrels by Grey Squirrels: A Test of the Interference Competition Hypothesis

Introduced American grey squirrels have replaced native red squirrels in most of the range currently occupied in Britain and northern Italy. The mechanisms of the replacement are not yet fully understood. We restated the commonly cited Interference Competition Hypothesis (ICH) that grey squirrels interfere with the behaviour of red squirrels in three possible ways: 1. by direct aggressive interactions; 2. by interrupting red squirrel mating-chases; or 3. by forcing red squirrels to actively avoid areas intensively used by grey squirrels. We compared the activity pattern, behaviour and reproductive performance of red squirrels in two study areas in northern Italy, one with only red squirrels (control area C1), the other with both species (experimental area E1). The following predictions were tested: 1. the total time spent in both intraspecific and interspecific interactions by red squirrels increases in the experimental area; 2. most interspecific interactions are aggressive, with grey squirrels being the dominant species; 3. the proportion of breeding female red squirrels that are unsuccessful at weaning offspring increases in area E1; 4. grey squirrels take part and interfere with red squirrel mating-chases, and thereby decrease the reproductive output of red squirrel females; 5. the activity pattern of red squirrels in the mixed-species area is shifted with respect to that in the control area to the hours of the day during which grey squirrels show little activity; and 6. red squirrels will shift their home range (or at least their core-area) when grey squirrel densities increase to avoid interspecific core-area overlap. Our results supported only the first prediction of the ICH: they failed to support all the other predictions. Moreover, the increase in the percentage of active time red squirrels spent interacting with other squirrels in the experimental study area was very small (only 1–2 min/day). Red squirrels did not avoid the woodland patches most intensively used by grey squirrels and the interspecific core-area overlap was similar to red squirrel intraspecific core-area overlap. This suggested that red squirrels avoided spatial overlap with grey squirrels in a similar manner as with conspecifics and that an increase in grey squirrel numbers will augment the intensity of resource competition. We therefore conclude that our results do not lend support to the Interference Competition Hypothesis and that interference competition by grey squirrels cannot explain the large-scale replacement of red by grey squirrels that has occurred in Britain and in Piedmont.     

Seasonal changes in the testis and epididymis of the American Grey squirrel, Sciurus carolinensis

The histological changes in the testis and epididymis of the American Grey squirrel, Sciurus carolinensis (Gmelin) during development and the testicular cycle are described. It was observed that male Grey squirrels, as a population, did not undergo a pronounced seasonal reproductive cycle, however, individual males did experience periods of reproductive inactivity which were asynchronous and not related to seasonal changes in the environment. It is suggested that the individual sexual cycles are due to an endogenous rhythm of gonadotrophin secretion by the pituitaries of these animals.     

Space invaders: effects of invasive alien Pallas’s squirrel on home range and body mass of native red squirrel

Alien species can affect native species through several ecological processes such as competition. Here we tested the hypothesis of interspecific competition for space and food resources between the native Eurasian red squirrel and the invasive Pallas’s squirrel introduced in Italy. We used an experimental study design comparing space and habitat use and body condition parameters of red squirrels between areas of co-occurrence with the Pallas’s squirrel and areas without it. There were no differences in mean home range size of red squirrels between red-only areas and red-Pallas. However, when Pallas’s squirrels were removed, the red squirrels increased their home ranges. Moreover, in the area of syntopy, red squirrels had a higher degree of intraspecific home range overlap than in the red-only area. We also found indirect evidence for competition for food with red squirrels having a poorer body condition when co-occurring with the alien species. We analyzed the body mass and size of red squirrels in the two areas and our results showed that red squirrels had a reduced body mass and size when in syntopy, confirming that the interspecific competition does not allow red squirrels to reach the optimum body condition that they would have if the competitor was not present. Moreover, tree-species niche overlap was very high and both species fed primarily on the same tree seeds. Differences in vegetation cover between areas are discussed. This is the first study that confirms the invasiveness of the Pallas’s squirrel also in terms of capability to compete with native species.     

Male Grey squirrel (Sciurus carolinensis) reproductive cycles in Britain

A synchronized seasonal cycle of sexual regression and development is described for the male Grey squirrel in Britain. Sexual development is maximal in December and May and minimal in August. The seasonal cycle can be overriden, but this is exceptional. Comparisons with previous work and the suggestion that the synchronized seasonal sexual cycle is general for Grey squirrels in Britain, and probably America, are made.     

Callosciurus squirrels: worldwide introductions,ecological impacts and recommendations to prevent the establishment of new invasive populations

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European red squirrel population dynamics driven by squirrelpox at a gray squirrel invasion interface

Infectious disease introduced by non‐native species is increasingly cited as a facilitator of native population declines, but direct evidence may be lacking due to inadequate population and disease prevalence data surrounding an outbreak. Previous indirect evidence and theoretical models support squirrelpox virus (SQPV) as being potentially involved in the decline of red squirrels (Sciurus vulgaris) following the introduction of the non‐native gray squirrel (Sciurus carolinensis) to the United Kingdom. The red squirrel is a major UK conservation concern and understanding its continuing decline is important for any attempt to mitigate the decline. The red squirrel–gray squirrel system is also exemplary of the interplay between infectious disease (apparent competition) and direct competition in driving the replacement of a native by an invasive species. Time series data from Merseyside are presented on squirrel abundance and squirrelpox disease (SQPx) incidence, to determine the effect of the pathogen and the non‐native species on the native red squirrel populations. Analysis indicates that SQPx in red squirrels has a significant negative impact on squirrel densities and their population growth rate (PGR). There is little evidence for a direct gray squirrel impact; only gray squirrel presence (but not density) proved to influence red squirrel density, but not red squirrel PGR. The dynamics of red SQPx cases are largely determined by previous red SQPx cases, although previous infection of local gray squirrels also feature, and thus, SQPV‐infected gray squirrels are identified as potentially initiating outbreaks of SQPx in red squirrels. Retrospective serology indicates that approximately 8% of red squirrels exposed to SQPV may survive infection during an epidemic. This study further highlights the UK red squirrel – gray squirrel system as a classic example of a native species population decline strongly facilitated by infectious disease introduced by a non‐native species. It is therefore paramount that disease prevention and control measures are integral in attempts to conserve red squirrels in the United Kingdom.     

The origins of bark-stripping by blue monkeys (Cercopithecus mitis): implications for management

Blue monkeys ( Cercopithecus mitis Wolf) on the Zomba Plateau, Malawi, damage exotic softwood plantations by bark-stripping. An ecological study was carried out to investigate the causes of this. The history of forestry practice on the Plateau reveals that, though bark-stripping did not start until 1976, plantation of exotic softwoods goes back to the beginning of the century. In the last 20 years the area of plantations close to the core of blue monkey habitat has increased substantially and some significant clearance of natural habitat has occurred. Bark-stripping is practised only by the groups which are in closest contact with the plantations. Its seasonality may be related to rainfall through sap quantity. There is no indication that the persistence of bark-stripping is the result of food shortage or agonistic encounters due to overcrowding. The most likely explanation for the genesis of bark-stripping is the encroachment of large blocks of plantations, combined with some clearance of natural habitat in 1973–74. The mechanism proposed for the initiation and maintenance of bark-stripping is a combination of temporary food shortage caused by travel through young plantations and a resultant acquired taste for sweet-tasting sap. Leaving areas of indigenous scrub adjacent to the core areas used by these primates and possibly the non-destruction of indigenous food trees within exotic plantations are suggested as ways of alleviating the problem in the future.     

Gliding locomotion of Siberian flying squirrels in low-canopy forests: the role of energy-inefficient short-distance glides

Short glides of less than 20 m seem energy inefficient for the Siberian flying squirrel Pteromys volans as with the northern flying squirrel Glaucomys sabrinus. However, Siberian flying squirrels in low-canopy forests frequently use short glides. Therefore, we sought to clarify the gliding patterns of Siberian flying squirrels for energy-efficient gliding transport in low-canopy forests (mean tree height, 15.3 m) in Hokkaido, Japan, based on records of 66 glides and 35 launch and landing trees. Mean launch height, landing height, and horizontal glide distance were 14.4, 2.7, and 21.4 m, respectively. For short distances, horizontal glide distance was strongly correlated with launch heights but not with launch tree height. For glides of more than 20 m, horizontal glide distance was significantly correlated with both launch height and launch tree height. The mean heights of launch and landing trees for short glides were 15.6 and 19.5 m, respectively. For long glides, these heights were 22.7 and 19.2 m. For short glides, mean launch tree height did not differ from overall mean tree height. However, for long glides, the mean launch tree height was greater than the overall mean tree height. Also, for short glides, the height of the landing tree was greater than that of the launch tree. Launch trees used for long glides were as high as the landing trees used in short glides. From these results, we conclude that Siberian flying squirrels in low-canopy forests save energy by gliding initially from a tree with sufficient height to permit a glide to a taller tree. This taller tree then permits long-distance glides that are energetically more efficient.     

Nest Site Selection by Western Gray Squirrels at Their Northern Range Terminus

ABSTRACT The western gray squirrel (Sciurus griseus) in Washington, USA, is limited to 3 disjunct areas and is a state threatened species. Information is lacking for the North Cascades population, which is the northernmost population for the species. Squirrels in this population exist without oaks (Quercus spp.) that provide forage and cavities for maternal nests elsewhere in their range. During May 2003 to August 2005, we studied selection of nest sites and nest trees by 18 radiocollared squirrels in Okanogan County, Washington. Without oak cavities, females reared their young in dreys. General nest-tree characteristics were similar to characteristics of western gray squirrel nest trees in Southeastern Cascades: relatively tall ponderosa pines (Pinus ponderosa) ≥ 40 cm diameter at breast height. Results from conditional logistic models determined that the odds of a squirrel selecting a tree for nesting increased with greater diameter at breast height and with infection by dwarf mistletoe (Arceuthobium spp.). Nest sites with high selection probability by squirrels had greater basal area and number of tree species than available unselected sites. Retention of forest patches that include a mix of conifer species or conifer and deciduous trees and moderate to high basal area could promote nesting opportunities, connectivity for arboreal travel, as well as abundance and diversity of hypogeous fungi. Experiments to test the efficacy of retaining untreated patches of varying size (including trees infected with mistletoe) on nesting by western gray squirrels within stands managed for fire suppression and forest health would provide important information about the effects of forest fuel management on arboreal wildlife.     

Effects of Logging Pattern and Intensity on Squirrel Demography

ABSTRACT We examined the effect of harvesting intensity and pattern on red squirrels (Tamiasciurus hudsonicus), northern flying squirrels (Glaucomys sabrinus), and yellow-pine chipmunks (Tamias amoenus) in mature inland Douglas-fir (Pseudotsuga menziesii glauca) forests in south-central British Columbia, Canada. We sampled squirrels 1 year before harvesting through 4 years after harvesting and estimated population parameters using open-population models. Relative to unharvested stands, each of the 3 species showed a strong response to tree removal. From 2 years to 4 years after logging, red squirrel density was 40% (SE = 7.1) lower in stands with 50% basal-area tree removal. From 1 year and up to 4 years after logging, northern flying squirrel density averaged 60% (SE = 5.2) lower in harvested treatments regardless of intensity or pattern of logging. In contrast, density of yellow-pine chipmunks increased markedly with increased logging intensity. Beginning 3 years after logging, yellow-pine chipmunk density was 734% (SE = 269) greater in stands with 50% basal-area tree removal. In the short term, harvesting intensity was a more important determinant of squirrel density than harvesting pattern. Retaining >10 m² per ha of live residual stand structure in mature inland Douglas-fir forests maintained habitat for forest-dependent species such as red squirrels and northern flying squirrels, albeit at lower densities.     

Managing anabolic steroids in pre-hibernating Arctic ground squirrels: obtaining their benefits and avoiding their costs

Androgens have benefits, such as promoting muscle growth, but also significant costs, including suppression of immune function. In many species, these trade-offs in androgen action are reflected in regulated androgen production, which is typically highest only in reproductive males. However, all non-reproductive Arctic ground squirrels, irrespective of age and sex, have high levels of androgens prior to hibernating at sub-zero temperatures. Androgens appear to be required to make muscle in summer, which, together with lipid, is then catabolized during overwinter. By contrast, most hibernating mammals catabolize only lipid. We tested the hypothesis that androgen action is selectively enhanced in Arctic ground squirrel muscle because of an upregulation of androgen receptors (ARs). Using Western blot analysis, we found that Arctic ground squirrels have AR in skeletal muscle more than four times that of Columbian ground squirrels, a related southern species that overwinters at approximately 0°C and has low pre-hibernation androgen levels. By contrast, AR in lymph nodes was equivalent in both species. Brain AR was also modestly but significantly increased in Arctic ground squirrel relative to Columbian ground squirrel. These results are consistent with the hypothesis that tissue-specific AR regulation prior to hibernation provides a mechanism whereby Arctic ground squirrels obtain the life-history benefits and mitigate the costs associated with high androgen production.