Return of Fire as a Restoration Tool: Long‐Term Effects of Burn Severity on Habitat Use by Mexican Fox Squirrels

After decades of suppression, fire is returning to forests of the western United States through wildfires and prescribed burns. These fires may aid restoration of vegetation structure and processes, which could improve conditions for wildlife species and reduce severe wildfire risk. Understanding response of wildlife species to fires is essential to forest restoration because contemporary fires may not have the same effects as historical fires. Recent fires in the Chiricahua Mountains of southeastern Arizona provided opportunity to investigate long‐term effects of burn severity on habitat selection of a native wildlife species. We surveyed burned forest for squirrel feeding sign and related vegetation characteristics to frequency of feeding sign occurrence. We used radio‐telemetry within fire‐influenced forest to determine home ranges of Mexican fox squirrels, Sciurus nayaritensis chiricahuae, and compared vegetation characteristics within home ranges to random areas available to squirrels throughout burned conifer forest. Squirrels fed in forest with open understory and closed canopy cover. Vegetation within home ranges was characterized by lower understory density, consistent with the effects of low‐severity fire, and larger trees than random locations. Our results suggest that return of low‐severity fire can help restore habitat for Mexican fox squirrels and other native wildlife species with similar habitat affiliations in forests with a historical regime of frequent, low‐severity fire. Our study contributes to an understanding of the role and impact of fire in forest ecosystems and the implications for forest restoration as fire returns to the region.     

Fox Squirrel Survival in Urban and Rural Environments

Abstract A number of studies on mammalian species that have adapted to urban areas suggest survival may be higher for urban populations than rural populations. We examined differences in fatalities between an urban and rural population of fox squirrels (Sciurus niger). We radiocollared (n = 50 rural, n = 78 urban) fox squirrels during approximately 2 years. We found monthly survival of rural fox squirrels (Ŝ = 0.936) was lower than urban fox squirrels (Ŝ = 0.976) over the same 12-month period. Nonetheless, when comparing a 24-month period of survival data on urban squirrels with an 18-month period on the rural squirrels (periods overlapped for 12 months), survival rates were more similar between urban (Ŝ = 0.938) and rural squirrels (Ŝ = 0.945). Our data suggest that sex and season may influence survival of urban squirrels and not rural squirrels. We also found that cause of fatalities differed between the urban and rural squirrels, with >60% of fatalities on the rural site caused by predation. In contrast, <5% of the fatalities on the urban site were caused by predation and >60% of urban fox squirrel fatalities were caused by motor vehicle collisions. This study illustrates the need to advance our ability to understand, predict, and mitigate effects of urbanization on wildlife resources.     

Differential Resource Use between Native and Introduced Gray Squirrels

Differences in habitat requirements and the spatial distribution of habitat for native and introduced species can determine outcomes of biological invasions. Introduced eastern gray squirrels (Sciurus carolinensis) have displaced native red squirrels (S. vulgaris) in Europe and have been implicated as a contributing factor to the decline of western gray squirrels (S. griseus) in North America. Eastern and western gray squirrels are associated with oak (Quercus spp.), but little is known about how these species interact. From April 2007 to April 2012, we radio-tracked sympatric eastern and western gray squirrels in western Washington to compare habitat use and evaluate competitive interactions. We developed resource utilization functions for each species and evaluated distribution of habitat on Joint Base Lewis-McChord, Washington, USA. Both species shared affinity for closed canopy forests and oaks; however, important differences in relationships with riparian areas and shrub cover resulted in low overlap in habitat distribution, which likely limited potential competitive interactions. Eastern gray squirrels appeared restricted to areas around wetlands likely because they supported the deciduous tree species that comprise habitat for this species in its native range. Use by western gray squirrels, but not eastern gray squirrels, significantly decreased with increasing shrub cover. Forestry practices that promote mixed oak-conifer with little shrub cover in uplands can benefit western gray squirrels and minimize interactions with eastern gray squirrels. © 2020 The Wildlife Society.     

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.     

Winter Versus Summer Habitat Selection in a Threatened Ground Squirrel

Hibernation is a strategy many species employ to survive periods of thermal stress or resource shortage (e.g., harsh thermal conditions, food limitations) and habitat requirements of hibernating species may differ between summer (the active season) and winter (during hibernation). Accounting for seasonal differences in habitat affinities will help ensure that management actions are more beneficial and land-use policies are more appropriate. The northern Idaho ground squirrel (Urocitellus brunneus) is a federally listed threatened species that is in decline and hibernates for approximately 8 months per year. We collared northern Idaho ground squirrels in Adams County, Idaho from 2013–2017. The majority of northern Idaho ground squirrels we collared selected hibernacula outside of the areas they used during the active season. Furthermore, habitat features of hibernacula locations differed from habitat features of active-season areas. Hibernacula locations had greater canopy closure compared to active-season locations (36.9% and 7.0% canopy closure, respectively) and hibernaculum habitat features (particularly distance to nearest log) influenced overwinter survival. Our results suggest that recovery efforts for northern Idaho ground squirrels should include protection and management for the full range of habitat conditions used throughout summer and winter. More broadly, we emphasize the need to identify and protect habitat during all seasons because habitat requirements can differ substantially during different portions of an animal's annual cycle and effective conservation will require management of year-round habitat needs. © 2020 The Wildlife Society.     

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.     

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.     

A systematic review into the suitability of urban refugia for the Eurasian red squirrel Sciurus vulgaris

1. Urban growth and intensification are projected to increase as the global human population increases. Historically, urban areas have been disregarded as suitable wildlife habitat, but it is now known that these areas can be biodiverse and that wildlife species can adapt to the environmental conditions. One such urban-dwelling species is the Eurasian red squirrel Sciurus vulgaris, which has suffered population declines in several countries throughout its range in recent decades.
2. The current published literature was systematically reviewed to determine whether or not urban habitats are suitable refugia for red squirrels, through identifying and discussing key topics regarding the urban ecology of red squirrels.
3. Urban environments can support higher population densities of red squirrels than rural areas, probably due to the widespread and reliable provision of anthropogenic supplemental food alongside natural food sources. The availability and quality of urban greenspaces are important determinants of the suitability of urban habitats for red squirrels, as they provide natural food sources and nesting sites. Despite the barriers present in urban landscapes (e.g. roads), red squirrels can still disperse and maintain gene flow at the population level.
4. Road traffic accidents appear to be a significant cause of mortality in some urban red squirrel populations, and seasonal peaks of mortality occur during the autumn months. Diseases (e.g. squirrelpox virus) can also be a significant cause of mortality, although effects differ between populations and depend on whether grey squirrels Sciurus carolinensis are present. Many of the predation events that affect red squirrels appear to be due to free-ranging domestic and feral cats Felis catus, although there is currently little evidence to suggest that predation is a limiting factor for urban red squirrel populations.
5. We conclude that urban areas can be suitable refugia for red squirrels, provided that high-quality greenspaces are maintained. Mitigation measures may also be necessary to reduce population mortality and to prevent disease outbreaks.

     

Importance of scale in nest-site selection by Arizona gray squirrels

Nests provide a place for individuals to rest, raise young, avoid predators, and escape inclement weather; consequently, knowledge of habitat characteristics important to nest placement is critical for managing species of conservation concern. Arizona gray squirrels (Sciurus arizonensis) are endemic to mountains of southwestern United States and northwestern Mexico. We investigated factors influencing nest-site selection at 4 spatial scales (forest-type, nest-site, nest-tree, and within-canopy placement) to provide ecological information and management recommendations for this sensitive species. Nest densities were 2.6 times higher in riparian than pine-oak woodlands. Nest sites had more large trees, snags, logs, and canopy cover and had lower slope. Arizona gray squirrels selected tall trees with more interlocking trees and tended to place nests adjacent to the main trunk. Regardless of scale, Arizona gray squirrels seemed to select nesting areas for their ability to provide protection from predators and the elements as well as access to food. Consequently, maintaining large trees with closed canopies and downed logs should be considered when determining land management plans. © 2011 The Wildlife Society.     

Fine-Scale Analysis of Mount Graham Red Squirrel Habitat Following Disturbance

ABSTRACT Habitat destruction and degradation are major factors in reducing abundance, placing populations and species in jeopardy. Monitoring changes to habitat and identifying locations of habitat for a species, after disturbance, can assist mitigation of the effects of human-caused or -amplified habitat disturbance. Like many areas in the western United States, the Pinaleño Mountains of southeastern Arizona, USA, have suffered catastrophic fire and large-scale insect outbreaks in the last decade. The federally endangered Mt. Graham red squirrel (Tamiasciurus hudsonicus grahamensis) is only found in the Pinaleño Mountains, and to assess effects of forest disturbance on habitat we modeled their potential habitat by identifying characteristics of cover surrounding their centrally defended middens. We classified high-spatial resolution satellite imagery into ground cover classes, and we used logistic regression to determine areas used by squirrels. We also used known midden locations in conjunction with slope, elevation, and aspect to create a predictive habitat map. Squirrels selected areas of denser forest with higher seedfall for midden sites. Among active middens, those in the densest and least damaged forests were occupied in more seasons than those in more fragmented and damaged areas. The future conservation of red squirrels and the return of healthy mature forests to the Pinaleño Mountains will rely on preservation of mixed conifer zones of the mountain and active restoration of spruce-fir forests to return them to squirrel habitat. Our ability to evaluate the spectrum of fine- to coarse-scale disturbance effects (individual tree mortality to area wide boundaries of a disturbance) with high-resolution satellite imagery shows the utility of this technique for monitoring future disturbances to habitat of imperiled species.     

Winter Roosting Ecology of Eastern Red Bats in Southwest Missouri

Abstract: Eastern red bats (Lasiurus borealis) have been found to overwinter in areas that can experience severe fluctuations in temperature. We examined the red bat's use of winter roosts in southwest Missouri, USA, for 2 winters (2003–2005). We found tree roosts in eastern red cedars (Juniperus virginiana) and hardwoods. Tree roost sites were located on the south side of trees, and we found roost trees on south-facing slopes. Roost sites occurred more frequently in the location with least canopy cover. Bats switched from tree roosts to leaf litter roosts when ambient temperatures approached or fell below freezing. We found habitat characteristics and aspect to be determining factors in the selection of leaflitter roosts. Management of overwintering red bats requires a diverse forest structure, including canopy gaps, stand-density variation, and leaf-bearing trees, including oaks (Quercus spp.).     

Characteristics of Mount Graham Red Squirrel Nest Sites in a Mixed Conifer Forest

Abstract: The Mount Graham red squirrel (Tamiasciurus hudsonicus grahamensis) is constrained to the Pinaleño Mountains in southeastern Arizona, USA. The population's endangered status and extensive forest damage from insects and fire warrants a better understanding of habitat variables important for nest site selection. We examined characteristics of cavity (n = 91) and drey (n = 38) nests and compared these to random sites (n = 113). Dreys were found primarily in Engelmann spruce (Picea engelmannii) and corkbark fir (Abies lasiocarpa var. arizonica). Cavity nests occurred primarily in aspen (Populus tremuloides) and corkbark fir. Squirrels selected nest sites with higher canopy cover and more corkbark fir, decayed logs, and living trees. Forest management plans emphasizing thinning must consider how altering these habitat characteristics could affect availability and suitability of tree stands for nesting squirrels.     

Diagnostic Genetic Marker That Differentiates Eastern Fox Squirrels From Eastern Gray Squirrels

Abstract The Delmarva fox squirrel (Sciurus niger cinereus) has been listed as endangered by the United States Department of Interior since 1967. A high-priority task for the recovery of this taxon is to determine its current geographic distribution. Toward this end, we have identified a microsatellite locus that unambiguously differentiates Delmarva fox squirrels from eastern gray squirrels (S. carolinensis), which frequently co-occur with Delmarva fox squirrels. Analysis of this marker in noninvasively collected hair samples will allow unequivocal identification of localities occupied by Delmarva fox squirrels with a minimum investment of funds, time, and effort because handling individuals will be unnecessary. This protocol will expedite site review in connection with the Endangered Species Act consultation process.     

Relative Roles of Grey Squirrels,Supplementary Feeding,and Habitat in Shaping Urban Bird Assemblages

Non-native species are frequently considered to influence urban assemblages. The grey squirrel Sciurus carolinensis is one such species that is widespread in the UK and is starting to spread across Europe; it predates birds’ nests and can compete with birds for supplementary food. Using distance sampling across the urbanisation intensity gradient in Sheffield (UK) we test whether urban grey squirrels influence avian species richness and density through nest predation and competition for supplementary food sources. We also assess how urban bird assemblages respond to supplementary feeding. We find that grey squirrels slightly reduced the abundance of breeding bird species most sensitive to squirrel nest predation by reducing the beneficial impact of woodland cover. There was no evidence that grey squirrel presence altered relationships between supplementary feeding and avian assemblage structure. This may be because, somewhat surprisingly, supplementary feeding was not associated with the richness or density of wintering bird assemblages. These associations were positive during the summer, supporting advocacy to feed birds during the breeding season and not just winter, but explanatory capacity was limited. The amount of green space and its quality, assessed as canopy cover, had a stronger influence on avian species richness and population size than the presence of grey squirrels and supplementary feeding stations. Urban bird populations are thus more likely to benefit from investment in improving the availability of high quality habitats than controlling squirrel populations or increased investment in supplementary feeding.     

Scavenging affects persistence of avian carcasses resulting from window collisions in an urban landscape

ABSTRACT Collisions with windows remain an important human‐related threat to bird survival in urban landscapes. Accurately estimating the magnitude of avian mortality at windows is difficult and may be influenced by many sources of error, such as scavenging of carcasses. Failure to account for removal of carcasses by scavengers can bias estimates of window mortality. We tested the hypothesis that carcass survival depends on local habitat factors known to influence scavenger behavior. Scavenger activity on bird carcasses was documented at 20 buildings in an urban landscape in northwestern Illinois for 1 week during each season of a year. Known‐fate models were used to relate carcass survival to local habitat composition and to evaluate temporal variation in survival. We also documented species of scavengers and the timing of scavenging using motion‐triggered cameras. Daily carcass survival was greater in winter than during spring, summer, and fall. Survival was related negatively to canopy cover (trees and shrubs within a 50‐m buffer) and window area, and positively to pavement cover. Using an exponential model of survival time, estimated mean time of survival of carcasses (t± SE) was 82.9 ± 11.7 d for winter and 11.8 ± 7.2 d for other seasons. Raccoons (Procyon lotor) scavenged more carcasses than other species. Our results suggest that (1) carcass survival times may be short at locations with preferred habitats of known scavengers and predictable sources of food, and (2) knowledge of scavenger distribution and activity can inform predictive models of persistence. In studies of bird‐window collisions, the influence of scavenger bias can be minimized by maintaining short time intervals between carcass searches. Search intervals can be inferred by estimating the number of days that a carcass should persist at a site, which can be calculated using predicted daily survival probabilities of carcasses at study buildings.     

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。     

Characterization of twenty-six polymorphic microsatellite markers for the fox squirrel (Sciurus niger) and their utility in gray squirrels (Sciurus carolinensis) and red squirrels (Tamiasciurus hudsonicus)

Twenty-six polymorphic microsatellite markers were developed for the fox squirrel (Sciurus niger). The number of alleles ranged from 2 to 11 and observed heterozygosities ranged from 0.037 to 0.815. Preliminary screening revealed that numerous loci also were polymorphic in other squirrel species: Sciurus carolinensis and Tamiasciurus hudsonicus. These loci should be useful tools for investigating the social structure, mating tactics and movement behavior of fox squirrels.     

Facilitation of tree seedling establishment in a sand dune succession

Abstract. A nucleated pattern of establishment by pine seedlings of Pinus strobus and P. resinosa around pre-established oak trees of Quercus rubra in a sand dune succession in Ontario, Canada was examined using field observations, seed planting and habitat manipulation. Densities of young pines beneath oak canopies were approximately six times greater than in treeless areas, and densities on the north sides of the trees were significantly greater than on the south sides. However, oaks younger than 35 yr showed no preferential establishment beneath them, while the pine population structures beneath older oaks indicated single periods of successful recruitment. Pine seed planted beneath and beyond oak canopies of three sizes germinated primarily beneath the canopies of medium and large-sized oaks, but subsequent survivorship of seedlings over two growing seasons was poor. Several micro-environmental conditions were changed by oak canopies, but only shade showed a pattern closely corresponding to that of seedling establishment. A habitat manipulation experiment confirmed the primary role of shade in facilitating pine seedling establishment beneath oaks. Failure of pine to continue recruiting successfully beneath facilitating trees is tentatively attributed to intraspecific competition among pine individuals.     

Daily Activity and Nest Occupation Patterns of Fox Squirrels (Sciurus niger) throughout the Year

The authors investigated the general activity and nest occupation patterns of fox squirrels in a natural setting using temperature-sensitive data loggers that measure activity as changes in the microenvironment of the animal. Data were obtained from 25 distinct preparations, upon 14 unique squirrels, totaling 1385 recording days. The animals were clearly diurnal, with a predominantly unimodal activity pattern, although individual squirrels occasionally exhibited bimodal patterns, particularly in the spring and summer. Even during the short days of winter (9 hours of light), the squirrels typically left the nest after dawn and returned before dusk, spending only about 7 hours out of the nest each day. Although the duration of the daily active phase did not change with the seasons, the squirrels exited the nest earlier in the day when the days became longer in the summer and exited the nest later in the day when the days became shorter in the winter, thus tracking dawn along the seasons. During the few hours spent outside the nest each day, fox squirrels seemed to spend most of the time sitting or lying. These findings suggest that fox squirrels may have adopted a slow life history strategy that involves long periods of rest on trees and short periods of ground activity each day.     

The effect of landscape structure on dispersal distances of the Eurasian red squirrel

Landscape structure can affect dispersal and gene flow in a species. In urban areas, buildings, roads, and small habitat patches make the landscape highly fragmented and can inhibit movement and affect dispersal behavior. Similarly, in rural forested areas, large open areas, such as fields, may act as barriers to movement. We studied how landscape structure affects natal dispersal distances of Eurasian red squirrels (Sciurus vulgaris) in an urban area and a rural area in Finland, by monitoring juvenile red squirrels with radio telemetry. We observed extremely long dispersal distances—up to 16 km—in the rural study area, but shorter distances—on average only half a kilometer—in the urban study area. The landscape structure affected the eventual dispersal paths; in the rural landscape, dispersers favored spruce dominated areas and avoided fields along their dispersal route, although they occasionally even crossed wide fields. In the urban landscape, squirrels preferred areas with deciduous or coniferous trees. The movement steps made by dispersers were longer in the more hostile landscape compared to forested areas. Despite these effects on movement path, the landscape structure only had a minor effect on straight line dispersal distances moved from the natal nest. In other words, individuals moved longer distances and were likely to circumvent barriers in their path, but this did not affect how far they settled from their natal home. This result indicates that, although landscape structure has obvious effects on movement, it still may have only a small effect on other aspects of the population, for example, gene flow.