Alarm Calls of California Ground Squirrels (Spermophilus beecheyi)

California ground squirrel alarm vocalizations were recorded in field and laboratory, and sonagraphically analysed. The contexts of both naturally occurring and experimentally elicited calls were noted in the field. The components of this graded system are chatters, chats and whistles. Chatters and chats are often elicited by terrestrial predators, whistles commonly by low flying raptors. Whistles are more commonly associated with cryptic behavior and flight than chatter-chats, but both call types usually elicit bipedal alert postures. These calls grade along a number of dimensions which may signal redundantly the level of excitation of the caller. We propose that the chatter-chat calls of highly aroused squirrels are composed of more and longer notes, occur at a higher rate, are less noisy and contain more frequency modulation. Whistles, however, are single-note calls that contain no frequency modulation, even though they are emitted by highly aroused squirrels and are long and noise free. Preliminary data suggest that: 1) chats are easier for a human ♀ to localize than whistles; 2) elevation of the head, by adopting bipedal postures and mounting promontories, enhances the audibility of alarms.     

Microsatellite loci for the Siberian flying squirrel,Pteromys volans

We report the isolation and characterization of polymorphic microsatellite loci for the Siberian flying squirrel Pteromys volans. The seven most useful loci had between six and 11 alleles and expected heterozygosities ranging from 0.477 to 0.866. We also tested the utility of these loci in other squirrel species, northern flying squirrels (Glaucomys sabrinus and G. volans) and the common red squirrel (Sciurus vulgaris). Three of the Siberian flying squirrel loci were polymorphic in other squirrel species, suggesting a limited potential for cross‐species use.     

Habitat and landscape correlates of southern flying squirrel use of red-cockaded woodpecker clusters

Southern flying squirrels (Glaucomys volans) can have significant negative impacts on red-cockaded woodpecker (Picoides borealis) reproductive success and group size. Although direct control of southern flying squirrels may be necessary in small red-cockaded woodpecker populations (<30 groups), creation of high quality habitat through landscape management is the preferred method for managing larger woodpecker populations. Thus, we determined the habitat and landscape factors within 100 m, 400 m, and 800 m of cluster centers that were related to southern flying squirrel use of red-cockaded woodpecker cavities at the Carolina Sandhills National Wildlife Refuge, South Carolina. At all spatial scales, the number of cavities in the cluster was the most influential variable determining use by southern flying squirrels. At the 400-m and 800-m scales, the amount of stream length was also positively associated with the presence of flying squirrels. The proximity and amount of hardwoods surrounding clusters were not related to southern flying squirrel use at any spatial scale; thus, removal or conversion of hardwood stands surrounding red-cockaded woodpeckers may not be necessary for reducing cavity kleptoparasitism by flying squirrels. However, when establishing recruitment clusters, areas with streams should be avoided and addition of artificial cavities to existing clusters should be done judiciously to minimize the number of excess cavities. Published 2012. This article is a U.S. Government work and is in the public domain in the USA.     

Winter site occupancy patterns of the northern flying squirrel in boreal mixed-wood forests

We assessed habitat use by the northern flying squirrel (Glaucomys sabrinus) to test the hypothesis that the species is not a late-seral coniferous forest specialist in boreal mixedwood forests of northwestern Québec. We monitored 149 pairs of nest boxes over 3 visits during February–April 2008. A total of 31 pairs of nest boxes were occupied during the study. The average (3.7, 2.8, and 1.8) and maximum (8, 6, and 4) numbers of individuals in nest boxes decreased from the first to last visit. Average air temperatures were below freezing and increased with the arrival of spring (?23.9, ?14.5, and ?3.7 °C). Using GIS software, we created a 430-m (median dispersal distance) buffer around each pair of nest boxes and extracted landscape composition variables from digital forest inventory maps. We created models that potentially explained site occupancy by the northern flying squirrel as well as detection probability. Model selection indicated that the area of coniferous forests negatively affected site occupancy by flying squirrels while temperature positively influenced detection probability. Our results support the hypothesis that the northern flying squirrel is not a habitat specialist, but can occupy a wider range of forest cover types than previously recognized in various landscape contexts.     

Role of the Siberian flying squirrel as an umbrella species for biodiversity in northern boreal forests

One of the potentially useful indirect shortcut methods in biodiversity conservation is the umbrella species concept. An umbrella species can be seen relatively demanding for the size of the area and probably also for certain habitat types: conservation management for the umbrella species would thus encompass other species preferring similar habitats but with smaller area requirements. As such, it has a comprehensive spatial aspect for landscape planning. We tested the role of the Siberian flying squirrel (Pteromys volans) as an umbrella species for wood dependent species among red-listed and old-growth forest associated polypores, epiphytic lichens and beetles. Flying squirrels inhabit home ranges of several to tens of hectares, and prefer mature spruce-dominated (Picea abies) mixed forests, which often have high amounts of dead wood. We carried out species surveys and trappings during 1 year from 20 mature spruce-dominated forest stands (altogether 162 ha), of which 12 were occupied by the flying squirrel. The amount of dead wood was higher in occupied stands than in unoccupied stands. We also found a tendency for a higher number of species and number of records in occupied stands, a relationship mostly due to the polypore species. The presence of the flying squirrel may reflect the habitat availability for species depending on dead and living wood, and assist in site selection of conservation areas. We suggest that the flying squirrel has potential as an umbrella species to partly enhance maintenance of biodiversity in northern boreal forests in Finland.     

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.     

A meta-analysis of forest age and structure effects on northern flying squirrel densities

Research on the impact of clearcut logging and partial harvesting practices on northern flying squirrels (Glaucomys sabrinus) has shown inconsistent and contrary results, limiting the use of this species as a management indicator species. Much of this variability in study results is due to the labor intensive nature of studying flying squirrels, resulting in small sampling sizes (average = 5.2 sites, n = 14) and high variation (CV = 0.59) across studies. We conducted a meta-analysis of relevant studies from North America to determine how forestry practices affect flying squirrel abundance. Mean effect size was −1.18 (P < 0.001; n = 14) for all studies, indicating a strong difference between control stands and those regenerating postclearcut or following partial harvesting. Our results support the association of northern flying squirrels with mature, uncut forest and their suitability as ecological indicators of these vegetation types. © 2011 The Wildlife Society.     

Survey for zoonotic rickettsial pathogens in northern flying squirrels, Glaucomys sabrinus, in California

Epidemic typhus, caused by Rickettsia prowazekii, is maintained in a southern flying squirrel (Glaucomys volans) sylvatic cycle in the southeastern United States. The northern flying squirrel (Glaucomys sabrinus) has not been previously associated with R. prowazekii transmission. A second rickettsial pathogen, Anaplasma phagocytophilum, infects dusky-footed woodrats (Neotoma fuscipes) and tree squirrels in northern California. Because northern flying squirrels or their ectoparasites have not been tested for these rickettsial pathogens, serology and polymerase chain reaction (PCR) were used to test 24 northern flying squirrels for R. prowazekii and A. phagocytophilum infection or antibodies. Although there was no evidence of exposure to R. prowazekii, we provide molecular evidence of A. phagocytophilum infection in one flying squirrel; two flying squirrels also were seropositive for this pathogen. Fleas and ticks removed from the squirrels included Ceratophyllus ciliatus mononis, Opisodasys vesperalis, Ixodes hearlei, Ixodes pacificus, and Dermacentor paramapertus.     

Strongyloides robustus and the northern sympatric populations of northern (Glaucomys sabrinus) and southern (G. volans) flying squirrels

Within North America, northern (Glaucomys sabrinus) and southern (Glaucomys volans) flying squirrels occupy distinct ranges with limited overlap. Sympatry in northern latitudes coincides with northern hardwood vegetation from Minnesota to New England. Strongyloides robustus is an intestinal parasite that infects both species but appears to be deleterious only to northern flying squirrels. As a result, S. robustus could be a critical determinant of flying squirrel population characteristics in at least some areas of sympatry. However, cold weather could potentially limit the distribution of S. robustus in northern climates. Therefore, we assessed fecal samples from both flying squirrel species to determine the presence of the nematode in Wisconsin. Strongyloides robustus was found in 12 flying squirrel scat samples and infected 52% of southern flying squirrels and 11% of northern flying squirrels. Prevalence of S. robustus infection for northern flying squirrels was substantially lower than previously reported from more southern regions. This is the northernmost documentation of S. robustus in flying squirrels and the first documentation of S. robustus parasitizing flying squirrels in Wisconsin.     

Flying squirrels and their ectoparasites: disseminators of epidemic typhus

Information gathered during the past decade indicates that the eastern flying squirrel, Glaucomys volans, is a zoonotic reservoir of Rickettsia prowazekii - causative agent of louse-borne (epidemic) typhus. The sporadic cases o f typhus that have occurred in the USA in association with flying squirrels provide evidence that flying squirrels can transmit R. prowazekii infection to humons. Strains of R. prowazekii, isolated from flying squirrels multiply readily in human body lice, but flying squirrel lice, although readily infected, are very host specific and tend not to bite humans. It may be that the infection is spread to humans in infective ectoporasite faeces aerosolized when the flying squirrels groom themselves. As Joseph McDade emphasizes in this article, current concepts of typhus epidemiology and control must be re-evaluated to take into account this zoonotic aspect.     

Seasonal energetics and torpor use in North American flying squirrels

Seasonal cold temperatures require mammals to use morphological, behavioural, or physiological traits to survive periods of extreme cold and food shortage. Torpor is a physiological state that minimizes energy requirements by decreasing resting metabolic rate (MR) and body temperature (T_b). Many rodent species are capable of torpor, however, evidence in northern and southern flying squirrels (Glaucomys sabrinus and Glaucomys volans, respectively) has remained anecdotal. We experimentally attempted to induce torpor in wild-caught flying squirrels by lowering ambient temperature (T_a) and measuring MR using open-flow respirometry. We also studied seasonal differences in MR and T_b at various T_a. Both MR and T_b provided evidence for torpor in flying squirrels, but only infrequent, shallow torpor. MR decreased infrequently and any decreases were rarely sustained for longer than one hour. We found a significant positive relationship between T_a and T_b only in G. volans, which suggests that G. volans is more susceptible to low T_a compared with G. sabrinus, possibly due to their small body size. We observed no substantive seasonal or interspecific differences in the relation between MR and T_a, with the exception that northern flying squirrels expended more energy at cold T_a during warm season trials than other species-season combinations. The infrequency of torpor use in our experiments suggests that other energy-saving strategies, such as social thermoregulation, may limit the reliance on torpor in this lineage.     

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.     

Patch occupancy by squirrels in fragmented deciduous forest: effects of behavior

We tested whether species-specific behavioral traits could explain patterns of habitat patch occupancy by five different squirrel species in Ontario, Canada: the northern and southern flying squirrel (Glaucomys sabrinus and Glaucomys volans), the North American red squirrel (Tamiasciurus hudsonicus), the eastern chipmunk (Tamias striatus), and the eastern gray squirrel (Sciurus carolinensis). Flying squirrel species exhibit group nesting in winter, which may put them at risk of extirpation in small patches with few individuals to contribute to group nests. Flying squirrels are also volant, potentially making non-treed matrix a barrier. Our surveys revealed that G. sabrinus was most likely to occur in large patches that were embedded in landscapes with low connectivity, and least likely to occur in small patches in highly connected landscapes. Conversely, G. volans was most likely to occur in large, well-connected patches and least likely to occur in small, unconnected patches. Patch occupancy by the cursorial squirrels was not strongly influenced by patch area or isolation. These findings reinforce previous studies suggesting that an understanding of species-specific traits such as behavior is an important consideration when interpreting habitat fragmentation effects.     

Contrasting seasonal dynamics in fleas of the Siberian flying squirrel (Pteromys volans) in Finland

Abstract.  1. The seasonal and spatial variation of the adult flea fauna (Siphonaptera) was examined in connection with live-trapping studies of the Siberian flying squirrel ( Pteromys volans ) in three study areas in southern Finland between 1997 and 2005.
2. The numerically dominant flea species of the Siberian flying squirrel were Tarsopsylla octodecimdentata octodecimdentata and Ceratophyllus (Monopsyllus) indages indages (Ceratophyllidae); the latter being a host specialist of the Siberian flying squirrel. Tarsopsylla octodecimdentata , which also commonly occurs on the red squirrel ( Sciurus vulgaris ), infected a large proportion of the host population throughout the winter, whereas C. indages occurred predominantly during the summer and autumn, being practically the only flea species in nests during the flying squirrels' breeding season and on juveniles still inhabiting their natal nest.
3. The use of nest boxes by flying squirrels did not have any positive effects on flea populations, but it may have had a negative impact on T.   octodecimdentata .
4. The potentially vulnerable C.   i.   indages is a predictable, widespread component in the flea fauna of the Siberian flying squirrel at various spatial scales, and it is likely to occur in most of the permanent flying squirrel populations in Finland.     

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.     

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.     

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.     

Species-specific and shared features in vocal repertoires of three Eurasian ground squirrels (genus <Emphasis Type="Italic">Spermophilus</Emphasis>)

Along to alarm calls, Eurasian ground squirrels of the genus Spermophilus also produce other call types toward potential predators and rival conspecifics. Individually identified 50 speckled (Spermophilus suslicus), 18 European (S. citellus) and 59 yellow (S. fulvus) ground squirrels were examined for interspecies differences in their vocal repertoires. A separate sample of 116 (90 adult and 26 juvenile) S. suslicus was examined for presence of ultrasound in their alarm calls. In addition, all tonal calls in all the three species were checked for presence of nonlinear phenomena. Calls were elicited by approaching animals in live-traps or near burrows; some types of vocalizations were also recorded during handling. Eight call types, three tonal and five wideband ones, were described. Vocal repertoires were remarkably similar between species, excluding the alarm calls, which were species-specific. Alarm calls with ultrasonic components were found in two individuals of S. suslicus. Concerning nonlinear phenomena, biphonation in alarm calls of S. suslicus, frequency jumps and sidebands in screams of S. citellus, frequency jumps and subharmonics in screams of S. fulvus were found. Results are discussed with literature evidence on audible and ultrasonic vocalizations in ground squirrels.     

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ABSTRACT

In most mammals, adults produce relatively low frequency vocalizations compared to those of juveniles. This rule is not maintained however at least in four species of ground squirrels, whose juveniles call at the adult's fundamental frequency. These findings have been obtained however with separate sets of juveniles and adults and no data is available concerning the ontogeny linked to these differences. Here we analyze the acoustic structure of alarm calls of the same Yellow Spermophilus fulvus and Speckled S. suslicus ground squirrel individuals, recorded as pups and as adults after hibernation. We found the fundamental frequencies of adults within the same frequency ranges as those of pups, in spite of the significant difference in body mass. In ground squirrels, severing the relationship between body size and call frequency removes some vocal cues to age. We discuss some functional hypotheses advanced to explain manipulations with fundamental frequencies in ground squirrels and other animals, and suggest the lack of data for discussing the mechanisms of such vocal tuning.