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.     

A nuclear perspective on endemism in northern flying squirrels (Glaucomys sabrinus) of the Alexander Archipelago, Alaska

We used microsatellites to examine populationstructure and genetic diversity in northernflying squirrels in the Alexander Archipelagoof Southeast Alaska, with an emphasis on theendemic Prince of Wales flying squirrel(Glaucomys sabrinus griseifrons). Previouswork showed this subspecific designationcoincided with a distinct mitochondrial lineageon eleven islands (the Prince of Wales [POW]complex). To obtain a nuclear perspective onthis lineage and to further investigate geneticdiversity among insular populations, weexamined six microsatellite loci in 233 flyingsquirrels representing eight populations inSoutheast Alaska and a population from interiorAlaska (seven island and two mainlandlocalities). Island populations have lowerheterozygosity and allelic diversity thanmainland populations. Overall, populationpairs show a pattern of isolation by distance,indicating there is little long-distance geneflow across the archipelago. Analyses ofmicrosatellite allele frequencies revealsignificant differences between the POW complexpopulations and others we examined, a findingcongruent with the mitochondrial data. Thepopulation from Mitkof Island, a non-POWcomplex island, also differs significantly fromother populations in allele frequencies. Thesix POW complex populations are geneticallyvery similar, suggesting current or recent geneflow among these islands, while there seems tobe no gene flow between the POW complex andother populations in Southeast Alaska. Ourdata corroborate mitochondrial DNA resultsindicating that G. s. griseifrons isgenetically distinct and suggest a generalpattern of isolation of insular flyingsquirrels in Southeast Alaska.     

The cost of living large: comparative gliding performance in flying lizards (Agamidae: Draco)

Despite exhibiting considerable interspecific variation in body mass, flying lizards of the genus Draco are isometric in their area-mass scaling relationships and exhibit no significant compensatory variation in wing aspect ratio. Thus, larger species are expected to be relatively poor gliders, in lieu of behavioral or physiological compensation, when compared with smaller congeners. Here we tested this hypothesis by conducting gliding performance trials for 11 Draco species spanning virtually the entire size range of the genus. We considered three primary performance variables: maximum velocity adjusted for wind conditions, height lost over a standard horizontal glide distance, and glide angle. Comparative analysis confirmed that larger species are relatively poor gliders and do not compensate substantially for their higher wing loadings via either behavioral or physiological mechanisms. Flying lizards were found to exhibit substantial context-dependent variation in glide performance, with smaller species often exhibiting extensive variation in height lost and glide angle between trials. Variation also was observed in empirically derived velocity profiles, with only a subset of individuals appearing to perform equilibrium glides. Such size-dependent variation in performance has important consequences for the ecology and evolution of flying lizards and other glissant taxa.     

Mechanical performance of aquatic rowing and flying

Aquatic flight, performed by rowing or flapping fins, wings or limbs, is a primary locomotor mechanism for many animals. We used a computer simulation to compare the mechanical performance of rowing and flapping appendages across a range of speeds. Flapping appendages proved to be more mechanically efficient than rowing appendages at all swimming speeds, suggesting that animals that frequently engage in locomotor behaviours that require energy conservation should employ a flapping stroke. The lower efficiency of rowing appendages across all speeds begs the question of why rowing occurs at all. One answer lies in the ability of rowing fins to generate more thrust than flapping fins during the power stroke. Large forces are necessary for manoeuvring behaviours such as accelerations, turning and braking, which suggests that rowing should be found in slow-swimming animals that frequently manoeuvre. The predictions of the model are supported by observed patterns of behavioural variation among rowing and flapping vertebrates.     

Site occupancy and cavity use by the northern flying squirrel in the boreal forest

Tree cavities are used as shelter and breeding nests by numerous avian and mammalian species. In cold environments, tree cavities are often proposed as the best winter nest choice because of the superior protection they offer from precipitation, wind, cold temperatures, and predators. As such, they represent a critical resource, which has the potential to limit population size of non-excavating species. We assessed factors affecting site occupancy in the boreal forest by northern flying squirrels, a secondary user of tree cavities, and to identify which nest type is preferred during the colder days of the autumn–winter period. We trapped flying squirrels twice in 59 aspen-dominated stands in the autumn period using low- (1.5 m above ground-level) and high-mounted (4 m) traps to determine site occupancy. A total of 85 individuals were captured on 2,880 trap-nights. During the winter period, we radio-tracked 26 individuals to 87 diurnal nests in 220 locations. None of the habitat variables considered (cavity availability, woody debris, and lateral cover) explained site occupancy. Detectability decreased with precipitation, and was lower using high traps than low traps. Both females and males used tree cavities (26%), external nests (39%), and ground nests (35%). In cold weather, females preferred ground nests, whereas males preferred external nests. Our results do not support the hypothesis that tree cavities represent a limiting factor to northern flying squirrels in cold environments. Instead, this species seems to be a generalist and is opportunistic, using a variety of nest types. Nevertheless, practices ensuring the persistence of large diameter live cavity trees, providing better insulative properties, are likely to increase the relative use of tree cavities as nest sites by northern flying squirrels. © 2011 The Wildlife Society.     

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.     

Locomotor performance and social dominance in male Anolis cristatellus

The proximal mechanisms determining social dominance are not well understood. We used the highly territorial lizard A. cristatellus to test two main hypotheses: (1) that male social dominance is associated with locomotor abilities; (2) that locomotor abilities (maximal performance), as measured in the laboratory, are correlated with behaviour in the field. In the field, we recorded locomotor behaviours and assertion displays, then characterized microhabitat use and thermal relations. In the laboratory, we measured maximum sprint running speed, endurance and morphometric characters, and assessed dominance by pairing males of similar body size in an experimental arena. In 72 of 77 interactions, one lizard (the ‘winner’) was unequivocally determined to be dominant over the other (the ‘loser’). Winners performed more assertion displays than losers before capture and also had higher endurance in laboratory tests. Although contestants were matched for snout-vent length, winners had significantly deeper and wider heads. However, we found no significant differences in field locomotor behaviours, perch or thermal characteristics, head length, or maximal sprint speed. Our findings support those of previous studies, and extend them in several ways. This is the first demonstration that assertion displays in the field are related to both locomotor performance and laboratory-assessed social dominance. Locomotor performance may directly affect social dominance by allowing some males to perform better in dyadic interactions. Alternatively, both locomotor performance and social dominance may be linked to a common underlying mechanism, such as variation in hormone levels, which are known to affect aggression, locomotor performance and morphology.     

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.     

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.     

Locomotor performance and muscle metabolic capacities: impact of temperature and energetic status

In aquatic ectotherms, muscle metabolic capacities are strongly influenced by exogenous factors, principally temperature and food availability. Seasonal changes in temperature lead many organisms to modify their metabolic machinery so as to maintain capacity even in "slower" cold habitats. Modifications of mitochondrial capacities are central in this response. The increases in protein-specific oxidative capacities of mitochondria during cold acclimation of temperate fishes do not occur during the evolutionary adaptation to cold in Antarctic species. Instead, Antarctic fishes tend to increase the proportion of fibre volume devoted to mitochondria, perhaps to facilitate intracellular distribution of oxygen and metabolites. Variation in energetic status can drastically modify muscle metabolic status, with glycolytic muscle changing more than oxidative muscle. This in turn impacts swimming performance. A decrease in the condition of cod leads endurance at speeds above Ucrit to drop by 70%. Sprint swimming is less affected, perhaps as it does not exhaust glycolytic muscle. We used interindividual variation in muscle metabolic capacities to identify correlates of swimming performance in stickleback and cod. Activities of cytochrome c oxidase in glycolytic muscle are a correlate of sprint swimming in stickleback (Gasterosteus aculeatus) and cod (Gadus morhua), whereas lactate dehydrogenase activities in glycolytic muscle are a correlate of cod endurance swimming. In scallops, gonadal maturation leads to virtually complete mobilisation of glycogen from muscle. This does not reduce the capacity of the scallops, Chlamys islandica and Euvola ziczac, to mount escape responses, but significantly slows their recuperation from exhaustive exercise. Muscle metabolic capacities fall in parallel with glycogen mobilisation. In the compromise between muscles' dual roles as a motor and a macromolecular reserve, a significant loss in locomotory ability occurs during gametogenesis and spawning. Reproductive fitness takes the upper hand over maintenance of performance.