Lucas A. Wauters

Anticipatory reproduction and population growth in seed predators.

Mast seeding, the intermittent, synchronous production of large seed crops by a population of plants, is a well-known example of resource pulses that create lagged responses in successive trophic levels of ecological communities. These lags arise because seed predators are thought capable of increasing reproduction and population size only after the resource pulse is available for consumption. The resulting satiation of predators is a widely cited explanation for the evolution of masting. Our study shows that both American and Eurasian tree squirrels anticipate resource pulses and increase reproductive output before a masting event, thereby increasing population size in synchrony with the resource pulse and eliminating the population lag thought to be universal in resource pulse systems.

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

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

Molecular phylogeography of European Sciurus vulgaris: refuge within refugia?

The red squirrel (Sciurus vulgaris) is a well‐known forest animal distributed all over Europe. Still, we are far from having a firm knowledge of the species’ phylogeography. This study investigates the genetic differentiation of S. vulgaris across the species’ Eurasian range, using sequence data from the mitochondrial DNA gene (D‐loop, 252 base pairs, cytochrome b, 359 base pairs), and eight variable autosomal microsatellite loci genotyped for 236 individuals. The results reveal the presence of two main mitochondrial phylogroups. The first clade comprises the individuals from the region of Calabria in southern Italy, belonging to the subspecies S. v. meridionalis, while the second clade contains the remainder of the studied individuals. Bayesian analysis of microsatellite genotypes resulted in three main clusterings corresponding to the three S. vulgaris subspecies: infuscatus, meridionalis and fuscoater. Geographical distribution of mtDNA haplotypes and mismatch analysis suggest a common refugium for the red squirrel across most of its present range from which expansion happened rather rapidly. The genotype mixing of italicus with northern populations could be a residual of postglacial expansion. The lack of mixing between the Calabrian lineage and the rest of European red squirrel haplotypes can be seen as evidence for distinct histories throughout the Pleistocene. Calabrian mtDNA probably diverged in an ice age contraction and remained isolated from the neighbouring squirrel populations until very recent times.

Comparing data of different survey methods for sustainable wildlife management in hunting areas: the case of Tarangire–Manyara ecosystem, northern Tanzania

Cost–benefit considerations of wildlife monitoring are essential, particularly, in areas outside national park boundaries, where resources for conducting wildlife censuses are scarce, but that, at the same time, are subject to high pressure for wildlife utilization, such as hunting. Large mammal survey data from various sources were collated and analyzed to investigate which methods are best suited for monitoring purposes at low cost in the Tarangire–Manyara ecosystem, northern Tanzania. Our results indicate that primary data (from aerial and road transects counts) that involve direct species observations, although sometimes very expensive, are required for establishing the status of the target species in terms of density or population size. Concomitantly, secondary data from various sources, such as interviews, hunting quota, and damage reports, obtained over wide areas and over longer periods of time, can provide important information on presence/absence and distribution of species within an area. In addition, the study revealed that hunting quotas set did not correlate with species abundance/numbers from the primary data surveys for most of the large mammals hunted within the ecosystem. For a better conservation and management of wildlife, in particular with respect to the forthcoming formation of Wildlife Management Areas, we propose an integrated approach to wildlife monitoring using primary and secondary data sources through the involvement of local people’s knowledge.

Exclusive core areas and intrasexual territoriality in Eurasian red squirrels (Sciurus vulgaris) revealed by incremental cluster polygon analysis

When animal home ranges overlap extensively in species lacking overt territorial behaviours, identifying exclusive core areas within individual ranges can be difficult. By analysing the size and overlap of successively smaller core areas among individual Eurasian red squirrels (Sciurus vulgaris), we determined exclusive areas within the home ranges of resident males and females. Possible effects of habitat composition and food supplies were explored by monitoring squirrels in different conifer forests and during years with low and high tree seed production. Using outlier-exclusive cores (OEC) revealed that the total ranges consisted of large sally zones (on average, 35% of the total minimum convex polygon [MCP] range) around home ranges with multi-nucleate cores. The mean OEC home range size did not differ between the sexes but was larger with poor food availability. Home ranges (99% incremental cluster polygons [ICP]) overlapped extensively between sexes (average overlap high food–low food: males by females 21–40%, females by males 43–45%) and among males (males by males 26–44%), while intrasexual overlap among females was low (9–10%). The overlap of inner cores among females rapidly approached zero, suggesting the intrasexual territoriality of 75% core areas. This was not the case among male squirrels, for which intrasexual overlap averaged only 4% at 50% but 18% at 75% core areas. Even the smallest inner cores had some degree of intersexual overlap, indicating that complete territoriality did not occur in this species. Female home ranges were more strongly affected by annual fluctuations in food supplies than male ranges. Females reduced the size of their food-based intrasexual territories when food availability increases. Males probably benefit from using larger home ranges and core areas, which overlap with the ranges of several females, by increasing their probability of successful mating.

Ljungan Virus and an Adenovirus in Italian Squirrel Populations

Abstract We report Ljungan virus infection in Eurasian red squirrels (Sciurus vulgaris) for the first time, and extend the known distribution of adenoviruses in both native red squirrels and alien gray squirrels (Sciurus carolinensis) to southern Europe.

Estimating and comparing food availability for tree-seed predators in typical pulsed-resource systems: alpine conifer forests.

Abstract Alpine conifers produce pulsed resources for the community of tree‐seed consumers. Here, we describe field and statistical methods to estimate spatio‐temporal variation in seed‐crops in forests with different species composition. Annual and between‐site variation in seed production was high and characterised by occurrence of mast‐crops followed by seed‐crop failure the following year. Seed‐crops of different species did not fluctuate in parallel. Abbreviations: DLSM, differences of least square means; CED, Cedrasco; OGA, Oga; VAL, Valfurva; BOR, Bormio; RHE, Rhemes; COG, Cogne

The strong and the hungry: bias in capture methods for mountain hares Lepus timidus

Abstract Estimating density, age and sex structure of wild populations is a key objective in wildlife management. Live trapping is frequently used to collect data on populations of small and medium-sized mammals. Ideally, sampling mammal populations by live capturing of individuals provides a random and representative sample of the target population. Trapping data may, however, be biased. We used live-capture data from mountain hares Lepus timidus in Scotland to assess sampling bias between two different capture methods. We captured hares using baited cage traps and long nets on five study areas in the Scottish Highlands. After controlling for the effects of body size, individuals caught in traps were lighter than individuals caught using long nets, suggesting that the body condition of hares differed between the capture methods. This tendency may reflect an increased risk-taking of individuals in poorer body condition and less aversion to entering traps in order to benefit from eating bait. Overall, we caught more adult hares than juveniles and more female hares than males. Our results show that estimates of density and population structure of mountain hares using live-capture data could be affected by the capture method used. We suggest that live-capture studies employ more than one capture method and test for heterogeneity in capture probability to minimise potential bias and achieve reliable estimates of population parameters.

Living on the Edge: Can Eurasian Red Squirrels (Sciurus vulgaris) Persist in Extreme High-elevation Habitats?

Abstract Studying intraspecific spatiotemporal variation in vital rates among populations over a range of environmental conditions is essential to reveal intrinsic and extrinsic factors affecting population dynamics. Mammal populations living at higher elevations often have higher adult survival, shorter breeding seasons, and lower reproductive output per season than at lower elevations. We studied dynamics of a Eurasian red squirrel (Sciurus vulgaris) population in high-elevation, mountain pine (Pinus mugo) forest with extreme winters, in the Central Italian Alps, and compared vital rates with populations in more productive habitats at lower elevations. Average density was 0.14 ± 0.07 squirrels ha−1 (range 0.07–0.30 ha−1), and numbers typically increased in summer–autumn as a result of seasonal reproduction and immigration. Mean persistence time was only 12.5 months, and there was a nearly complete population turnover in only two years. Local survival and recruitment rate were correlated with seasonal population growth rate, and partial effect of survival explained 80% of variation in growth rate. While reproductive rate in mountain pine habitat was more similar than in more productive habitats at lower elevations, density and autumn–winter survival were much lower. Thus, red squirrels did not show the adaptations observed in several other mammal species, but might invest heavily in early reproduction to compensate for short life expectancy.

Measurement of fecal glucocorticoid metabolite levels in Eurasian red squirrels ( Sciurus vulgaris ): effects of captivity, sex, reproductive condition, and season

The causes and consequences of physiological stress in wildlife are of great interest in a wide range of biological disciplines including understanding how environmental changes affect species fate and persistence. In some areas, the Eurasian red squirrel (Sciurus vulgaris) faces local extinction because of the invasive Eastern gray squirrel (Sciurus carolinensis). Whether or not physiological stress is induced by the presence of invasive species or contributes to local declines in Eurasian red squirrels is unknown. Here, we develop an assay that can be used to quantify physiological stress in fecal samples from Eurasian red squirrels to eventually address these questions. We captured free-living squirrels (6 females, 11 males) and placed them into captivity for 48 h. Fecal glucocorticoid metabolite (FGM) concentrations in female and male squirrels were significantly higher 24 and 32 h after initial captivity with a lag time to peak excretion ranging from 24 to 36 h. We measured FGM concentrations in free-living squirrels (37 females, 45 males) over a 3-year period. Lactating free-living squirrels had higher FGM concentrations compared to nonbreeding or pregnant squirrels but there were no differences in FGM concentrations in males with scrotal and abdominal testes. Free-living squirrels had the highest FGM concentrations in the winter and lowest in the summer (winter > autumn > summer). Squirrels kept in captivity for 4–48 h had significantly higher FGM concentrations than free-living squirrels (111 fecal samples from 82 squirrels). FGM concentrations in captive but not wild squirrels were significantly repeatable. We found no sex differences and no association with body mass in FGM concentrations in captive or free-living squirrels. Our results indicate that this assay can accurately quantify physiological stress in Eurasian red squirrels, which may be useful for future studies to document how the invasive Eastern gray squirrel contributes to local extinction.