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How to send red squirrel bodies for post mortem

Advice on how and where to send dead red squirrels for post mortem can be found here.

2020

• Anderson, N. E. & Lurz, P. W. W. (2020) The density of Smith’s bush squirrel, Paraxerus cepapi, in Luambe National Park, Zambia. African Journal of Ecology 58(2): 332-336.

 

• Bamber, J., Shuttleworth, C. & Hayward, M. (2020) Do Differing Levels of Boldness Influence the Success of Translocation? A Pilot Study on Red Squirrels (Sciurus vulgaris). Animals 10: 1748.

Conservation translocations, including reintroductions, are practices that are vital to restoring biodiversity and ecosystem function within conservation schemes globally. Sadly, population translocations have had a poor success rate historically. At a time where biodiversity is constantly decreasing, improving translocation success is vital for future conservation schemes. Often, to improve success, the selection of individuals is based on genetic characteristics and behaviours linked directly to survival. Further development to improve selection is proposed within this paper using animal personality. The study took place opportunistically to test how personality, in particular an animal’s boldness/timidness, may influence a population restoration of red squirrels into the Ogwen Valley, North Wales. Despite frequent translocations, data on how boldness and timidness may affect the establishment of this species are low. Testing was performed on key survival behaviours and boldness/timidness pre-release. This was performed via video data collection and identification of key behaviours that could be used to identify boldness or behaviours that could be linked to reduced fitness once released. Encounters at different distance intervals were monitored post-release via camera trapping to identify if boldness/timidness may change the furthest encounter distance of focal animals away from their release site. Relationships between the period for an individual to reappear post-threat was significantly linked to boldness, with other behavioural results and the encounter distance also showing trends of a potential relationship. Our results indicate that bolder individuals have a higher chance of expressing behavioural traits that will increase exposure to risks and, therefore, reduce the likelihood of successfully establishing populations. However, the small sample size of this study means that further research is needed. We suggest that during early stages of conservation translocation programmes, personality testing for boldness should become common practice, and we recommend selecting timid individuals for an initial release to improve population establishment, with bolder individuals utilised later to expand population distribution.

• Bamber, J. A., Shuttleworth, C. M., Hayward, M. W. & Everest, D. J. (2020) Reinstating trophic cascades as an applied conservation tool to protect forest ecosystems from invasive grey squirrels (Sciurus carolinensis). Food Webs 25: e00164.

Abstract: Red squirrel (Sciurus vulgaris) range within the United Kingdom (UK) has retracted significantly due to the spread of an Invasive Alien Species, the North American Eastern grey squirrel (Sciurus carolinensis). Where grey squirrels are sympatric, red squirrel populations decline through inter-specific competition and squirrelpox virus (SQPV) infection. Grey squirrel eradication from the island of Anglesey facilitated the complete restoration of native red squirrels. Although native species recovery delivered significant ecological and economic benefits, the eradication extended only to a narrow sea-channel boundary, across which grey squirrel dispersal continues to occur. Hence, the long-term sustainability of Anglesey’s red squirrel population is vulnerable to grey squirrel re-establishment without continuous intervention. Recent research has demonstrated that as pine marten (Martes martes) landscape use intensity increases, so too does red squirrel occupancy, likely linked to parallel declines in grey squirrel occupancy. Restoration of this mustelid predator is a potential tool to deliver sustainable grey squirrel control by restoring a missing trophic component, depressing grey squirrel incursion rates onto Anglesey, reducing red squirrel exposure to SQPV. Recent UK pine marten translocations have sourced animals under licence from wild Scottish populations. We explore the alternative use of captive-bred founders, simultaneously introducing new genetic variability against a limited diversity within extant populations. A current conservation translocation is paired with an ongoing assessment of founder behaviour and ‘personality’ measured before release. We then highlight the multi-disciplinary approach to delivering applied red squirrel conservation programmes in the face of invasive species.

• Bamidele, A. & Akinpelu, A. (2020) Comparison of Cranial and External Morphology of Tree Squirrels (Funiscurus leucogenys) in Selected Locations of Rainforest in Nigeria. Journal of Scientific Research and Reports 25(5): 1-12.

Abstract: This study looked at the differences in cranial morphology of tree squirrel species (F. leucogenys) from four different locations in Rainforest part of Nigeria. The squirrels were captured through the use of locally fabricated live traps made of wire-mesh and steel. Trapped specimens were immediately transferred to the laboratory in captivity cages, where they were euthanized in a bell-jar containing chloroform-soaked cotton wool. Specimens were then preliminarily identified to the generic level, using an identification key. The skull of 131 tree squirrels were prepared using Long Island Natural History Museum guide and the sex of the specimens was determined by visual inspection of the external genitalia. The skull and other body parameters were measured using digital venier caliper. The results showed that the body parameters (HBL, TL, TBL, EL, HFL and BW) measured were slightly different from one location to another. Also, the cranial measure showed some similarities among some locations (Ile-Ife, Emure-Ekiti and Ado-Ekiti), while measurement on squirrels from Sekona was different from other three locations. In conclusion, the cranial measurements of the tree squirrels shows there was no new species of F. leucogeny from the sampled locations.

• Cerri, J., Bertolino, S., Genovesi, P., Carnevali, L. & Monaco, A. (2020) Blacklists do not necessarily make people curious about invasive alien species. A case study with Bayesian structural time series and Wikipedia searches about invasive mammals in Italy.

Blacklists of invasive alien species are a popular tool to manage and prevent biological invasions. Furthermore, by providing accessible examples of invasive alien species and by having a certain media resonance, they can in principle raise the awareness and make laypeople curious towards this topic. However, no study ever tested for this side-effect of blacklists. We tested if the implementation of the first blacklist of invasive alien species of the European concern, by the European Union in August 2016, increased visits to Wikipedia pages about invasive alien mammals in Italy. We adopted Bayesian Structural Time Series, using native mammals as a synthetic control, and we considered both invasive alien mammals that appeared on the list and those which were not included. Following the publication of the first European blacklist of invasive alien species, there was no increase in the amount of weekly visits to the Wikipedia pages about invasive mammals. This was true both for species that were included in the list and those which were not. Rather increased search volumes were syncronous to other events that had media resonance. Our results indicate that important policymaking initiatives, do not necessarily raise public awareness about biological invasions, even when these policies, such as blacklists, are easy to understand and have a certain media coverage. We emphasize the importance of coupling them with adequate communication campaigns and also to develop communication guidelines for the media.

• Cooper, A. & Jackson, D. (2020) Dune Gardening? A critical view of the contemporary coastal dune management paradigm. Area

Abstract: Modern coastal dune management is viewed largely through the prism of dune ecology. Achieving maximum biodiversity and preserving priority species are the primary objectives and management is based on interventions (grazing, mowing, burning, reseeding, and artificial destabilisation) to achieve that purpose. Under non-managed conditions, dune vegetation tends to evolve temporally following well-established succession patterns that lead to a low diversity scrub or woodland. Achieving high diversity involves resisting succession so as to preserve the more biodiverse, earlier stages. The net Accepted Article This article is protected by copyright. All rights reserved effect of management is to create dunes with a network of vegetation types that conform to human wish lists. Rather than natural environments, such interventions reduce dunes to the status of parklands. The natural status of NW European dunes under the conditions of relatively stable sea-level and generally limited sand supply over the past few thousand years, and certainly since the Little Ice Age (1650-1800), is to be at a relatively well-vegetated and advanced stage of succession. Modern dune management effectively amounts to a form of ‘dune gardening’ to maintain a status that is not natural (i.e. attuned to the ambient environment) under contemporary conditions nor those of the past two centuries. Current practice is based on a human value-judgement that views natural succession as the enemy. This view and the resulting actions to resist natural change are at variance with the more pervasive environmental management goal of maintaining natural systems by resisting human intervention. Such an approach reduces dune resilience to global climate change. We advocate an approach that views dunes as combined geomorphological and ecological systems capable of change in response to changing environmental drivers.

• Côrte‐Real, J., Lopes, A., Rebelo, H., Lopes, J., Amorim, F., Pita, R., Correia, J., Melo, P., Beja, P., Esteves, P. & Abrantes, J. (2020) Adenovirus emergence in a red squirrel (Sciurus vulgaris) in Iberian Peninsula. Transboundary and Emerging Diseases.

Abstract: Adenoviruses (AdV) are pathogens capable of infecting animals and humans leading to a wide spectrum of diseases. They have a widespread geographical dissemination and infect several species, including red squirrels (Sciurus vulgaris). In the Iberian Peninsula, there are no reports of infection of red squirrels by AdV. In this study, we report for the first time the detection of squirrel AdV in a red squirrel trapped in central Portugal. The phylogenetic analysis shows that the sequences obtained for the hexon and the penton base genes were similar to the Korean squirrel AdV strain and strongly divergent from the strains present in red squirrels in Europe. These findings indicate the emergence of a new strain circulating in Europe and raise concerns regarding the conservation of the red squirrel.

• Cox, P., Morris, P., Hennekam, J. & Kitchener, A. (2020) Morphological and functional variation between isolated populations of British red squirrels (Sciurus vulgaris). Journal of Zoology 312(4): 271-283.

Isolation due to habitat fragmentation can lead to morphological and functional variation between populations, with the effect being well documented in rodents. Here, we investigated whether such morphological variation could be identified between British populations of the Eurasian red squirrel (Sciurus vulgaris). This species was once widespread across Great Britain, but suffered a severe population decline across the 20th century, leaving a highly fragmented distribution. The aim was to test for morphological and biomechanical variation of the mandible between the remaining British red squirrel populations, and between British and continental European red squirrels. Linear and geometric morphometric methods were used to analyse shape in a sample of over 250 red squirrel hemi‐mandibles from across Britain plus a sample from Germany representing the central European subspecies. Procrustes ANOVA identified significant shape variation between populations, with particularly distinct differences being noted between red squirrels from Germany and several British red squirrel populations, which may reflect their evolutionary history. Linear biomechanical measurements showed that the red squirrels from Formby and Jersey had a significantly lower mechanical advantage of the temporalis muscle than other British populations, suggesting they were less efficient at gnawing. This functional difference may be related to many factors, such as founder effect, potential inbreeding and/or past supplemental feeding with less mechanically resistant food items. The current distribution of red squirrels (Sciurus vulgaris) in Great Britain is highly fragmented. This study demonstrates significant variation in the mandible between British red squirrel populations, both in morphology and in biomechanical proxies. The squirrels at Formby on the Lancashire coast are particularly notable for their reduced mechanical advantage of the temporalis. We speculate that such variation may be driven by differences in evolutionary history and/or diet.

• Dantzer, B., McAdam, A. G., Humphries, M. M., Lane, J. E. & Boutin, S. (2020) Decoupling the effects of food and density on life history plasticity of wild animals using field experiments: insights from the steward who sits in the shadow of its tail, the North American red squirrel. Journal of Animal Ecology n/a(n/a).

Summary: Long-term studies of wild animals provide the opportunity to investigate how phenotypic plasticity is used to cope with environmental fluctuations, and how the relationships between phenotypes and fitness can be dependent upon the ecological context. Most previous studies have only investigated life history plasticity in response to changes in temperature, yet wild animals often experience multiple environmental fluctuations simultaneously. This requires field experiments to decouple which ecological factor induces plasticity in fitness-relevant traits to better understand their population-level responses to those environmental fluctuations. For the past 32 years, we have conducted a long-term integrative study of individually marked North American red squirrels (Tamiasciurus hudsonicus Erxleben) in the Yukon, Canada. We have used multi-year field experiments to examine the physiological and life history responses of individual red squirrels to fluctuations in food abundance and conspecific density. Our long-term observational study and field experiments show that squirrels can anticipate increases in food availability and density, thereby decoupling the usual pattern where animals respond to, rather than anticipate, an ecological change. As in many other study systems, ecological factors that can induce plasticity (such as food and density) co-vary. However, our field experiments that manipulate food availability and social cues of density (frequency of territorial vocalizations) indicate that increases in social (acoustic) cues of density in the absence of additional food can induce similar life history plasticity, as does experimental food supplementation. Changes in the levels of metabolic hormones (glucocorticoids) in response to variation in food and density are one mechanism that seems to induce this adaptive life history plasticity. Although we have not yet investigated the energetic response of squirrels to elevated density or its association with life history plasticity, energetics research in red squirrels has overturned several standard pillars of knowledge in physiological ecology. We show how a tractable model species combined with integrative studies can reveal how animals cope with resource fluctuations through life history plasticity.

• Dastjerdi, A., Benfield, C., Everest, D., Stidworthy, M. & Zell, R. (2020) Novel enteric viruses in fatal enteritis of grey squirrels. Journal of General Virology.

Astro- and kobuviruses infect both humans and animals. Here, we report on the disease history, detection and genomic characterization of novel astro- and kobuviruses from fatal diarrhoea of two juvenile grey squirrels. The virus particles had enterovirus-like morphology and a diameter of 28-32 nm. Next-generation sequencing confirmed astro- and kobuviruses and sequence analysis revealed typical astrovirus and picornavirus genome organizations. The astrovirus ORF2 sequence clustered with a clade of unassigned astroviruses, with marmot and rodent mamastroviruses as closest relatives. For the kobuvirus, divergences greater than 49.4 % for P1 and 43.5 % in the non-structural proteins indicated a novel species. However, phylogenetic analysis of the 3D polymerase showed that it clustered with that of the newly classified ludopivirus A1, suggesting a previous recombination event in the evolution of the kobuvirus. Our data provide further insights into the diversity of astro- and kobuviruses and broaden the spectrum of viruses infecting grey squirrels.

• Deb, J. C., Forbes, G. & MacLean, D. A. (2020) Modelling the spatial distribution of selected North American woodland mammals under future climate scenarios. Mammal Review 50(4): 440-452.

Abstract: North America has a diverse array of mammalian species. Model projections indicate significant variations in future climate conditions of North America, and the habitats of woodland mammals of this continent may be particularly sensitive to changes in climate. We report on the potential spatial distributions of 13 wide-ranging, relatively common species of North American woodland mammals under future climate scenarios. We examined the potential influence of the mean and seasonal climate variables on the distribution of species. Presence-only occurrence records of species, four predictor variables, two future climate scenarios (Representative Concentration Pathways 4.5 and 8.5), and two time steps (current and 2070) were used to build species? distribution models using a maximum entropy algorithm (MaxEnt). Our results suggested that overall, 11 of the 13 species are likely to gain climatically suitable space (regions where climate conditions will be similar to those of area currently occupied) at the continental scale, but American marten Martes americana and ?woodland? caribou Rangifer tarandus are likely to lose suitable climate range by 2070. Furthermore, climate space is likely to be expanding northwards under future climate scenarios for most of the mammals, and many jurisdictions in the border region between Canada and the USA are likely to lose iconic species, such as moose Alces alces. We identified regions as potential in situ and ex situ climate change refugia, which are increasingly considered to be important for biodiversity conservation. The model results suggest significant implications for conservation planning for the 13 mammalian species under global climate change, especially at fine spatial scales. Numerous species that are presently common at their southern range edge will be functionally or completely extirpated in 50 years. The potential in situ and ex situ climate change refugia could provide an effective support for adaptive strategies aimed at species conservation planning.

• Fotis, A., Patel, S. & Chavez, A. (2020) Habitat-based isolating barriers are not strong in the speciation of ecologically divergent squirrels (Tamiasciurus douglasii and T. hudsonicus). Behavioral Ecology and Sociobiology 74.

Abstract: Adaptations to different environments between closely related species can be important drivers of reproductive isolation during speciation due to either habitat isolation between species or reduced fitness in hybrids that possess suboptimal adaptations. Hybrid zones are useful natural arenas to explore how ecologically divergent species compete for habitat in sympatry and how possible differences in their habitat use may contribute to the speciation process. We investigated habitat selection by sister species of tree squirrels, the Douglas squirrel and the red squirrel, that have evolved in different forest types in allopatry and hybridize in a transitional forest. We first used genome-wide SNP data and admixture analyses to classify individuals into parental or hybrid classes. Next, we estimated home ranges with radio telemetry data and then used a novel ground-based lidar system to measure forest canopy structure of squirrel home ranges, midden sites, and marginally used forest habitat. We found hybrids consisting of multiple hybrid classes were intermixed with both parental species in the same forest with varying canopy structure complexity. On average, Douglas squirrels utilized forests with slightly greater structural complexity than either red squirrels or hybrids, while marginally used forests were the least structurally complex. Interestingly, hybrid squirrels were not relegated to marginal habitat and were successful in mating among each other and with both parental species. As such, our study suggests that prezygotic-ecological isolation and postzygotic-hybrid infertility, and postzygotic ecological inviability of hybrids are not strong barriers in the speciation process between Douglas squirrels and red squirrels.

• Hämäläinen, S., Fey, K. & Selonen, V. (2020) Search strategies in rural and urban environment during natal dispersal of the red squirrel. Behavioral Ecology and Sociobiology 74(10): 124.

Abstract: Decisions made during natal dispersal highly influence the future fitness of the animal. The selection of future home range affects, for example, food supply and mating success. In order to select between potential sites, dispersing individuals may use different search strategies, and for example, aim to compare quality of different sites by revisiting them. Alternatively, dispersers visit new sites without comparison until a suitable site is located. By using radio telemetry, we studied search strategies during natal dispersal in both urban and rural environments. We examined what kind of sites juvenile red squirrels (Sciurus vulgaris) choose as their home range from all visited sites and whether they display revisit behavior during natal dispersal. We found that most of the dispersing individuals perform revisits while searching for the settlement site. In the rural study area, there were also individuals (mostly long-distance dispersers) that did not perform revisits and settled in the last visited area. The landscape characteristics did not explain the amount of revisits individuals made, but the amount of revisits declined with site’s distance to natal nest and dispersal distance of individuals. The effect of landscape characteristics on settlement decision was small, but surprisingly, juvenile red squirrels in the rural area settled in sites with more built area compared with sites only visited. However, the decision-making of red squirrels during natal dispersal seems to be driven mainly by other factors (potentially food availability and conspecific density) than landscape characteristics. We conclude that revisiting behavior is linked to dispersal strategy of red squirrels and can differ between the sexes and habitats of the species.

• Herzschuh, U. (2020) Legacy of the Last Glacial on the present-day distribution of deciduous versus evergreen boreal forests. Global Ecology and Biogeography 29(2): 198-206.

Abstract: Issue Despite their rather similar climatic conditions, eastern Eurasia and northern North America are largely covered by different plant functional types (deciduous or evergreen boreal forest) composed of larch or pine, spruce and fir, respectively. I propose that these deciduous and evergreen boreal forests represent alternative quasi-stable states, triggered by their different northern tree refugia that reflect the different environmental conditions experienced during the Last Glacial. Evidence This view is supported by palaeoecological and environmental evidence. Once established, Asian larch forests are likely to have stabilized through a complex vegetation?fire?permafrost soil?climate feedback system. Conclusion With respect to future forest developments, this implies that Asian larch forests are likely to be governed by long-term trajectories and are therefore largely resistant to natural climate variability on time-scales shorter than millennia. The effects of regional human impact and anthropogenic global warming might, however, cause certain stability thresholds to be crossed, meaning that irreversible transitions occur and resulting in marked consequences for ecosystem services on these human-relevant time-scales.

• Hollingsworth, P., O’Brien, D., Ennos, R., Ahrends, A., Ballingall, K., Brooker, R., Burke, T., Cavers, S., Dawson, I., Elston, D., Kerr, J., Marshall, D., Neaves, L., Pakeman, R., Trivedi, C., Wall, E., Wright, F., Yahr, R., Bean, C. & Ogden, R. (2020) Scotland’s Biodiversity Progress to 2020 Aichi Targets: Aichi Target 13 – Genetic Diversity Maintained – Supplementary Report 2020.

Abstract: As there is no agreed national list of species of socio-economic and/or cultural value for Scotland, a set of criteria for selecting species has been developed. These include: • Species prioritised for conservation value • Species identified as being culturally important • Species providing important ecosystem services • Game species • Species collected for food or medicine Using these criteria, 26 species were selected for initial assessment. For each species, we produced a Genetic Scorecard, outlining: • Relevant genetic conservation issues for the species in question • The importance of its genetic diversity on an international scale • An evaluation of the genetic risks facing in situ populations • A statement of confidence in the assessment • The degree to which representation in ex situ collections mitigate against genetic diversity loss (further explanation of these maps is provided in Section 4 of the report). • An overall ‘traffic light’ score of genetic risks and whether current conservation actions are effective.

• Jezkova, T. (2020) Founder takes more: Interspecific competition affects range expansion of North American mammals into deglaciated areas. Journal of Biogeography n/a(n/a).

Abstract: Aim I assess the impact of interspecific competition on species’ post-glacial range expansions into previously glaciated areas. I hypothesize that expansion of one species (the founder) after the last glacial maximum (LGM) into areas that were glaciated has hindered expansion of its competitor (the latecomer). If true, I predict that range and niche sizes of two congeners are more disproportionate within the previously glaciated areas (i.e. the areas of post-glacial expansion) than within areas that were not glaciated. I also predict that niche partitioning is reduced within previously glaciated areas. If true, range and climatic niche overlap of two congeners is smaller in the previously glaciated areas than in non-glaciated areas. Location North America. Taxon Non-volant mammals. Methods I calculate species range size for 87 congeneric species pairs and climatic niche hypervolume for 71 pairs, separately for areas that were and were not glaciated during the LGM. I then compare (a) range and climatic niche size ratio and (b) range and climatic niche overlap for each species pair between areas that were and were not glaciated using a paired t test. Results Congeneric species pairs exhibit more disproportionate range areas and climatic niche hypervolumes, and smaller range and climatic niche overlap within previously glaciated areas, in comparison with areas that were not glaciated during the LGM. Main conclusions Interspecific competition has likely hindered post-glacial range expansion of some species into deglaciated areas. Similarly to the founder effect documented on the intraspecific level, the founder species may hinder the expansion of the latecomer. Consequently, one species may expand over very large areas whereas its competitor may expand very little or not at all. Additionally, niche partitioning within the deglaciated areas may be reduced as the founder expands and fills biotopes that would otherwise be utilized by its competitor.

• Johnston, A., Vander Haegen, M. & West, S. (2020) Differential Resource Use between Native and Introduced Gray Squirrels. The Journal of Wildlife Management 84: 726-738.

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. Introduced eastern gray squirrels share affinity for oaks with native western gray squirrels but are limited to riparian areas with diverse deciduous trees and shrubs in Washington. In contrast, western gray squirrels frequently inhabit conifer‐dominated uplands, which limits interspecies interactions and provides opportunities for managers to exploit habitat differences between species to conserve western gray squirrels.

• Kostrzewa, A. & Krauze-Gryz, D. (2020) The choice and handling of supplemental food by red squirrels in an urban park. Behavioural Processes: 104153.

One of the main factors that allow the red squirrel (Sciurus vulgaris) to live in cities may be the availability of supplemental food provided by people. In this article, preferences of red squirrels in relation to supplemental food, how it is handled, and whether change seasonally, is investigated. A cafeteria test was performed, in which squirrels could choose one from six kind of nuts. Three types of nuts were used: walnuts, hazelnuts and peanuts, with unshelled and shelled nuts of each offered. In most cases squirrels took an offered nut. If a nut was not taken, it was mostly due to lack of interest, without signs of fear. Regardless of season, whole, unshelled walnuts were selected most frequently – the most high-energy and least perishable from offered food. The least frequently selected overall were peanuts. Caching behaviour changed seasonally – nuts were mostly eaten in spring and hoarded in autumn and it took less time to cache nuts in autumn and winter. It may reflect optimal caching strategy at the time when food is abundant and there is a desire to secure as big a store of food as possible. Results from the study suggest that supplemental food is an important component of squirrels’ diet and have an impact on their caching strategies.

• Lawton, C., Hanniffy, R., Victoria, Molloy, Guilfoyle, C., Michael, Stinson & Reilly, E. (2020) All-Ireland squirrel and pine marten survey 2019. Irish Wildlife Manuals, No. 121. National Parks and Wildlife Service, Department of Culture, Heritage and the Gaeltacht, Ireland.

Executive Summary: In 2019, a survey was conducted across the island of Ireland on the distribution and status of three mammal species, the native Red Squirrel Sciurus vulgaris and Pine Marten Martes martes, and the invasive Grey Squirrel Sciurus carolinensis. The Grey Squirrel was introduced to Ireland just over 100 years ago, and has since spread to cover most of the eastern half of the island. The Grey Squirrel competes with the Red Squirrel for resources, and carries a disease that is fatal to the native species. As a result, the spread of Grey Squirrel had been mirrored by a retraction in Red Squirrel range and a reduction in numbers. The distributions of the two squirrel species have been observed through a series of surveys running since the middle of the 20th century. In recent surveys however, it became apparent that in some parts of Ireland there had been a change in the fortunes of the two species. In surveys conducted in 2007 and 2012 it was evident that the Grey Squirrel had disappeared from parts of the midlands of Ireland. This disappearance was linked in subsequent studies to the re-emergence of the Pine Marten, a native carnivore of Ireland. The Pine Marten had previously almost disappeared but has made a considerable recovery after becoming protected under Irish and EU legislation. High densities of Pine Marten were found in the midlands of Ireland, in the areas where Grey Squirrel had disappeared. Meanwhile, Red Squirrel has made a recovery in some of these areas and seemed capable of co-habiting woodlands with the native carnivore. The current survey sought to update the distribution maps for the three species in recognition of the rapidly changing situation. The survey team was a collaborative effort between academics and NGO officers from both Republic of Ireland and Northern Ireland, and used the online recording facilities offered by the National Biodiversity Data Centre (RoI) and Centre for Environmental Data and Recording (NI). The survey was publicised using print, broadcasting and social media, with the public invited to contribute sightings as part of a citizen science effort. Three thousand four hundred and seven records were received during the course of the survey, a large increase on the number received in previous surveys. Grey Squirrel records however had decreased in number, and the range covered by the species had dropped considerably, with 37.8% fewer hectads (10 km x 10 km squares) occupied than in 2012. The distribution gap in the midlands now stretches across nine counties, with Grey Squirrel having functionally disappeared from Co. Fermanagh, Co. Monaghan and parts of Co. Meath and Co. Kildare since the 2012 survey. Red Squirrel sightings have increased considerably, and it has returned to parts of the midlands from which it had disappeared. The number of sightings of Pine Marten, previously considered an elusive animal, was very high, with signs that its core range has expanded from the west and midlands to include parts of Northern Ireland and Co. Wicklow. Again, there is evidence of the negative correlation between the Grey Squirrel and Pine Marten, and it is in the areas where the Pine Marten has increased most significantly that this squirrel has disappeared. The Grey Squirrel continues to thrive in some areas, in particular in urban regions around Belfast and Dublin. There was some evidence of further Grey Squirrel spread into the southwest of the island. The survey was very successful and underlined the benefits of citizen science and using the collective knowledge of an informed and enthusiastic public. The collaborative nature of the study, bringing together colleagues from institutions across both parts of the island, was very productive. The status of the two native species is very good, with very important populations in their strongholds in the west and midlands of Ireland. However, recommendations are made to ensure that this remains the case, with further monitoring required to allow early intervention if conservation at a local or national level is required

• Leaver, L., Ford, S., Miller, C., Yeo, M. & Fawcett, T. (2020) Learning is negatively associated with strength of left/right paw preference in wild grey squirrels (Sciurus carolinensis). Learning and Behaviour.

Abstract: Cerebral laterality, via hemispheric specialisation, has been evidenced across the animal kingdom and linked to cognitive performance in a number of species. Previously it has been suggested that cognitive processing is more efficient in brains with stronger hemispheric differences in processing, which may be the key fitness benefit driving the evolution of laterality. However, evidence supporting a positive association between cognitive performance and lateralization is mixed: data from studies of fish and birds show a positive relationship whereas more limited data from studies of mammals suggest a weak or even negative relationship, suggesting the intriguing possibility of a mammal/non-mammal divide in the nature of this relationship. Here, we report an empirical test examining the relationship between lateralization and cognitive performance in wild grey squirrels (Sciurus carolinensis) by measuring left/right paw preference as a behavioural assay of cerebral lateralization and learning speed as an assay of cognitive efficiency. We carried out a motor-based laterality test using a reaching paradigm and measured learning speed on a problem-solving task. In accordance with the suggestion of a mammal/non-mammal divide, we found a negative relationship between strength of paw preference and performance on the learning task. We discuss this finding in light of niche-specific adaptations, task-specific demands and cognitive flexibility.

• Leaver, L., Lea, S., Chow, P. & McLaren, I. (2020) Behavioral Flexibility: a review, a model and some exploratory tests. Learning and Behaviour

Abstract: This paper aims to explore and clarify the concept of behavioral flexibility. A selective literature review explores how the concept of behavioral flexibility has been used in ways that range from acknowledging the fact that animals’ behavior is not always bounded by instinctual constraints, to describing the variation between species in their capacity for innovative foraging, a capacity that has repeatedly been linked to having a brain larger than would be predicted from body size. This wide range of usages of a single term has led to some conceptual confusion. We seek to find a more precise meaning for behavioral flexibility, by representing it within a simple formal model of problem solving. The key to our model is to distinguish between an animal’s state of knowledge about the world and its observable behavior, using a construct of response strength to represent that underlying knowledge. We model behavioral flexibility as a parameter in the function that transforms response strengths into observable response probabilities. We test this model in simulations based on some recent experimental work on animal problem solving. Initial results show that parametric manipulation can mimic some of the behavioral effects that have been attributed to flexibility.

Lurz, P. & Kortland, K. (2020) An assessment of long-term forest management policy options for red squirrel conservation in Scotland. Hystrix

A spatially explicit mathematical model was developed to assess the population viability of red squirrels (Sciurus vulgaris) in designated forest strongholds in Scotland under the implementation of two forest management policies: a specific Stronghold Management for red squirrel conservation (SM) compared to the multi-purpose UK Forestry Standard (UKFS) for sustainable forest management. The study showed that, in the presence of grey squirrels (Sciurus carolinensis), the SM policy provides an advantage to red squirrels over grey squirrels when compared to the UKFS, and its implementation supports red squirrel conservation efforts. When grey squirrels are not present, there is no discernible benefit in the SM policy compared to the UKFS. The model results therefore indicate that species-specific forest management for red squirrel conservation in the absence of sympatric grey squirrels would not be required. This would allow less prescriptive forest management options that maintain viable red squirrel populations to be explored. The study also identified forest regions that, due to their composition, are capable of sustaining a viable red squirrel population, in the presence of grey squirrels, without the application of specific forest management policy. They can be considered ‘natural strongholds’. Selecting such natural strongholds may afford more flexibility to conserve red squirrel populations whilst simultaneously delivering other multi-species conservation and forest management objectives. We review our findings in terms of criteria that were used in the original stronghold designation in Scotland and discuss how our work can be used to inform a forthcoming review of stronghold management policy by Scottish Forestry. Furthermore, the findings can inform red squirrel conservation strategies in other regions and the modelling techniques can be adapted to a wide range of conservation settings.

• Maranesi, M. (2020) Seasonal Expression of NGF and Its Cognate Receptors in the Ovaries of Grey Squirrels (Sciurus carolinensis). Animals 10: 1558.

The grey squirrel is an invasive alien species that seriously threatens the conservation of the native red squirrel species. With the aim of characterizing the reproductive physiology of this species due to its great reproductive success, the function of the ovarian nerve growth factor (NGF) system was analyzed in a grey squirrel population living in central Italy. During the breeding and nonbreeding seasons, the ovarian presence, distribution, and gene expression of NGF, neurotrophic tyrosine kinase receptor 1 (NTRK1), and nerve growth factor receptor (NGFR), as well as NGF plasma concentrations, were evaluated in female grey squirrels. NGF was found in the luteal cells and in the thecal and granulosa cells of follicles, while NTRK1 and NGFR were only observed in follicular thecal and granulosa cells. NGF and NGFR transcripts were almost two-fold greater during the breeding season, while no seasonal differences were observed in NTRK1 gene expression. During the breeding season, NGFR was more expressed than NTRK1. Moreover, no changes were observed in NGF plasma levels during the reproductive cycle. The NGF system seems to be involved in regulating the ovarian cycle mainly via local modulation of NGF/NGFR, thus playing a role in the reproductive physiology of this grey squirrel population.

• Maranesi, M. & Mercati, F. (2020) Reproductive Traits of an Invasive Alien Population of Grey Squirrel (Sciurus carolinensis) in Central Italy. Animals 10: 738.

Abstract: The reproductive cycle of an invasive alien Italian grey squirrel population was studied to understand its adaptation and limit its spread, in order to conserve the autochthonous red squirrel. Female and male genital traits were evaluated throughout the reproductive cycle, including the ovary, uterus, testicle, epididymis, seminiferous tubule morphometry, and germinative epithelium histology. Moreover, individual female fecundity was determined by counting uterine scars. Ovary width and uterus weight, length, and width reached their highest values in the luteal and pregnancy phases. On conducting a histological evaluation of the testicular germinal epithelium, four morphotypes related to the different reproductive phases of the male squirrels were identified: immature, pubertal, spermatogenesis, and regressive. Testicle and epididymis weights and seminiferous tubule diameters reached their largest values during spermatogenesis. Uterine scar analysis showed that 69% of the females had given birth to one or two litters, while 31% had no uterine scars. Litters were larger in the first breeding period than in the second; annual fecundity was 4.52 ± 1.88 uterine scars/female. Umbrian grey squirrels have adapted to their non-native range, showing two annual mating periods at times similar to those in their native range, and high reproductive success.

• McNicol, C. M., Bavin, D., Bearhop, S., Bridges, J., Croose, E., Gill, R., Goodwin, C. E. D., Lewis, J., MacPherson, J., Padfield, D., Schofield, H., Silk, M. J., Tomlinson, A. J. & McDonald, R. A. (2020) Postrelease movement and habitat selection of translocated pine martens Martes martes. Ecology and Evolution n/a(n/a).

Abstract: Monitoring postrelease establishment and movement of animals is important in evaluating conservation translocations. We translocated 39 wild pine martens Martes martes (19 females, 20 males) from Scotland to Wales. We released them into forested areas with no conspecifics in 2015, followed by a second release in 2016, alongside the previously released animals. We used radio-tracking to describe postrelease movement and habitat selection. Six martens (15%) were not re-encountered during the tracking period, of which four undertook long-distance dispersal. For the remaining individuals, we characterized two phases of movement, ?exploration? followed by ?settlement,? that differed between releases. In the first release, martens remained in exploration phase for a mean of 14.5 days (SE = 3.9 days) and settled at a mean distance of 8.7 km (SE = 1.8 km) from release sites, whereas martens released in year two, alongside resident conspecifics, traveled away from release sites at a faster rate, settling sooner, at a mean of 6.6 days (SE = 1.8 days), but further, at a mean distance of 14.0 km (SE = 1.7 km) from release sites. Animals released in year one did not exhibit habitat preferences overall but within forests they favored recently felled areas, whereas animals released in year two showed strong selection for forested habitat but did not discriminate between forest types. The presence of conspecifics appeared influential for settlement and site fidelity of translocated martens and was associated with more rapid but more distant dispersal of the later cohort. Releases of animals in close proximity appeared to promote site fidelity and rapid establishment of ranges in the recipient environment.

• McNicol, C. M., Bavin, D., Bearhop, S., Ferryman, M., Gill, R., Goodwin, C. E. D., MacPherson, J., Silk, M. J. & McDonald, R. A. (2020) Translocated native pine martens Martes martes alter short-term space use by invasive non-native grey squirrels Sciurus carolinensis. Journal of Applied Ecology 57(5): 903-913.

Abstract Predators can shape the distributions and dynamics of their prey through direct and indirect mechanisms. Where prey animals are regarded as pests, the augmentation of predator populations might offer a potential tool in their management. Declines in invasive non-native grey squirrel Sciurus carolinensis populations in Ireland and Scotland have been related to an increase in range and density of native pine marten Martes martes populations. These reductions in grey squirrel abundance have, in turn, been linked to recovery of native red squirrels Sciurus vulgaris. Taking the opportunity presented by a conservation translocation of pine martens from Scotland to Wales, we investigated the short-term effects of exposure to translocated martens on the space use and survival of resident grey squirrels. Grey squirrel range size and daily distance travelled increased significantly with increasing exposure to martens but we found no effect of marten exposure on the recapture probability (i.e. apparent survival) of the sampled squirrels within the study time frame. This is suggestive of contemporary, non-lethal effects changing the ranging or foraging regimes of squirrels, due either to predator avoidance and/or earlier lethal effects associated with a reduction in intraspecific competition. Synthesis and applications. Our evaluation mimics the conditions experienced by grey squirrels at the front edge of naturally recovering pine marten populations and presents direct evidence that pine marten translocations could play an influential role in the behaviour and dynamics of invasive non-native grey squirrel populations. Translocations of native predators, undertaken primarily for biodiversity conservation, could therefore find additional application in managing the ecological and economic impacts of invasive non-native prey.

• Mead, D., Fingland, K., Cripps, R., Miguez, R., Smith, M., Corton, C., Oliver, K., Skelton, J., Betteridge, E., Dolucan, J., Dudchenko, O., Omer, A., Weisz, D., Aiden, E., Fedrigo, O., Mountcastle, J., Jarvis, E., McCarthy, S., Sims, Y. & Blaxter, M. (2020) The genome sequence of the Eurasian red squirrel, Sciurus vulgaris Linnaeus 1758. Wellcome Open Research 5: 18.

Abstract: We present a genome assembly from an individual male Sciurus vulgaris (the Eurasian red squirrel; Vertebrata; Mammalia; Eutheria; Rodentia; Sciuridae). The genome sequence is 2.88 gigabases in span. The majority of the assembly is scaffolded into 21 chromosomal-level scaffolds, with both X and Y sex chromosomes assembled.

• Menéndez, I., Gómez Cano, A. R., Cantalapiedra, J. L., Peláez-Campomanes, P., Álvarez-Sierra, M. Á. & Hernández Fernández, M. (2020) A multi-layered approach to the diversification of squirrels. Mammal Review n/a(n/a).

Abstract: The shape of the tree of life is the result of shifting diversification rates, and identifying the factors driving these shifts is one of the main aims in evolutionary biology. Various biotic and abiotic factors have been proposed to have an impact on mammal diversification, such as climatic and tectonic changes, the acquisition of new traits, and expansion into new ecosystems or landmasses. We used phylogenetic comparative methods to quantify the influence of potential drivers on diversification patterns in extant squirrels (Sciuridae, Rodentia). We conducted a multilayer approach, comparing diversification rates among squirrel lineages depending on their degree of biome specialisation, biogeographic realm occupancy, locomotion adaptations, and presence in mountainous regions. We generated the most complete phylogeny of squirrels to date, encompassing almost 80% of the extant species, and applied multiple and binary state-dependent diversification models. All the traits examined showed an influence on diversification rates. The biome specialist lineages showed the highest speciation rates, suggesting a major role of bioclimatic specialisation on macroevolutionary patterns. A single major event, the Miocene?Pleistocene radiation of terrestrial-adapted lineages in North America, left a signal that was recovered in two of our analyses. Both the Nearctic lineages and the terrestrial-adapted lineages showed high speciation rates, highlighting the fact that that major evolutionary episodes may produce confounding effects in state-dependent diversification models. Ancestral reconstructions showed that cold and warm intervals in Earth?s history had different effects on squirrels? diversification, depending on their climatic affinities. Tropical and arboreal squirrels evolved predominantly in the warm intervals, while terrestrial and cold-adapted squirrels radiated in the cold intervals. Our findings suggest that, while global climatic shifts are key for the speciation processes in mammalian lineages, lineage-specific ecological adaptations are critical modulators of the responses of lineages to such environmental shifts, in an interplay that ultimately affects their diversification patterns.

• Menzies, A. K., Studd, E. K., Majchrzak, Y. N., Peers, M. J. L., Boutin, S., Dantzer, B., Lane, J. E., McAdam, A. G. & Humphries, M. M. (2020) Body temperature, heart rate, and activity patterns of two boreal homeotherms in winter: Homeostasis, allostasis, and ecological coexistence. Functional Ecology n/a(n/a).

Abstract Organisms survive environmental variation by combining homeostatic regulation of critical states with allostatic variation of other traits, and species differences in these responses can contribute to coexistence in temporally variable environments. In this paper, we simultaneously record variation in three functional traits?body temperature (Tb), heart rate and activity?in relation to three forms of environmental variation?air temperature (Ta), photoperiod and experimentally manipulated resource levels?in free-ranging snowshoe hares and North American red squirrels to characterize distinctions in homeotherm responses to the extreme conditions of northern boreal winters. Hares and squirrels differed in the level and precision of Tb regulation, but also in the allostatic pathways necessary to maintain thermal homeostasis. Hares demonstrated a stronger metabolic pathway (through heart rate variation reflective of the thermogenesis), while squirrels demonstrated a stronger behavioural pathway (through activity variation that minimizes cold exposure). As intermediate-sized, winter-active homeotherms, hares and squirrels share many functional attributes, yet, through the integrated monitoring of multiple functional traits in response to shared environmental variation, our study reveals many pairwise species differences in homeostatic and allostatic traits, that both define and are defined by the natural history, functional niches and coexistence of sympatric species. A free Plain Language Summary can be found within the Supporting Information of this article.

• Montgomery, I. (2020) The dynamics of pine marten predation on red and grey squirrels. Mammalian Biology – Zeitschrift für Säugetierkunde: 3.

Invasive alien species pose one of the greatest threats to global biodiversity. In parts of Europe, introduced eastern grey squirrels (Sciurus carolinensis) have caused regional extinctions of the native red squirrel (Sciurus vulgaris). However, exposure to pine martens (Martes martes) has been demonstrated to reverse the competitive outcome between red and grey squirrels. The mechanism whereby this effect occurs remains unclear. It is hypothesised that direct predation, facilitated by a lack of behavioural response, is the mechanism driving this relationship. We review the literature and reanalyse a new dataset to provide further data on the occurrence of both squirrel species in the scats of pine marten. Both squirrel species occurred in the scats of pine marten confirming its role as a predator of these species. Predation of grey squirrels was significantly higher than red squirrels and was recorded only in spring and summer. Our review provides evidence for the mechanism driving the current decline in grey squirrels in Ireland and Scotland and supports the hypothesis that in the presence of a shared predator, direct predation influences the outcome of species interactions between native red and non-native grey squirrels.

• Nannarone, S., Moretti, G., Bellocchi, F., Menchetti, L. & Bufalari, A. (2020) A Comparative Study of Intramuscular Alfaxalone- or Ketamine-Based Anesthetic Mixtures in Gray Squirrels Undergoing Gonadectomy: Clinical and Physiologic Findings. Animals 10: 1402.

The gray squirrel is one of the most common invasive species in Europe, whose presence is dangerous for the survival of the European red squirrel. To cope with this biological invasion and to safeguard biodiversity, the LIFE+U-SAVEREDS project aims to protect the red squirrel, by limiting the growth of the current population of gray squirrels and simultaneously promoting their eradication with surgical sterilization. This study compares two different anesthetic protocols, including dexmedetomidine (40 µg/kg) and midazolam (0.3 mg/kg) associated with ketamine (15 mg/kg; n = 25 squirrels) or alfaxalone (5 mg/kg; n = 22 squirrels). A blinded investigator evaluated the quality and onset of sedation, intraoperative anesthesia, and recovery, as well as the physiologic parameters for each animal. Alfaxalone provided a good quality of anesthesia with limited cardiovascular effects (p < 0.05) and good intraoperative myorelaxation. Ketamine induced complete relaxation in a shorter time (p < 0.05) and a rapid (p < 0.001) and excellent (p < 0.05) recovery. Despite the overall superiority of ketamine, alfaxalone appeared to be an adequate alternative anesthetic drug that can be administered without requiring intravascular access. It should be rapidly metabolized and excreted; however, it requires the combination of longer acting sedatives/myorelaxants to prevent a poor recovery quality.

• Niu, H., Wang, Z., Huang, G., Peng, C., Zhang, Z. & Zhang, H. (2020) Responses of a scatter-hoarding squirrel to conspecific pilfering: a test of the reciprocal pilferage hypothesis. Animal Behaviour 170: 147-155.

Abstract: The reciprocal pilferage hypothesis argues that, despite the occurrence of pilferage, scatter-hoarding behaviour could evolve if cache loss is compensated by gains pilfered from others. However, this model has not been strictly tested because of the difficulty associated with tracking caches, including ownership, over long periods. Using infrared radiation cameras, we tracked caching of Juglans regia seeds by groups of two Père David’s rock squirrels, Sciurotamias davidianus, within a large natural enclosure. Our goal was to quantify how squirrels responded to the presence of a conspecific when seed hoarding and pilfering and test the reciprocal pilferage hypothesis. We found that the numbers of seeds harvested from the seed sources and pilfered from conspecifics was initially low, increasing as seeds were removed and then dropping off once few seeds remained at the seed sources. Additionally, the number of seeds scatter hoarded increased with a decreasing number of seeds remaining at the sources. Seeds harvested from the source and pilfered from competitors were at first cached randomly within the enclosure and then centralized to the low-competition area near the nest as seed sources declined. Overall, pilferage was not high. The proportions of seeds pilfered did not vary between conspecifics and were positively correlated with each other over trials. More seeds were harvested from the seed sources than pilfered by competitors at the early stage of hoarding. These results suggest that, under conspecific competition, squirrels appeared first to compete for food at the sources, then for caches with each other as food sources decreased, and finally cache ownership became relatively stable. Squirrels compensated for cache loss by both harvesting food from the sources and pilfering caches from its competitor. The amount of seeds gained from pilfering was sufficient to replace pilfered caches, supporting the reciprocal pilferage hypothesis.

• Nyakatura, J. A., Baumgarten, R., Baum, D., Stark, H. & Youlatos, D. (2019) Muscle internal structure revealed by contrast-enhanced μCT and fibre recognition: The hindlimb extensors of an arboreal and a fossorial squirrel. Mammalian Biology 99: 71-80.

In individuals of similar body mass representing closely related species with different lifestyles, muscle architectural properties can be assumed to reflect adaptation to differing, lifestyle-related functional demands. We here employ a fibre recognition algorithm on contrast-enhanced micro-computed tomography (μCT) scans of one specimen each of an arboreal (Sciurus vulgaris) and a fossorial (Spermophilus citellus) sciuromorph rodent. The automated approach accounts for potential heterogeneity of architectural properties within a muscle by analysing all fascicles that compose a muscle. Muscle architectural properties (volume, fascicle length and orientation, and force-generating capacity) were quantified in 14 hindlimb (hip, knee, and ankle) extensor muscles and compared between specimens. We expected the arboreal squirrel to exhibit greater force-generating capacity and a greater capacity for length change allowing more powerful hindlimb extension. Generally and mostly matching our expectations, the S. vulgaris specimen had absolutely and relatively larger extensor muscles than the S. citellus specimen which were thus metabolically more expensive and demonstrate the relatively larger investment into powerful hindlimb extension necessary in the arboreal context. We conclude that detailed quantitative data on hindlimb muscle internal structure as was gathered here for a very limited sample further lends support to the notion that muscle architecture reflects adaptation to differential functional demands in closely related species with different locomotor behaviours and lifestyles.

• Paniw, M., Childs, D. Z., Armitage, K. B., Blumstein, D. T., Martin, J. G. A., Oli, M. K. & Ozgul, A. (2020) Assessing seasonal demographic covariation to understand environmental-change impacts on a hibernating mammal. Ecology Letters 23(4): 588-597.

Abstract: Natural populations are exposed to seasonal variation in environmental factors that simultaneously affect several demographic rates (survival, development and reproduction). The resulting covariation in these rates determines population dynamics, but accounting for its numerous biotic and abiotic drivers is a significant challenge. Here, we use a factor-analytic approach to capture partially unobserved drivers of seasonal population dynamics. We use 40 years of individual-based demography from yellow-bellied marmots (Marmota flaviventer) to fit and project population models that account for seasonal demographic covariation using a latent variable. We show that this latent variable, by producing positive covariation among winter demographic rates, depicts a measure of environmental quality. Simultaneously, negative responses of winter survival and reproductive-status change to declining environmental quality result in a higher risk of population quasi-extinction, regardless of summer demography where recruitment takes place. We demonstrate how complex environmental processes can be summarized to understand population persistence in seasonal environments.

• Parker, A. & Benkman, C. (2020) Enhanced seed defenses potentially relax selection by seed predators against serotiny in lodgepole pine. Ecology and Evolution.

Abstract: Serotiny, the retention of seeds in a canopy seed bank until high temperatures cause seeds to be released, is an important life history trait for many woody plants in fire‐prone habitats. Serotiny provides a competitive advantage after fire but increases vulnerability to predispersal seed predation, due to the seeds being retained in clusters in predictable locations for extended periods. This creates opposing selection pressures. Serotiny is favored in areas of high fire frequency, but is selected against by predispersal seed predators. However, predation also selects for cone traits associated with seed defense that could reduce predation on serotinous cones and thereby relax selection against serotiny. This helps explain the elevated defenses in highly serotinous species. However, whether such interactions drive variation in seed defenses within variably serotinous populations has been studied rarely. We investigated the effects of phenotypic selection exerted by red squirrel (Tamiasciurus hudsonicus) predation on Rocky Mountain lodgepole pine (Pinus contorta latifolia) seeds. Squirrels preferentially harvested cones with more and larger seeds, indicating a preference for a higher food reward. We found evidence for stronger selection on trees with serotinous cones, which presumably accounts for the elevated defenses of and lower predation on serotinous compared to non‐serotinous cones. Lower levels of predation on serotinous cones in turn lessen selection against serotiny by squirrels. This has important implications because the frequency of serotiny in lodgepole pine has profound consequences for post‐fire communities and ecosystems widespread in the Rocky Mountains.

• Passantino, G., Tursi, M., Vercelli, C., Filippi, I., DeCaro, N., Tinelli, A., Valente, L., Leone, R. & Zizzo, N. (2020) Systematic Pathologic Findings Report of Callosciurus finlaysonii (Horsfield, 1823) (Rodentia, Sciuridae) Squirrels from Maratea area (South Italy) to Investigate Species-Specific Pathologies, Reliability of CO2 Euthanasia Method, and Possible Use as Environmental Sentinels. Animals 10(10).

The aim of the present study was to macroscopically and microscopically describe the main pathological findings occurring in this invasive species, in order to better understand the real risks for naïve animals and humans. The present study was conducted on Callosciurus finlaysonii squirrels (n = 165), captured in the Maratea area and euthanatized with CO2 according to a population control of invasive species of the Italian Agriculture Ministry project (ex CIPE project) and conferred to the Department of Veterinary Medicine of Bari (Italy). Macroscopic analysis demonstrated heart, lung, and liver congestion, probably due to the euthanasia method, and variable lesions of bowel, liver, and kidney. The microscopically examination showed the presence of lymphocytic infiltration in the lower layers of the bowel, suggesting enteritis. To the best of our knowledge, this is the first systemic report of gross and microscopical anatomopathological lesions in C.finlaysonii, in South Italy. The results could be useful to fill a gap of knowledge of this species in Italy.

• Peplinski, J. & Brown, J. S. (2020) Distribution and diversity of squirrels on university and college campuses of the United States and Canada. Journal of Mammalogy 101(4): 930-940.

Abstract: Squirrels on North American college and university campuses have been the subject of much animated conversation in informal venues, but a systematic assessment of their distribution across this habitat type has not been undertaken until now. We collected reports of squirrel species’ presence and absence from faculty experts at 536 campuses in Canada and the continental United States, and found that squirrels are nearly ubiquitous on campuses (95% had at least one species, and 40% had three or more), but that only a select few species are common campus residents. Foremost among these is the eastern gray squirrel (Sciurus carolinensis—on 62% of surveyed campuses), followed by the eastern fox squirrel (Sciurus niger), eastern chipmunk (Tamias striatus), North American red squirrel (Tamiasciurus hudsonicus), and woodchuck (Marmota monax). Eastern gray and fox squirrels partition campuses with one another, both in their shared native/core range and on the West Coast. While these invasive eastern species are present on campuses on the West Coast, notably, contingency analysis did not provide evidence that they are the primary factor discouraging western gray squirrels (Sciurus griseus) from using these habitats. The inventory and analyses presented here can provide a basis for longitudinal studies both within and across campuses and may be productively combined with initiatives that involve students in research.

• Perlut, N. (2020) Long-Distance Dispersal by Eastern Gray Squirrels in Suburban Habitats. Northeastern Naturalist 27: 195.

Abstract: Natal dispersal by Sciurus carolinensis (Eastern Gray Squirrel) is poorly un-derstood, given so rarely reported, yet dispersal patterns in small mammals can affect seed dispersal and predation, as well as population dynamics of predators. Herein, I document long-distance dispersal by 3 Eastern Gray Squirrels from the suburban coastal campus of the University of New England in Biddeford, ME. Mean dispersal distance was 10.1 km (min–max = 6.3–14.5 km), occurring in random directions (SW, S, NW). These results, combined with the previous studies, better describe the distribution of natal dispersal by Eastern Gray Squirrel—critical information in understanding population processes and potentially developing effective landscape-management strategies.

• Pm, H., O’Brien, D., Yahr, Neaves, Ahrends, Kt, B., Brooker, R., Burke, Cavers, S., Dawson, Da, E., Kerr, Df, M., Rj, P., Trivedi, Wall, Wright, Ogden, R. & Ennos, R. (2020) Scotland’s Biodiversity Progress to 2020 Aichi Targets Conserving Genetic Diversity -Development of a national approach for addressing Aichi Biodiversity Target 13 that includes wild species Contributors.

Abstract: Aichi Target 13 (T13) focuses on the conservation of genetic diversity. • Major challenges in implementing T13 are that the type of genetic diversity to conserve is not clearly defined, and that key issues in genetic conservation vary across different sectors (e.g., forestry vs agriculture vs other species of socio-economic importance). • In Scotland and the UK more widely, baseline mechanisms are well established for assessing and reporting on genetic diversity in species of agricultural importance (e.g., rare livestock breeds, crop wild relatives), and a methodology has been established for ornamental plants. • A new UK Strategy for Forest Genetics Resources was launched in 2019, creating a framework for linking forest trees into T13 reporting. • However, there is no clear strategy to deal with ‘other species of socio-economic importance’ in Scotland, the UK or indeed elsewhere, and addressing this gap is the major focus of this report. • There is a lack of guidance for identifying focal species of socio-economic importance, and no clear mechanism for addressing T13 for these species once they have been identified. • To address this, we have identified a set of criteria for defining terrestrial and freshwater species of socio-economic importance in Scotland, and selected an initial list of 26 species.

• Sainsbury, A. W., Chantrey, J., Ewen, J. G., Gurnell, J., Hudson, P., Karesh, W. B., Kock, R. A., Lurz, P. W. W., Meredith, A. & Tompkins, D. M. (2020) Implications of squirrelpox virus for successful red squirrel translocations within mainland UK. Conservation Science and Practice 2(6): e200.

Abstract: Remnant red squirrel populations in the UK mainland are threatened by squirrelpox viral disease and the reservoir of the squirrelpox virus, the invasive grey squirrel, is expanding its range. Until this threat can be effectively mitigated, there is a high risk from disease outbreaks, following proposed conservation translocation of red squirrels.

• Santicchia, F., Wauters, L., Piscitelli, A., Dongen, S., Martinoli, A., Preatoni, D., Romeo, C. & Ferrari, N. (2020) Spillover of an alien parasite reduces expression of costly behaviour in native host species. Journal of Animal Ecology.

Abstract: Understanding the effects of invasive alien species (IAS) on native host–parasite relationships is of importance for enhancing ecological theory and IAS management. When IAS and their parasite(s) invade a guild, the effects of interspecific resource competition and/or parasite‐mediated competition can alter existing native host–parasite relationships and the dependent biological traits such as native species’ behaviour. We used a natural experiment of populations of native red squirrels Sciurus vulgaris that were colonized by the alien grey squirrel Sciurus carolinensis, comparing repeated measurements of red squirrel parasite infection and personality with those taken in sites where only the native species occurred. We explored two alternative hypotheses: (a) individual differences in personality traits (activity and/or sociability) of native red squirrel positively affect the probability of macroparasite spillover and thus the likelihood to acquire the alien’s parasitic helminth Strongyloides robustus; (b) the combined effects of grey squirrel presence and parasite infection result in a reduction of costly personality traits (activity and/or exploration). Using data from 323 arena tests across three experimental (native species and IAS) and three control sites (only native species), we found negative correlations between native species’ activity and infection with S. robustus in the sites invaded by the alien species. Activity was also negatively correlated with infection by its native helminth Trypanoxyuris sciuri but only when grey squirrels were present, while in the red‐only sites there was no relationship of T. sciuri infection with any of the personality traits. Moreover, individuals that acquired S. robustus during the study reduced their activity after infection, while this was not the case for animals that remained uninfected. Our results show that parasite‐mediated competition is costly, reducing activity in individuals of the native species, and altering the native host–native parasite relationships. The authors show how an alien parasite reduces the expression of activity, an energetically costly personality trait. The results also suggest that transmission of alien parasites may alter the native host–native parasite relationship: only in areas invaded by alien grey squirrels and their macroparasites, infection by the native oxiurid helminth reduced native hosts’ activity.

• Schilling, A.-K., Lurz, P. & Meredith, A. (2020) Field anaesthesia of Eurasian red squirrels (Sciurus vulgaris). Veterinary Anaesthesia and Analgesia.

 

• Schulze, V., Lurz, P., Ferrari, N., Romeo, C., Steele, M., Marino, S., Mazzamuto, M., Calvignac-Spencer, S., Schlottau, K., Beer, M., Ulrich, R. & Ehlers, B. (2020) Search for polyoma-, herpes-, and bornaviruses in squirrels of the family Sciuridae. Virology Journal 17.

Abstract: Background Squirrels (family Sciuridae) are globally distributed members of the order Rodentia with wildlife occurrence in indigenous and non-indigenous regions (as invasive species) and frequent presence in zoological gardens and other holdings. Multiple species introductions, strong inter-species competition as well as the recent discovery of a novel zoonotic bornavirus resulted in increased research interest on squirrel pathogens. Therefore we aimed to test a variety of squirrel species for representatives of three virus families. Methods Several species of the squirrel subfamilies Sciurinae, Callosciurinae and Xerinae were tested for the presence of polyomaviruses (PyVs; family Polyomaviridae) and herpesviruses (HVs; family Herpesviridae), using generic nested polymerase chain reaction (PCR) with specificity for the PyV VP1 gene and the HV DNA polymerase (DPOL) gene, respectively. Selected animals were tested for the presence of bornaviruses (family Bornaviridae), using both a broad-range orthobornavirus- and a variegated squirrel bornavirus 1 (VSBV-1)-specific reverse transcription-quantitative PCR (RT-qPCR). Results In addition to previously detected bornavirus RNA-positive squirrels no more animals tested positive in this study, but four novel PyVs, four novel betaherpesviruses (BHVs) and six novel gammaherpesviruses (GHVs) were identified. For three PyVs, complete genomes could be amplified with long-distance PCR (LD-PCR). Splice sites of the PyV genomes were predicted in silico for large T antigen, small T antigen, and VP2 coding sequences, and experimentally confirmed in Vero and NIH/3T3 cells. Attempts to extend the HV DPOL sequences in upstream direction resulted in contiguous sequences of around 3.3 kilobase pairs for one BHV and two GHVs. Phylogenetic analysis allocated the novel squirrel PyVs to the genera Alpha- and Betapolyomavirus, the BHVs to the genus Muromegalovirus, and the GHVs to the genera Rhadinovirus and Macavirus. Conclusions This is the first report on molecular identification and sequence characterization of PyVs and HVs and the detection of bornavirus coinfections with PyVs or HVs in two squirrel species. Multiple detection of PyVs and HVs in certain squirrel species exclusively indicate their potential host association to a single squirrel species. The novel PyVs and HVs might serve for a better understanding of virus evolution in invading host species in the future.

• Shuttleworth, C. M., Brady, D., Cross, P., Gardner, L., Greenwood, A., Jackson, N., McKinney, C., Robinson, N., Trotter, S., Valle, S., Wood, K. & Hayward, M. W. (2020) Recalibrating risk: Implications of squirrelpox virus for successful red squirrel translocations within mainland UK. Conservation Science and Practice n/a(n/a): e321.

 

• Steele, M. A. & Yi, X. (2020) Squirrel-Seed Interactions: The Evolutionary Strategies and Impact of Squirrels as Both Seed Predators and Seed Dispersers. Frontiers in Ecology and Evolution 8(259).

For many squirrel species, their intense – arguably coevolutionary – interactions with seed and nut producing trees have significantly shaped their biology and diversity. Here we provide an overview of this relationship in a range of forest types worldwide. We first review the evidence for how forest composition (conifer, hardwood, mixed hardwood and overall diversity of tree species) influences interactions between squirrels and seed trees and, ultimately, the role of squirrels as either seed predators or seed dispersers. We review, for example, the intense selective pressure squirrels exert on conifer trees as seed predators and the diversity of morphological traits and behavioral strategies that allow them to efficiently exploit this critical resource. In contrast, we show how the squirrel’s role shifts to one of seed disperser in hardwood forests and how the specifics of this interaction varies further with forest structure, forest composition and climatic conditions. We then review the growing evidence for the tight ecological and evolutionary dance between the squirrels and the oaks that has shaped the biology of both across the globe. We show how a suite of seed (acorn) characteristics (e.g., chemical gradients, germination schedules, seedling morphology and tolerance-resistance strategies) are all intimately tied to the scatter-hoarding decisions of several squirrel species. And, based on studies in oak forests in Central America, Mexico, North America, and Eurasia, we also highlight the behavioral strategy of embryo excision now reported for six species across at least four genera of squirrels. This behavior, glaringly absent in other scatter-hoarding rodents worldwide, is now known be an innate trait in at least two species, one in Asia and another in North America. We review extensive recent research on one species of squirrel, the Siberian chipmunk (Tamias sibiricus), which exhibits a suite of behavioral strategies unique to that of other squirrels that independently contributes to seed dispersal and establishment. Finally, we outline numerous remaining questions concerning plants and other taxa of squirrels still open to investigation.

• Studd, E. K., Menzies, A. K., Siracusa, E. R., Dantzer, B., Lane, J. E., McAdam, A. G., Boutin, S. & Humphries, M. M. (2020) Optimisation of energetic and reproductive gains explains behavioural responses to environmental variation across seasons and years. Ecology Letters 23(5): 841-850.

Abstract Animals switch between inactive and active states, simultaneously impacting their energy intake, energy expenditure and predation risk, and collectively defining how they engage with environmental variation and trophic interactions. We assess daily activity responses to long-term variation in temperature, resources and mating opportunities to examine whether individuals choose to be active or inactive according to an optimisation of the relative energetic and reproductive gains each state offers. We show that this simplified behavioural decision approach predicts most activity variation (R2 = 0.83) expressed by free-ranging red squirrels over 4 years, as quantified through accelerometer recordings (489 deployments; 5066 squirrel-days). Recognising activity as a determinant of energetic status, the predictability of activity variation aggregated at a daily scale, and the clear signal that behaviour is environmentally forced through optimisation of gain, provides an integrated approach to examine behavioural variation as an intermediary between environmental variation and energetic, life-history and ecological outcomes.

• Thévenin, C., Morin, A., Kerbiriou, C., Sarrazin, F. & Robert, A. (2020) Heterogeneity in the allocation of reintroduction efforts among terrestrial mammals in Europe. Biological Conservation 241: 108346.

Abstract: Reintroductions offer a powerful tool to reverse adverse anthropogenic impacts on biodiversity by restoring extirpated populations within the indigenous range of species. Reintroductions have become popular and have been increasingly used over the last decades. However, this species-centred conservation approach has been criticized for being taxonomically biased and for focusing on large and charismatic species. Studies investigating taxonomic biases in the allocation of reintroduction efforts at large scale generally consider taxonomic bias within and among higher taxa (e.g. vertebrates, plants), by comparing the number of reintroduced species within a taxon to its prevalence in nature. Here, we show that the bias is even more striking when accounting for the differences in the number of implemented programs among reintroduced species. We conducted a comprehensive search of the peer-reviewed and grey literature to inventory reintroduction programs of European terrestrial mammals. We identified 28 species that have been reintroduced at least one time. For each reintroduced mammal, we extensively searched two literature search engines and found 414 relevant publications, which described 375 distinguishable reintroduction programs implemented in Europe from the early 20th century to 2013. We used the number of implemented programs and the number of associated publications to investigate the distribution of reintroduction efforts among species. Our results show a substantial heterogeneity in the allocation of reintroduction efforts, with 68% of implemented reintroductions in Europe involving only three species: the beaver (Castor fiber), the Alpine ibex (Capra ibex) and the European bison (Bison bonasus).

• Turkia, T., Jousimo, J., Tiainen, J., Helle, P., Rintala, J., Hokkanen, T., Valkama, J. & Selonen, V. (2020) Large-scale spatial synchrony in red squirrel populations driven by a bottom-up effect. Oecologia.

Abstract: Spatial synchrony between populations emerges from endogenous and exogenous processes, such as intra- and interspecific interactions and abiotic factors. Understanding factors contributing to synchronous population dynamics help to better understand what determines abundance of a species. This study focuses on spatial and temporal dynamics in the Eurasian red squirrel (Sciurus vulgaris) using snow-track data from Finland from 29 years. We disentangled the effects of bottom-up and top-down forces as well as environmental factors on population dynamics with a spatiotemporally explicit Bayesian hierarchical approach. We found red squirrel abundance to be positively associated with both the abundance of Norway spruce (Picea abies) cones and the predators, the pine marten (Martes martes) and the northern goshawk (Accipiter gentilis), probably due to shared habitat preferences. The results suggest that red squirrel populations are synchronized over remarkably large distances, on a scale of hundreds of kilometres, and that this synchrony is mainly driven by similarly spatially autocorrelated spruce cone crop. Our research demonstrates how a bottom-up effect can drive spatial synchrony in consumer populations on a very large scale of hundreds of kilometres, and also how an explicit spatiotemporal approach can improve model performance for fluctuating populations.

• Twining, J. P., Montgomery, W. I., Price, L., Kunc, H. P. & Tosh, D. G. (2020) Native and invasive squirrels show different behavioural responses to scent of a shared native predator. Royal Society Open Science 7: 191841.

Abstract: Invasive species pose a serious threat to native species. In Europe, invasive grey squirrels (Sciurus carolinensis) have replaced native red squirrels (Sciurus vulgaris) in locations across Britain, Ireland and Italy. The European pine marten (Martes martes) can reverse the replacement of red squirrels by grey squirrels, but the underlying mechanism of how pine martens suppress grey squirrels is little understood. Research suggests the reversal process is driven by direct predation, but why the native red squirrel may be less susceptible than the invasive grey squirrel to predation by a commonly shared native predator, is unknown. A behavioural difference may exist with the native sciurid being more effective at avoiding predation by the pine marten with which they have a shared evolutionary history. In mammals, olfactory cues are used by prey species to avoid predators. To test whether anti-predator responses differ between the native red squirrel and the invasive grey squirrel, we exposed both species to scent cues of a shared native predator and quantified the responses of the two squirrel species. Red squirrels responded to pine marten scent by avoiding the feeder, increasing their vigilance and decreasing their feeding activity. By contrast, grey squirrels did not show any anti-predator behaviours in response to the scent of pine marten. Thus, differences in behavioural responses to a shared native predator may assist in explaining differing outcomes of species interactions between native and invasive prey species depending on the presence, abundance and exposure to native predators.

• Twining, J. P., Montgomery, W. I. & Tosh, D. G. (2020) Declining invasive grey squirrel populations may persist in refugia as native predator recovery reverses squirrel species replacement. Journal of Applied Ecology n/a(n/a).

Abstract: Invasive species pose one of the most serious global threats to biodiversity. Investigations into the interactions of native and non-native species focus on the impacts of single species, despite being embedded in a network of direct and indirect interactions between multiple species and their environments. We developed 1-km2 resolution, single-species and multi-species occupancy models using quantitative camera trap data collected by citizen scientists at 332 sites in a regional survey comprising the 14,130 km2 of Northern Ireland. Recent research suggests that native red and invasive grey squirrels in Britain and Ireland are linked by resource and disease-mediated competition, and by a shared enemy, the European pine marten. We demonstrate that the presence of the pine marten reverses red squirrel replacement by grey squirrels on a regional basis, with red squirrel occupancy positively affected by exposure to pine marten. In contrast, the grey squirrel has a strongly negative response to the presence of pine marten. Modelling habitat suitability of pine marten and both squirrel species suggests that, despite the potentially strong effect of a recovering population of pine martens in controlling grey squirrel distribution, the latter is likely to persist in urban refugia that are either inaccessible or avoided by the pine marten. Synthesis and applications. Sustainable recovery of both the red squirrel and the pine marten across the wider landscapes of Ireland, Scotland and Northern England seems probable, while in Southern England and Wales, although possible, chances are more remote. Increasing native woodland cover on a landscape scale to facilitate pine marten recovery may assist in realizing this scenario. Despite the ongoing recovery of the pine marten, and resultant declines in grey squirrels, isolated populations are likely to persist in urban areas. Human-lead control of grey squirrel populations in urban refugia requires urgent funding and implementation to avoid the development of novel genotypes in grey squirrel source populations that enable the species to better avoid predation by the pine marten.

• Wang, B. & Yang, X. (2020) Scatter-hoarding rodent foraging preference on nutrient content is mediated by seed size. Current Zoology 66(4): 445-446.

ABstract: Seed dispersal and predation by scatter-hoarding rodents play an important role in seedling regeneration, community structure and composition, and species coexistence (Vander Wall 2010; Zhang et al. 2016). Rodents generally exhibit seed species preference during their scatter-hoarding behavioral processes (Xiao et al. 2006; Gong et al. 2015; Zhang et al. 2016), and seed traits are a key factor in explaining rodent foraging preference (Wang and Chen 2009; Vander Wall 2010; Zhang et al. 2016). Numerous studies have explored how the diversity of seed traits influences rodent foraging behavior but have shown inconsistent results (Blate et al. 1998; Moles et al. 2003; Xiao et al. 2006; Wang and Chen 2009). These disparate results are likely because of the complicated interactions among traits (Gong et al. 2015; Zhang et al. 2016). In other words, the effect of a given seed trait on rodent foraging behavior may be influenced by other seed traits. However, few studies have directly targeted the interactive effects among seed traits, possibly because of the lack of nondestructive methods for measuring many seed traits, which makes it difficult to compare among individual seeds. Theoretically, multispecies studies provide a potential way to test the interactive effects of different seed traits on rodent foraging preference, by using the mean values of seed traits for each species. However, until now, few studies have included enough species to perform a statistical analysis of the interactive effects among traits (but see Gong et al. 2015; Zhang et al. 2016). Furthermore, large intraspecific variation often exists in many seed traits (Wang and Ives 2017), which may bias the results of interspecific comparisons by using mean values. Artificial seed experimental systems provide an opportunity to ascertain the joint effects of several seed traits by precisely controlling each target trait individually (Wang and Chen 2009, 2012; Wang and Yang 2014). However, all these studies only pointed out the potential interactive effects among traits, and did not analyze how one seed trait would influence the magnitude or intensity of the effect of another trait on rodent foraging preference. In this study, we used artificial seeds to target two important seed traits—seed size and nutrient content—to further explore how one seed trait influences the effect magnitude of another trait on the foraging preference of scatter-hoarding rodents (see Supplementary Material).

• Werner, C. S. & Nunn, C. L. (2020) Effect of urban habitat use on parasitism in mammals: a meta-analysis. Proceedings of the Royal Society B: Biological Sciences 287(1927): 20200397.

Abstract: Rates of urbanization are increasing globally, with consequences for the dynamics of parasites and their wildlife hosts. A small subset of mammal species have the dietary and behavioural flexibility to survive in urban settings. The changes that characterize urban ecology—including landscape transformation, modified diets and shifts in community composition—can either increase or decrease susceptibility and exposure to parasites. We used a meta-analytic approach to systematically assess differences in endoparasitism between mammals in urban and non-urban habitats. Parasite prevalence estimates in matched urban and non-urban mammal populations from 33 species were compiled from 46 published studies, and an overall effect of urban habitation on parasitism was derived after controlling for study and parasite genus. Parasite life cycle type and host order were investigated as moderators of the effect sizes. We found that parasites with complex life cycles were less prevalent in urban carnivore and primate populations than in non-urban populations. However, we found no difference in urban and non-urban prevalence for parasites in rodent and marsupial hosts, or differences in prevalence for parasites with simple life cycles in any host taxa. Our findings therefore suggest the disruption of some parasite transmission cycles in the urban ecological community.

• Zwolak, R. & Sih, A. (2020) Animal personalities and seed dispersal: A conceptual review. Functional Ecology n/a(n/a).

Abstract: Consistent individual tendencies in behaviour, or behavioural types, are likely to impact the dynamics and outcomes of animal-mediated seed dispersal. We review the extant literature on this issue and outline a conceptual overview to guide this emerging field. We provide an overview of possible ways in which behavioural types can affect animal-mediated seed dispersal. We summarize theoretical mechanisms linking behavioural types with seed dispersal outcomes and review how behavioural types might affect each stage of seed dispersal, beginning with fruit encounter and harvest, and ending with events that take place after seed deposition. Since behavioural types involve correlations among different behaviours (i.e. behavioural syndromes), they can generate unexpected associations between different decisions that are involved in seed dispersal, with conflicting (or reinforcing) effects on different stages of seed dispersal. Thus, we draw particular attention to trade-offs faced by seeds dispersed by individuals with different behavioural tendencies. We also note that since seed dispersal is a multiplicative process with different stages, disperser behavioural types that provide moderately efficient dispersal at each stage will be better for plants than behavioural types that are very efficient at some steps, but inefficient on others. Finally, we provide testable predictions on the links between behavioural types and characteristics of seed dispersal, including, for example, influences on the probability of seed harvest, dispersal distance, deposition sites and condition of dispersed seeds. We argue that investigating the links between behavioural types and animal-mediated seed dispersal will provide a better mechanistic understanding of seed dispersal and plant regeneration. A free plain language summary can be found within the Supporting Information of this article.

2019

• Amspacher, K., Bauer, B., Waldron, J., Wiggers, E. & Welch, S. (2019) Sciurus niger niger (Southern Fox Squirrel) Density and the Diurnal Patterns, Occupancy, and Detection of Sympatric Southern Fox Squirrels and S. carolinensis (Eastern Gray Squirrel) on Spring Island, South Carolina.

Abstract: Sciurus niger niger (Southern Fox Squirrel, hereafter, SFS) are habitat specialists within the Pinus palustris (Longleaf Pine) ecosystem of the southeastern US whose populations are declining due to habitat loss and fragmentation. Sciurus carolinensis (Eastern Gray Squirrel, hereafter, EGS) are generalists found throughout the eastern US that have historically avoided competition with SFSs through habitat-partitioning. Spring Island, SC, once reported the densest population of SFSs in the southeast, but residents of the island community recently reported decreasing SFS populations and increasing EGS populations. We used baited game-cameras at stratified random points to estimate SFS density, observe patterns of SFS and EGS diurnal activity, and examine the influence of habitat structure and management techniques (e.g., prescribed fire) on SFS and EGS occurrence on Spring Island. Our estimate of SFS density on Spring Island (0.28-0.97 SFSs/ha) was similar to the high density previously reported (0.758 SFSs/ha). SFSs and EGSs had similar diurnal patterns, but SFSs were most active around midday while EGSs were most active in early morning and evening hours. EGS occupancy was negatively associated with fire frequency. EGS detection probability was negatively associated with maximum daily temperature. These data suggest that habitat management via prescribed fire has allowed stable populations of both SFSs and EGSs to persist despite anthropogenic land changes.

• Amuno, S., Shekh, K., Kodzhahinchev, V. & Niyogi, S. (2019) Neuropathological changes in wild muskrats (Ondatra zibethicus) and red squirrels (Tamiasciurus hudsonicus) breeding in arsenic endemic areas of Yellowknife, Northwest Territories (Canada): Arsenic and cadmium accumulation in the brain and biomarkers of oxidative stress. Science of The Total Environment: 135426.

Abstract: The brain is one of the critical organs particularly susceptible to the damaging effects of chronic arsenic poisoning and there is a growing body of evidence that suggest that oxidative stress plays a key role in the pathogenesis of neurodegenerative disorders. The aim of this present work was to comparatively assess biomarkers of oxidative stress and status of antioxidant enzyme activities in the brains of muskrats and squirrels breeding in arsenic endemic areas, specifically near the vicinity of the abandoned Giant mine site (~2 km radius), and an intermediate location approximately 20 km from the mine area and in reference locations spanning 52–105 km from the city of Yellowknife, Northwest Territories (Canada). Analysis included measurement of total arsenic and cadmium concentration in the nails, brain, and stomach content of muskrats and squirrels, in addition to biochemical evaluation of lipid peroxidation levels and antioxidant enzymes defense: catalase (CAT), superoxide dismutase (SOD) and glutathione peroxidase (GPx) in the brain tissues. The results revealed that arsenic concentration in the nails of muskrats collected closest to the vicinity of the mine area was in the range of 11 to 35.1 times higher than those from the reference site. The maximum concentration of arsenic in the nails of muskrats from the intermediate location was 47.6 times higher than the maximum concentration observed in the reference muskrats. Cadmium was generally undetected in the nails of muskrats and squirrels from the three sampling locations. Arsenic in the gut contents of muskrats from the arsenic affected area was 4.5 to 49.1 times higher than those from the reference site. Cadmium levels in the guts of muskrats from the mine area almost doubled those from the reference site. Arsenic accumulated in the nails of squirrels from the areas closest to the mine but was undetected in the squirrel nails from the reference location. The maximum arsenic levels in the stomach content of squirrels from the mine area was ~40 times higher than those from the reference site. Arsenic did not accumulate in the brains of muskrats, but cadmium was detected in a few brains of muskrats. Brains of squirrels from the mine area and intermediate locations accumulated both arsenic and cadmium. The brains of squirrels and muskrats from the arsenic affected area showed no evidence of increased lipid peroxidation compared to the animals from the reference site. However, SOD, CAT and GPx activities in the brains of animals from the arsenic endemic areas tended to be higher compared to the control sites. This is the first study documenting evidence of oxidative stress and altered antioxidant enzyme activities in brains of wild rodent population in arsenic endemic areas of Canada.

• Asari, Y. & Yanagawa, H. (2019) Home Range and Nest Arrangement of the Siberian Flying Squirrel Pteromys volans in an Urban Edge, Northern Japan. Polish Journal of Ecology 67(2): 159-167.

Forest fragmentation by urbanizing and developing roads and fields is common in Japan; thus, the Siberian flying squirrel (Pteromys volans) population may decrease in the future. This study examined the effects of urbanization on P. volans by clarifying their home ranges and nest arrangements. Ten flying squirrels were captured and radio-telemetrically tracked from 2005 to 2007 in Obihiro City in eastern Hokkaido (Northern Japan). Males had significantly larger 100% MCP (minimum convex polygon) areas (mean ± SD; 6.5 ± 5.1 ha) and 95% MCP areas (3.6 ± 1.3 ha) than females (1.8 ± 1.0 ha and 1.0 ± 0.5 ha, respectively). Also the core area of males (1.2 ± 1.1 ha) was larger than females (0.2 ± 0.2 ha), though this difference was not significant. Males and females used more nests located inside than outside of their core areas. Distance from the core area edge to the nest located outside the core was longer for males (78.6 ± 139.9 m) than for females (39.7 ± 39.9 m), but this difference was not significant. Siberian flying squirrels in urban forests likely show similar ecology with flying squirrels in consecutive forests; however, they might change their behavior to share resources with others because urban forests restrict nest and food resources.

• Bertrand, O. C., San Martin-Flores, G. & Silcox, M. T. (2019) Endocranial shape variation in the squirrel-related clade and their fossil relatives using 3D geometric morphometrics: contributions of locomotion and phylogeny to brain shape. Journal of Zoology 0(0).

Abstract: Landmark-based 3D geometric morphometrics has rarely been employed to understand the relationship between endocranial shape, phylogeny and ecology. The goal of this study is to examine the endocranial morphology of members of the squirrel-related clade by using these methods, and to develop a multi-faceted framework for studying brain evolution applicable to other groups. The squirrel-related clade is taxonomically and ecologically diverse, and includes tree squirrels, the mountain beaver and dormice. Virtual endocasts for Ischyromyidae, a primitive group of rodents likely to be related to the squirrel-related clade, were also included. Thirty landmarks were taken on virtual endocasts derived from 32 extant and extinct species. The results show that endocranial shape and size are significantly correlated in that smaller endocasts are relatively wider laterally than larger endocasts. The principal components analysis (PCA) reveals that endocranial shape is clearly distinct for Sciuridae, Aplodontidae, Gliridae and Ischyromyidae. Endocranial shape variation is associated with changes in the development of the neocortex, cerebellum (including the paraflocculi) and olfactory bulbs. The Kmult test shows that endocranial shape reflects phylogenetic relationships among the four families and within Sciuridae. In the PCA analysis, flying squirrels show the most distinct endocranial morphology among squirrels, overlapping the least with other tribes and subfamilies. This result suggests that gliding may have imposed specific constraints on cranial shape. The endocasts of fossil and modern fossorial Aplodontidae have a shape similar to those of Ischyromyidae. This similarity could be the result of homoplasy related to fossorial specialization in later occurring Aplodontidae. The fossil Sciurini Protosciurus is outside the range of variation for modern squirrels, suggesting that the emergence of the modern squirrel endocranial bauplan may have not been established until after the early Miocene. From the data gathered, phylogeny and locomotion both impacted endocranial shape in our rodent sample.

Bono, J. M., Pigage, H. K., Wettstein, P. J., Prosser, S. A. & Pigage, J. C. (2018) Genome-wide markers reveal a complex evolutionary history involving divergence and introgression in the Abert’s squirrel (Sciurus aberti) species group. BMC Evolutionary Biology 18(1): 139.

Abstract: Background Genetic introgression between divergent lineages is now considered more common than previously appreciated, with potentially important consequences for adaptation and speciation. Introgression is often asymmetric between populations and patterns can vary for different types of loci (nuclear vs. organellar), complicating phylogeographic reconstruction. The taxonomy of the ecologically specialized Abert’s squirrel species group has been controversial, and previous studies based on mitochondrial data have not fully resolved the evolutionary relationships among populations. Moreover, while these studies identified potential areas of secondary contact between divergent lineages, the possibility for introgression has not been tested. Results We used RAD-seq to unravel the complex evolutionary history of the Abert’s squirrel species group. Although some of our findings reinforce inferences based on mitochondrial data, we also find significant areas of discordance. Discordant signals generally arise from previously undetected introgression between divergent populations that differentially affected variation at mitochondrial and nuclear loci. Most notably, our results support earlier claims (disputed by mitochondrial data) that S. aberti kaibabensis, found only on the north rim of the Grand Canyon, is highly divergent from other populations. However, we also detected introgression of S. aberti kaibabensis DNA into other S. aberti populations, which likely accounts for the previously inferred close genetic relationship between this population and those south of the Grand Canyon. Conclusions Overall, the evolutionary history of Abert’s squirrels appears to be shaped largely by divergence during periods of habitat isolation. However, we also found evidence for interbreeding during periods of secondary contact resulting in introgression, with variable effects on mitochondrial and nuclear markers. Our results support the emerging view that populations often diversify under scenarios involving both divergence in isolation and gene flow during secondary contact, and highlight the value of genome-wide datasets for resolving such complex evolutionary histories.

• Broughton, R. K. (2019) Current and future impacts of nest predation and nest-site competition by invasive eastern grey squirrels Sciurus carolinensis on European birds. Mammal Review 50(1): 35-81.

Abstract The eastern grey squirrel (hereafter ?grey squirrel?) is considered one of the most damaging invasive alien species in Europe, with negative effects on native ecosystems. Despite it being widely perceived as a significant predator of bird eggs and chicks and as a competitor for nest sites, evaluation of the grey squirrel?s impact on European bird populations has been hindered by limited empirical data. The aim was to review the incidence of grey squirrels as nest predators of and nest-site competitors with European birds, and to use this information to identify species at potential risk of negative effects from within the grey squirrel?s expanding range in continental Europe. A comprehensive literature review was conducted, and the data were used alongside additional new data, to assess nest predation and competition by grey squirrels in their current European range. Bird species were grouped by nest-site type, which was used to predict the impact on similar species groups in regions of continental Europe predicted to be colonised by grey squirrels in the current century. Camera-monitoring and field evidence for 12 bird species and 12420 nests in Britain showed that grey squirrels rarely depredated eggs or chicks, affecting just 0.5% of nests. Nest-site competition was also minor, with grey squirrels occupying 0.8% of 122 small tree cavities and 14% of 57 larger cavities. At least 69 bird species in continental Europe could be exposed to potential nest predation or competition from expanding grey squirrel populations within the current century, but population-level effects currently appear to be unlikely. Current evidence shows that grey squirrels are unlikely to be significant predators of or competitors with nesting birds in their present or projected range in Europe. However, further studies of more species in different regions would be valuable, particularly in urban and suburban habitats.

• Chow, P. K. Y., Lea, S., Hempel de Ibarra, N. & Robert, T. (2019) Inhibitory control and memory in the search process for a modified problem in grey squirrels, Sciurus carolinensis. Animal Cognition 22.

Abstract: Inhibiting learned behaviours when they become unproductive and searching for an alternative solution to solve a familiar but different problem are two indicators of flexibility in problem solving. A wide range of animals show these tendencies spontaneously, but what kind of search process is at play behind their problem-solving success? Here, we investigated how Eastern grey squirrels, Sciurus carolinensis, solved a modified mechanical problem that required them to abandon their preferred and learned solution and search for alternative solutions to retrieve out-of-reach food rewards. Squirrels could solve the problem by engaging in either an exhaustive search (i.e., using trial-and-error to access the reward) or a ‘backup’ solution search (i.e., recalling a previously successful but non-preferred solution). We found that all squirrels successfully solved the modified problem on their first trial and showed solving durations comparable to their last experience of using their preferred solution. Their success and high efficiency could be explained by their high level of inhibitory control as the squirrels did not persistently emit the learned and preferred, but now ineffective, pushing behaviour. Although the squirrels had minimal experience in using the alternative (non-preferred) successful solution, they used it directly or after one or two failed attempts to achieve success. Thus, the squirrels were using the ‘backup’ solution search process. Such a process is likely a form of generalisation which involves retrieving related information of an experienced problem and applying previous successful experience during problem solving. Overall, our results provide information regarding the search process underlying the flexibility observable in problem-solving success.

• Creley, C., Muchlinski, A. & Shilling, F. (2019) An Ecological Niche Model to Predict Range Expansion of the Eastern Gray Squirrel in California. Bulletin of the Southern California Academy of Sciences (1): 58-70.

Abstract: The eastern gray squirrel, Sciurus carolinensis (EGS) has been introduced to California and has expanded its geographic range since initial introductions. In this study we projected the potential future geographic range of the EGS in California using Maxent to create an ecological niche model. Location data were obtained over the time period of 2004–2015 from museum specimens, wildlife rehabilitation centers, the California Department of Public Health, the California Roadkill Observation System, and non-iNaturalist citizen science observations. Research grade data from iNaturalist was obtained over the time period of 2004–2018. Range and habitat suitability maps were developed by mapping in ArcGIS. Three threshold selection methods were used to create different estimates of the potential future range of the EGS in California. The first method used the 10th percentile logistic threshold, the second used the minimum training presence logistic threshold, and the third used Jenks Natural Breaks. We propose that Jenks Natural Breaks has distinct advantages over the other two methods for estimating the potential future range of the introduced EGS in California, because it provides information on the habitat suitability ranking throughout California, whereas the other methods only provide a binary suitable/unsuitable map.

• Curay, J., Romero, V. & Brito, J. (2019) Small non-volant mammals of the Reserva Geobotánica Pululahua, Ecuador. Mammalia 83(6): 574.

We provide insights into the diversity of small non-volant mammals (SNVMs) in the Reserva Geobotánica Pululahua, a peri-urban natural area of Quito, Pichincha province, Ecuador. Based on Sherman-like traps and pitfall traps, we recorded 21 species of SNVMs. They represent 40% of the total species reported for the entire province, and include six species that are endemic to the northwestern slope of the Ecuadorian Andes. Despite its proximity to the city of Quito (less than 5 km), Pululahua seems to have an average richness compared with other similar localities in the Ecuadorian Andes. Our results are a baseline for future conservation plans for SNVMs.

• Derbridge, J. J. & Koprowski, J. L. (2019) Experimental removals reveal dietary niche partitioning facilitates coexistence between native and introduced species. Ecology and Evolution 0(0).

Abstract: Niche overlap between native species and ecologically similar invaders can lead to competitive exclusion of threatened native species, but if two such species also co-occur naturally elsewhere, interactions between native and introduced populations may mirror coevolved niche partitioning that reduces competition and promotes coexistence. A single, insular population of Fremont?s squirrel (Tamiasciurus fremonti) the Mount Graham red squirrel (MGRS; T. f. grahamensis) in the Pinaleño Mountains, Arizona, USA, is critically endangered and resource competition with introduced Abert’s squirrels (Sciurus aberti) may threaten its long-term persistence. The species are naturally synoptic in other mountain sites, and both consume diets comprised primarily of conifer seeds and fungi. We conducted experimental removals of introduced Abert’s squirrels and used stable isotope analysis of diets before and after removals, and of diets in naturally syntopic populations to test the hypothesis that dietary niche partitioning can facilitate coexistence between native and introduced species. We also developed a novel approach to determine the influence of fluctuating food availability on carbon enrichment in consumers. Mount Graham red squirrels and introduced Abert’s squirrels partitioned the dietary niche similarly to naturally syntopic populations. Removals had no apparent effect. Diet of MGRS was more closely linked to availability of resources than to presence of Abert’s squirrels. Flexible dietary niche of introduced Abert’s squirrels may have allowed them to exploit a resource opportunity in syntopy with MGRS. Variable food production of MGRS habitat may intensify competition in poor years, and territorial defense against non-native Abert’s squirrels likely imposes fitness costs on individual MGRS. Similarity in our model species? diets may make MGRS more vulnerable to competition if climate change eliminates the advantages of larder-hoarding. Where introduced populations of ecologically similar species are better adapted to changing conditions, they may ultimately replace southern peripheral populations of native species.

• Di Febbraro, M., Menchetti, M., Russo, D., Ancillotto, L., Aloise, G., Roscioni, F., Preatoni, D. G., Loy, A., Martinoli, A., Bertolino, S. & Mori, E. (2019) Integrating climate and land-use change scenarios in modelling the future spread of invasive squirrels in Italy. Diversity and Distributions 0(0).

Abstract: Aim The establishment and spread of invasive alien species may be influenced by several mutually interacting factors, whose understanding is paramount to develop effective biosecurity policies. However, studies focused on modelling spatially explicit patterns of future invasion risk have so far focused on species response to climate change impacts, while land-use change has been neglected. We investigated how the interplay between climate and land-use change could affect the future potential distribution and dispersal corridors of four alien squirrels introduced to Europe (Sciurus carolinensis, Callosciurus finlaysonii, Callosciurus erythraeus and Eutamias sibiricus). Location and Methods Our study was conducted in Italy. We used Species Distribution Models and circuit theory methods to test whether future scenarios based only on climate change predict a different effect on range and connectivity of alien squirrel populations, compared to scenarios that include both climate and land-use changes. Results Scenarios based only on climate change predicted a range increase and a high geographic stability (>50%) for most species, with different, yet limited, effects on connectivity corridors. Conversely, scenarios based on both climate and land-use change showed a loss in range extent and a low geographic stability (<50%) of both range and dispersal corridors for most species. Main conclusions Scenarios considering both climate and land-use change provide predictions on invasion risk that overturn those including only climate change. The effect of global warming alone would lead to a considerable range expansion of all species. Conversely, when land-use change is added, a potential loss in suitable habitat and dispersal corridors is predicted for alien squirrels, hence limiting their range expansion. We recommend using multiple drivers in models to obtain reliable predictions for implementing biosecurity policies related to invasive alien species.

• Dunham, N. T., McNamara, A., Shapiro, L., Phelps, T., Wolfe, A. N. & Young, J. W. (2019) Locomotor kinematics of tree squirrels (Sciurus carolinensis) in free-ranging and laboratory environments: Implications for primate locomotion and evolution. Journal of Experimental Zoology Part A: Ecological and Integrative Physiology 331(2): 103-119.

Abstract The grasping capabilities and gait kinematics characteristic of primates are often argued to be adaptations for safely moving on small terminal branches. The goal of this study was to identify whether Eastern gray squirrels (Sciurus carolinensis), arboreal rodents that frequently move and forage on small branches, lack primate-like grasping and gait patterns, and arguably represent extant analogs of a stem primate ancestor?adjust gait kinematics to narrow and nonhorizontal branches. We studied locomotor kinematics of free-ranging and laboratory-housed squirrels moving over various substrates. We used high-speed video to film (a) a population of free-ranging squirrels moving on natural substrates and (b) laboratory-housed squirrels moving on horizontal poles. Substrates were coded as small, medium, or large relative to squirrel trunk diameter, and as inclined, declined, or horizontal. Free-ranging squirrels used more gallops and half-bounds on small- and medium-sized substrates, and more high-impact bounds, with reduced limb-lead durations, on declined substrates. Laboratory squirrels moved at higher speeds than free-ranging squirrels and responded to decreasing diameter by using more gallops and half-bounds, lowering speed, and?controlling for speed?increasing mean duty factor, mean number of supporting limbs, and relative forelimb lead duration. Our inability to detect substantial diameter or orientation-related gait adjustments in the wild may be due to a limited accounting of confounding influences (e.g., substrate compliance). Ultimately, studies assessing stability measures (e.g., center of mass fluctuations and peak vertical force) are required to assess whether primates’ enhanced grasping and gait patterns engender performance advantages on narrow or oblique substrates.

• Falvo, C. A., Koons, D. N. & Aubry, L. M. (2019) Seasonal climate effects on the survival of a hibernating mammal. Ecology and Evolution 9(7): 3756-3769.

Abstract Global climate change and associated regional climate variability is impacting the phenology of many species, ultimately altering individual fitness and population dynamics. Yet, few studies have considered the effects of pertinent seasonal climate variability on phenology and fitness. Hibernators may be particularly susceptible to changes in seasonal climate since they have a relatively short active season in which to reproduce and gain enough mass to survive the following winter. To understand whether and how seasonal climate variability may be affecting hibernator fitness, we estimated survival from historical (1964?1968) and contemporary (2014?2017) mark?recapture data collected from the same population of Uinta ground squirrels (UGS, Urocitellus armatus), a hibernator endemic to the western United States. Despite a locally warming climate, the phenology of UGS did not change over time, yet season-specific climate variables were important in regulating survival rates. Specifically, older age classes experienced lower survival when winters or the following springs were warm, while juveniles benefited from warmer winter temperatures. Although metabolic costs decrease with decreasing temperature in the hibernacula, arousal costs increase with decreasing temperature. Our results suggest that this trade-off is experienced differently by immature and mature individuals. We also observed an increase in population density during that time period, suggesting resources are less limited today than they used to be. Cheatgrass is now dominating the study site and may provide a better food source to UGS than native plants did historically.

• Fisher, D. N., Wilson, A. J., Boutin, S., Dantzer, B., Lane, J. E., Coltman, D. W., Gorrell, J. C. & McAdam, A. G. (2019) Social effects of territorial neighbours on the timing of spring breeding in North American red squirrels. Journal of Evolutionary Biology 0(0).

Abstract Organisms can affect one another’s phenotypes when they socially interact. Indirect genetic effects occur when an individual’s phenotype is affected by genes expressed in another individual. These heritable effects can enhance or reduce adaptive potential, thereby accelerating or reversing evolutionary change. Quantifying these social effects is therefore crucial for our understanding of evolution, yet estimates of indirect genetic effects in wild animals are limited to dyadic interactions. We estimated indirect phenotypic and genetic effects, and their covariance with direct effects, for the date of spring breeding in North American red squirrels (Tamiasciurus hudsonicus) living in an array of territories of varying spatial proximity. Additionally, we estimated indirect effects and the strength of selection at low and high population densities. Social effects of neighbours on the date of spring breeding were different from zero at high population densities but not at low population densities. Indirect phenotypic effects accounted for a larger amount of variation in the date of breeding than differences attributable to the among-individual variance, suggesting social interactions are important for determining breeding dates. The genetic component to these indirect effects was however not statistically significant. We therefore showcase a powerful and flexible method that will allow researchers working in organisms with a range of social systems to estimate indirect phenotypic and genetic effects, and demonstrate the degree to which social interactions can influence phenotypes, even in a solitary species.

• Flaherty, M. & Lawton, C. (2019) The regional demise of a non-native invasive species: the decline of grey squirrels in Ireland. Biological Invasions

Following the introduction of the grey squirrel (Sciurus carolinensis) to Ireland in 1911, the species quickly established and spread to cover the eastern half of Ireland. Historically, the River Shannon has delineated the western boundary of its distribution in Ireland, however the factors limiting the spread of the species westwards were unclear. The aims of this study were to assess the current squirrel distribution in the area directly bordering the River Shannon, and to identify habitat types and landscape characteristics that could be facilitating or impeding the spread of grey squirrels in Ireland. The current distribution was established through hair tube and live trapping surveys and through sightings from a citizen science survey. Grey squirrels are absent or in very low numbers in much of the study area. In some areas, red squirrels have reappeared where they previously had been displaced by the grey squirrel. Discriminant function analysis was used to identify significant differences in habitat types and landscape characteristics between a region with high grey squirrel occurrence records and a region where they are now rare. Several landscape attributes were found to be significantly different, including the presence of pine marten, water bodies, peatland and coniferous forests. The area in which grey squirrels have disappeared overlaps with the core pine marten population range, and in a landscape that is more fragmented than the areas in which grey squirrel are continuing to be invasive. The demise of the grey squirrel in Ireland is more widespread than previously believed.

• Franklin, C. M. A., Macdonald, S. E. & Nielsen, S. E. (2019) Can retention harvests help conserve wildlife? Evidence for vertebrates in the boreal forest. Ecosphere 10(3): e02632.

Abstract Retention harvesting, or the approach of leaving live mature trees behind during forest harvest, is used in natural disturbance-based management to mitigate the effects of logging on biodiversity. However, responses of many boreal vertebrates to variable retention harvesting are unknown. We investigated the influence of different retention levels in forest harvests on stand use by wildlife 15-18 yr post-harvest using a combination of surveys of wildlife signs (scats, middens) and camera trapping. Site-level measures of forest structure, including canopy cover, horizontal cover, tree height, tree diameter, basal area, cover of downed coarse woody material, and understory plant cover, were used to describe post-harvest differences in habitats used by common wildlife species in northwest Alberta’s boreal forest. Stand use of six species (black bear, coyote, fisher, red squirrel, wolverine, woodland caribou) increased with level of retention, while stand use of two species (grouse, snowshoe hare) declined with retention level. Retention level did not significantly affect stand use of five species (American marten, Canada lynx, deer, gray wolf, moose). Higher levels of retention characterized by greater canopy cover, basal area, and abundance of deadwood were associated with use of forest habitats by late-seral species. Woodland caribou, a species of conservation concern, was detected only in harvested stands with at least 20% retention. Greater understory and horizontal cover characterized lower levels of retention being attractive for early-seral species. These findings demonstrate the value of retention harvesting for conservation of wildlife species in boreal forest, while highlighting the challenge of managing forests for multiple species with different habitat preferences.

• Gibbs, J. P., Buff, M. F. & Cosentino, B. J. (2019) The Biological System—Urban Wildlife, Adaptation, and Evolution: Urbanization as a Driver of Contemporary Evolution in Gray Squirrels (Sciurus carolinensis). Understanding Urban Ecology: An Interdisciplinary Systems Approach. Hall, M. H. P.& Balogh, S. B. Cham, Springer International Publishing: 269-286.

Urbanization dramatically reduces levels of regional biodiversity, yet urban environments can serve as refuges for biodiversity because some biological stressors are more prevalent outside cities than inside. We evaluated whether urbanization can promote rare genotypes, focusing on gray squirrels (Sciurus carolinensis), which have genetically based color morphs visible to the naked eye. Combining data on 6681 occurrences of gray squirrels derived from an Internet-based, participatory research program (SquirrelMapper) and mined from an online image sharing platform (Flickr), we found the probability of the black morph increases with the extent of urban land cover. An internet-based game that crowdsourced search times of humans to find gray squirrels revealed a distinct camouflage advantage of gray morphs over black in early successional forests where hunting occurs but not in urban areas where hunting is prohibited. The black morph was strongly underrepresented (9%) within a sample of road-killed squirrels in contrast to its frequency (33%) among live squirrels in the same area, likely due to the black morph’s greater conspicuousness on pavement, which in turn facilitates driver avoidance and thereby favors the black morph in cities where vehicles are the primary source of squirrel mortality. Together these processes can generate remarkably steep phenotypic clines along urbanization gradients and will likely continue to shape morphological evolution of the species given that hunting pressures are declining throughout the species’ range while road traffic intensifies. This study suggests cities can serve as refuges for rare genotypes by neutralizing selective pressures that favor a widespread morph in the rural landscape while creating novel selective pressures against the widespread morph within urban areas.

• Hardouin, E., Baltazar-Soares, M., Schilling, A.-K., Butler, H., García Rodríguez, O., Crowley, E., Liang, W.-J., Meredith, A., Lurz, P. W., Forster, J., Kenward, R. E., Hodder, K. H., A Correspondence, E., H Hardouin, K. & Hodder (2019) Conservation of genetic uniqueness in remaining populations of red squirrels (Sciurus vulgaris L.) in the South of England. Ecology and Evolution:

The Eurasian red squirrel (Sciurus vulgaris) is an emblematic species for conservation, and its decline in the British Isles exemplifies the impact that alien introductions can have on native ecosystems. Indeed, red squirrels in this region have declined dramatically over the last 60 years due to the spread of squirrelpox virus following the introduction of the gray squirrel (Sciurus carolinensis). Currently, red squirrel populations in Britain are fragmented and need to be closely monitored in order to assess their viability and the effectiveness of conservation efforts. The situation is even more dramatic in the South of England, where S. vulgaris survives only on islands (Brownsea Island, Furzey Island, and the Isle of Wight). Using the D‐loop, we investigated the genetic diversity and putative ancestry of the squirrels from Southern England and compared them to a European dataset composed of 1,016 samples from 54 populations. We found that our three populations were more closely related to other squirrels from the British Isles than squirrels from Europe, showed low genetic diversity, and also harbored several private haplotypes. Our study demonstrates how genetically unique the Southern English populations are in comparison with squirrels from the continental European range. We report the presence of four private haplotypes, suggesting that these populations may potentially harbor distinct genetic lineages. Our results emphasize the importance of preserving these isolated red squirrel populations for the conservation of the species. Our study demonstrates how genetically unique the Southern English populations are in comparison with squirrels in the non‐threatened European range. We report the presence of four private haplotypes, suggesting that these populations may potentially harbour distinct genetic lineages. Our results emphasise the importance of preserving these isolated red squirrel populations for the conservation of the species.

• Holmes, P., Everest, D. J., Spiro, S., Wessels, M. & Shuttleworth, C. (2019) First report of dermatophilosis in wild European red squirrels (Sciurus vulgaris). Veterinary Record Case Reports 7(3): e000838.

The carcases of two wild red squirrels (Sciurus vulgaris) from Anglesey, North Wales, which had severe skin lesions, were presented to the Animal and Plant Health Agency. One was found dead in December 2017 and the second was found moribund in January 2018 but died 2 hours later. Both were examined at postmortem. The affected skin consisted of raised, firm, crusty lesions with pale subcutaneous necrotic tissue over the head particularly the face, eyelids and forehead. In addition, one had similar lesions on the scrotal and perineal area and along one hind leg extending to the foot. Postmortem examination, bacteriology and histopathology confirmed the presence of disease due to Dermatophilus congolensis with typical gross and pathological lesions. Squirrelpox virus was not detected. Here, we report the first description of dermatophilosis in European red squirrels, another important differential diagnosis of skin disease in a nationally declining population.

• Hutchinson, M. (2019) Releasing grey squirrels into the wild. Veterinary Record 23 February 2019: 257-258.

• Hämäläinen, S., Fey, K. & Selonen, V. (2018) Habitat and nest use during natal dispersal of the urban red squirrel (Sciurus vulgaris). Landscape and Urban Planning 169: 269-275.

As urban environments differ from the natural environment, the ability of a species to move in and use variable land composition types determines its fate in the urban environment. In many mammalian species, the selection of home ranges mainly occurs during natal dispersal. Thus, habitat selection of juvenile individuals greatly determines where animals are found in the city. Here, our goal is to understand how an originally forest-dwelling rodent selects components of an urban habitat. We used radio telemetry to record the habitat and nest use of Eurasian red squirrels (Sciurus vulgaris) in the city of Turku in southwestern Finland, with the main focus on habitat use during natal dispersal. We found that although the red squirrels preferred areas with more trees available than in the surrounding urban landscape they nevertheless often used sites with only a few trees. This result was highlighted by the analysis of the home range scale, as the area used did not differ greatly from the habitat composition of the available landscape. Juvenile red squirrels used the deciduous land cover type more often during movements in the natal area than during dispersal and explorative movements; however, they also settled in areas with less deciduous land cover type than in the natal area. Our results show that in urban areas movements of red squirrels are not restricted to their natural habitat type, which is a coniferous forest, but that they seem to be well adapted to urban areas, being able to utilize urban structures.

• Jagiello, Z. A., Dyderski, M. K. & Dylewski, Ł. (2019) What can we learn about the behaviour of red and grey squirrels from YouTube? Ecological Informatics 51: 52-60.

Abstract: Citizen science has emerged as an important tool in biology, ecology, and conservation studies. In this paper, we examined YouTube (YT) videos featuring the behaviour of red (Sciurus vulgaris) and grey (Sciurus carolinensis) squirrels in Europe, in both urban and forest habitats. Our study shows that specific behaviours of these squirrels, such as aggression, grooming and calling, have frequently been recorded on YT videos. The present paper uses this open data for the first time to compare differences between the frequency of some types of behaviour in two habitats. Based on detrended correspondence analysis, we show a significant difference in sets of behaviours between two squirrel species. Our investigation show that YT can be a source for monitoring the behaviour of wild species, especially in urban habitats, thus affording insights into the species plasticity of urban individuals. YT, as part of citizen science, is a potential source of information in behavioural ecology.

• Jokinen, M., Hanski, I., Numminen, E., Valkama, J. & Selonen, V. (2019) Promoting species protection with predictive modelling: Effects of habitat, predators and climate on the occurrence of the Siberian flying squirrel. Biological Conservation 230: 37-46.

Abstract: Species distribution models (SDMs) can be used to predict species occurrence and to seek insight into the factors behind observed spatial patterns in occurrence, and thus can be a valuable tool in species conservation. In this study, we used MaxEnt software to explain the occurrence of a protected forest-dwelling species, the Siberian flying squirrel. We produce occurrence maps covering the main distribution area for the species in the European Union. Using an exceptionally extensive presence-absence dataset collected with a standardized method, we evaluated the relative role of predation pressure, climate, and amount of habitat affecting flying squirrel occurrence. We found that regional variation in mean winter temperature had relatively large predictive power for flying squirrel occurrence. In addition, the regional abundance of flying squirrels was partly explained by differences in predation pressure. The results also support the conclusion that areas with older forests and nearby agricultural areas are optimal for the species. Our study shows that multiple factors affect the species’ occurrence in large spatial scales. We also conclude that climate is having a large effect on species occurrence, and thus the changing climate has to be taken into account in conservation planning. Our results help conservation managers in targeting surveys and protection measures on various spatial scales, and decision makers in focusing on the factors that drive the species’ occurrence. Our results also indicate that we would need additional tools and measures in the EU for achieving a favourable conservation status of those species that occur in commercial forests.

• Kelt, D. A., Heske, E. J., Lambin, X., Oli, M., Orrock, J. L., Ozgul, A., Pauli, J. N., Prugh, L., Sollmann, R. & Sommer, S. (2019) Advances in population ecology and species interactions in mammals. Mammalia 100(3): 965-1007.

Abstract: The study of mammals has promoted the development and testing of many ideas in contemporary ecology. Here we address recent developments in foraging and habitat selection, source–sink dynamics, competition (both within and between species), population cycles, predation (including apparent competition), mutualism, and biological invasions. Because mammals are appealing to the public, ecological insight gleaned from the study of mammals has disproportionate potential in educating the public about ecological principles and their application to wise management. Mammals have been central to many computational and statistical developments in recent years, including refinements to traditional approaches and metrics (e.g., capture-recapture) as well as advancements of novel and developing fields (e.g., spatial capture-recapture, occupancy modeling, integrated population models). The study of mammals also poses challenges in terms of fully characterizing dynamics in natural conditions. Ongoing climate change threatens to affect global ecosystems, and mammals provide visible and charismatic subjects for research on local and regional effects of such change as well as predictive modeling of the long-term effects on ecosystem function and stability. Although much remains to be done, the population ecology of mammals continues to be a vibrant and rapidly developing field. We anticipate that the next quarter century will prove as exciting and productive for the study of mammals as has the recent one.

• Kobayashi, S., Denda, T., Placksanoi, J., Waengsothorn, S., Aryuthaka, C., Panha, S. & Izawa, M. (2019) The pollination system of the widely distributed mammal-pollinated Mucuna macrocarpa (Fabaceae) in the tropics. Ecology and Evolution 0(0).

Abstract Although the pollinators of some plant species differ across regions, only a few mammal-pollinated plant species have regional pollinator differences in Asia. Mucuna macrocarpa (Fabaceae) is pollinated by squirrels, flying foxes, and macaques in subtropical and temperate islands. In this study, the pollination system of M. macrocarpa was identified in tropical Asia, where the genus originally diversified. This species requires ‘explosive opening’ of the flower, where the wing petals must be pressed down and the banner petal pushed upward to fully expose the stamens and pistil. A bagging experiment showed that fruits did not develop in inflorescences (n = 66) with unopened flowers, whereas fruits developed in 68.7% of inflorescences (n = 131) with opened flowers. This indicated that the explosive opening is needed for the species to reproduce. Four potential pollinator mammals were identified by a video camera-trap survey, and 78.3% and 60.1% of monitored inflorescences (n = 138) were opened by gray-bellied squirrels (Callosciurus caniceps) and Finlayson’s squirrels (C. finlaysonii), respectively, even though more than 10 mammal species visited flowers. Nectar was surrounded by the calyx, and the volume and sugar concentration of secreted nectar did not change during the day. This nectar secretion pattern is similar to those reported by previous studies in other regions. These results showed that the main pollinators of M. macrocarpa in the tropics are squirrels. However, the species’ nectar secretion pattern is not specifically adapted to this particular pollinator. Pollinators of M. macrocarpa differ throughout the distribution range based on the fauna present, but there might not have been no distinctive changes in the attractive traits that accompanied these changes in pollinators.

• Kumar, A., Melotti, J. R., Cooley, T. M. & Fitzgerald, S. D. (2019) Mortality Due to Toxoplasmosis in Suburban Eastern Fox Squirrels (Sciurus niger) in Michigan, USA. Journal of Wildlife Diseases 55(1): 213-217, 215.

Three separate mortality events affecting wild eastern fox squirrels (Sciurus niger) were investigated in suburban areas within southeastern Michigan, US over a 3-yr period from the summer of 2015 through the winter of 2017. A total of seven squirrels were submitted for investigation. The squirrels were generally in fair to good body condition with moderate fat deposits. The tissues that most commonly exhibited gross or histologic lesions included the lungs, liver, and heart, whereas spleen and brain exhibited lesions less frequently. Lung lesions in all seven squirrels consisted of moderate interstitial pneumonia with necrosis and moderate to high numbers of protozoal organisms. Livers in four out of seven squirrels had multifocal necrosis associated with low to moderate numbers of protozoal organisms. Three out of seven brains examined had mild nonsuppurative meningoencephalitis with widely scattered protozoal cysts. Protozoal organisms observed in various tissues were strongly immunoreactive to Toxoplasma gondii antibody by immunohistochemical staining. Other primary disease conditions tested for included West Nile virus infection, pesticides, and anticoagulants. Toxoplasma gondii can cause disease and mortality in a variety of wild squirrel species, especially near human settlements, and would merit more attention.

• Lilly, M. V., Lucore, E. C. & Tarvin, K. A. (2019) Eavesdropping grey squirrels infer safety from bird chatter. PloS one 14(9): e0221279.

When multiple species are vulnerable to a common set of predators, it is advantageous for individuals to recognize information about the environment provided by other species. Eastern gray squirrels (Sciurus carolinensis) and other small mammals have been shown to exploit heterospecific alarm calls as indicators of danger. However, many species–especially birds—emit non-alarm auditory cues such as contact calls when perceived predator threat is low, and such public information may serve as cues of safety to eavesdroppers. We tested the hypothesis that eavesdropping gray squirrels respond to “bird chatter” (contact calls emitted by multiple individuals when not under threat of predation) as a measure of safety. We compared vigilance behavior of free-ranging squirrels in the presence of playbacks of bird chatter vs non-masking ambient background noise lacking chatter after priming them with a playback recording of a red-tailed hawk (Buteo jamaicensis) call. Squirrels responded to the hawk call playbacks by significantly increasing the proportion of time they spent engaged in vigilance behaviors and the number of times they looked up during otherwise non-vigilance behaviors, indicating that they perceived elevated predation threat prior to the playbacks of chatter or ambient noise. Following the hawk playback, squirrels exposed to the chatter treatment engaged in significantly lower levels of vigilance behavior (i.e., standing, freezing, fleeing, looking up) and the decay in vigilance behaviors was more rapid than in squirrels exposed to the ambient noise treatment, suggesting squirrels use information contained in bird chatter as a cue of safety. These findings suggest that eastern gray squirrels eavesdrop on non-alarm auditory cues as indicators of safety and adjust their vigilance level in accordance with the vigilance level of other species that share the same predators.

• Lioy, S., Marsan, A., Balduzzi, A., Wauters, L., Martinoli, A. & Bertolino, S. (2019) The management of the introduced grey squirrel seen through the eyes of the media. Biological Invasions.

Abstract: Communication plans are fundamental for the success of conservation programs, especially when dealing with alien species. In a media-saturated society the effectiveness of communication could be evaluated through the information reported by the media, as an indicator of public attitudes towards a particular issue. We evaluated the effectiveness of a communication campaign by analysing the perception of the media towards management activities to control grey squirrel populations. A media content analysis was performed to classify the news in categories regarding their attitude towards the project. A total of 166 articles were analysed, which corresponds to a mean of one article every 9 days along the 4-years project. News followed peaks of interest in specific periods, generally related with milestones of the project. Half of the negative news regarded management activities in an urban area, despite the softer management approach adopted, with surgical sterilization of animals instead of their euthanasia after live-trapping. A reinforcement of the communication focused in Liguria inverted the trend of negative news which decreased over time, indicating the effectiveness of communication activities. Public support is fundamental for the successful implementation of eradication and control plans. Considering the role of media in informing and driving public opinion, media monitoring is routinely included in the development of management projects. We suggest that a media content analysis can be used as a performance indicator to evaluate the communication effectiveness of conservation projects, helping to identify negative trends so to proactively react.

• Marr, M. M. & MacLeod, N. (2019) Geographical variation in Eurasian red squirrel (Sciurus vulgaris L., 1758) mandibles and the issue of subspecies-level organization: a failure of history? Biological Journal of the Linnean Society 128(2): 337-359.

Sciurus vulgaris is a widespread, highly polytypic tree squirrel species, under which a large number of subspecies have been described. This study tests the robustness of the current subspecific classification by using geometric morphometrics to quantify morphological variation in mandible shape, along with canonical variates analysis to test hypotheses of morphological distinctiveness. Patterns of mandible allometry were examined in eight out of 16 currently recognized subspecies. The significance of the discriminate functions was tested statistically, and the iterative jackknife procedure was applied to evaluate stability of the subspecies-specific discriminant functions. Applying criteria that focus on shape diagnosability, rather than mean group differences, indicates that most regional subspecific groupings show intergradations and continuity in mandible shape and size and that allometric effects on mandible shape are negligible. Evidence of a distinct subspecies confined to the Iberian Peninsula (Sciurus vulgaris infuscatus) and a discrete group originating from an extinct, 19th century population in Dorset, UK were identified based on these mandibular data. All other regional subspecific groupings were not diagnosably different. These results suggest that most red squirrel subspecies might represent non-diagnosable morphological variants whose taxonomic validity seems doubtful. More generally, our results highlight the importance of applying objective, quantitative and reproducible criteria to the issue of subspecies delimitation.

• Martinig, A. R., McAdam, A. G., Dantzer, B., Lane, J. E., Coltman, D. W. & Boutin, S. (2019) The new kid on the block: immigrant males win big whereas females pay fitness cost after dispersal. Ecology Letters n/a(n/a).

Abstract: Dispersal is nearly universal; yet, which sex tends to disperse more and their success thereafter depends on the fitness consequences of dispersal. We asked if lifetime fitness differed between residents and immigrants (successful between-population dispersers) and their offspring using 29 years of monitoring from North American red squirrels (Tamiasciurus hudsonicus) in Canada. Compared to residents, immigrant females had 23% lower lifetime breeding success (LBS), while immigrant males had 29% higher LBS. Male immigration and female residency were favoured. Offspring born to immigrants had 15?43% lower LBS than offspring born to residents. We conclude that immigration benefitted males, but not females, which appeared to be making the best of a bad lot. Our results are in line with male-biased dispersal being driven by local mate competition and local resource enhancement, while the intergenerational cost to immigration is a new complication in explaining the drivers of sex-biased dispersal.

• McRobie, H. R., Moncrief, N. D. & Mundy, N. I. (2019) Multiple origins of melanism in two species of North American tree squirrel (Sciurus). BMC Evolutionary Biology 19(1): 140.

Background While our understanding of the genetic basis of convergent evolution has improved there are still many uncertainties. Here we investigate the repeated evolution of dark colouration (melanism) in eastern fox squirrels (Sciurus niger; hereafter “fox squirrels”) and eastern gray squirrels (S. carolinensis; hereafter “gray squirrels”). Results We show that convergent evolution of melanism has arisen by independent genetic mechanisms in two populations of the fox squirrel. In a western population, melanism is associated with a 24 bp deletion in the melanocortin-1-receptor gene (MC1RΔ24 allele), whereas in a south-eastern population, melanism is associated with a point substitution in the agouti signalling protein gene causing a Gly121Cys mutation. The MC1R∆24 allele is also associated with melanism in gray squirrels, and, remarkably, all the MC1R∆24 haplotypes are identical in the two species. Evolutionary analyses show that the MC1R∆24 haplotype is more closely related to other MC1R haplotypes in the fox squirrel than in the gray squirrel. Modelling supports the possibility of gene flow between the two species. Conclusions The presence of the MC1R∆24 allele and melanism in gray squirrels is likely due to introgression from fox squirrels, although we cannot completely rule out alternative hypotheses including introgression from gray squirrels to fox squirrels, or an ancestral polymorphism. Convergent melanism in these two species of tree squirrels has evolved by at least two and probably three different evolutionary routes.

• Mendes, C. & Koprowski, J. (2019) Does caching strategy vary with microclimate in endangered Mt. Graham red squirrels? PloS one 14: e0224947.

Food hoarding is a common behavior used by a variety of animals to cope with periods of low food availability. At the retreating edge of species’ distribution, the stressful environment and unfavourable climate conditions may impose severe costs on hoarding behavior. Since relict populations are hotspots for evolution and adaptation, and considering that food hoarding behavior has a strong evolutionary basis, we decided to evaluate the occurrence of behavioral variability in the amount of food cached by the endangered Mount Graham red squirrel (Tamiasciurus fremonti grahamensis). We tested the variation in cache size in response to microclimate, soil relief, vegetation, food availability and squirrel sex. The number of pits excavated by squirrels to cache cones was used as a proxy of cache size and was affected by mountain slope aspect and density of trees. More pits were excavated in the northeast facing slopes. The density of trees negatively affects the cache volume on southwest slopes, but not on northeast slopes. The sex of the resident squirrel also affects the number of pits in the squirrel midden, with males excavating 47% more pits than females. Males and females also presented different responses to the mountain slope aspect, with females excavating more pits on northeastern slopes than on southwestern slopes, whereas male cache size did not vary with the slope aspect. Finally, the squirrel’s caching behavior did not vary in response to midden microclimate variation, a result with possible implications for the survival of the Mt Graham red squirrels, given the predicted temperature increases in the region due to climate change.

• Mendes, C. P., Koprowski, J. L. & Galetti, M. (2019) NEOSQUIRREL: a data set of ecological knowledge on Neotropical squirrels. Mammal Review 0(0).

Abstract: The squirrels (Sciuridae), with 292 species, make up the second most diverse family of rodents. Squirrels play important roles as seed and spore dispersers and seed predators in all regions where they occur. In Neotropical regions, around 28 species of squirrel are recognised. However, our knowledge of the ecology of the Neotropical Sciuridae is severely incomplete, lacking in the most basic ecological information for most species. We reviewed the literature in English, Spanish, and Portuguese, for all squirrel species in the Neotropical biogeographic region, summarising ecological interactions between squirrels and the local biota, population density records, the number of publications, and the distribution of study sites. We found information for 20 squirrel species (71% of the recognised species), from 15 countries, in 48 publications containing 126 population density records and 155 publications containing 649 ecological interactions. The most studied species were Guerlinguetus brasiliensis, Notosciurus granatensis, and Sciurus variegatoides, with 53% of all publications, whereas for eight species of Microsciurus, we found no publications. The density of Neotropical squirrels varied from 0.08 to 100 individuals per km2 and was negatively correlated with forest area. Neotropical squirrels were recorded eating 174 plant taxa, five fungus taxa, four invertebrate taxa, and one species of vertebrate. Palms were common in the diet of squirrels (30 palm species, 27% of feeding records). Squirrels cached 28 plant species, of which 15 were palms. Sixty-five taxa of parasites are documented to occur in Neotropical squirrels, and the most common were Enderleinellus lice. Zoonotic parasites, including trypanosomes, Leptospira spp., Leishmania spp., and plague were also reported. Our review reveals the main information gaps in the current knowledge about the ecology of Neotropical Sciuridae and maps the geographic distribution of the available information throughout South and Central America. Squirrels often thrive in small forest fragments and fulfil important roles as seed dispersers and prey for mesopredators.

Millar, C. I. & Hickman, K. T. (2019) New Records for Douglas’s Squirrel (Tamiasciurus douglasii) in Two Mountain Ranges of the Great Basin. Western North American Naturalist 79(1): 99-109.

Abstract: New records of Douglas’s squirrel (Tamiasciurus douglasii) document the species in 2 mountain ranges of the Great Basin where tree squirrels have not been previously reported, including 2 sites in the White Mountains, California, and 1 site in the Desert Creek Mountains, Nevada. In the White Mountains, squirrels were photographed by a camera trap on 9 different dates in winter and spring 2016–2017 at a site on the east side of the range crest near the Crooked Creek Field Station of the White Mountain Research Center (3125 m). In early winter 2018, two Douglas’s squirrels were observed and photographed 25 km distant and 716 m lower in Leidy Canyon (2409 m) on the lower east side of the White Mountains, 3.5 km west of the Nevada state line. In the Desert Creek Mountains, Douglas’s squirrels were observed on 4 days in autumn 2017 and winter 2018 along Desert Creek from 2005 m to 2307 m. We present evidence to suggest that at least the White Mountains records represent recent colonization(s) and, as such, call into consideration the question of how montane mammals are able to migrate into isolated mountain ranges of the Great Basin during warm Holocene climates.

• Mitsuzuka, W., Kato, M. & Oshida, T. (2019) Seasonal pelage color change of two sympatric arboreal squirrel species in the subarctic region. The European Zoological Journal 86: 443-451.

Some mammal species exhibit pelage color change with seasonal molt. Seasonal molt and pelage color change are beneficial to thermoregulation and concealment, associated with seasonal environmental change. The Eurasian red squirrel Sciurus vulgaris Linnaeus, 1758 and the Siberian flying squirrel Pteromys volans (Linnaeus, 1758) are arboreal and sympatrically distributed in the subarctic northern Eurasian Continent and Sakhalin and Hokkaido islands. We expect that diurnal S. vulgaris may demonstrate more conspicuous difference between summer and winter pelages than nocturnal P. volans, because of its protective coloration in each season. To test this conjecture, we investigated their seasonal pelage color change. To diminish the effect of geographic variation in pelage color, we chose S. vulgaris orientis Thomas, 1906 and P. volans orii (Kuroda, 1921), which are endemic subspecies of Hokkaido Island, Japan. We used skin and stuffed specimens and frozen materials and categorized them into two pelage groups (summer and winter pelages) based on collection date. Pelage color characteristics were measured with a spectrophotometer for lightness, redness and yellowness. Countershading was examined by comparing dorsal and ventral lightness. Both subspecies showed lighter winter pelage than summer pelage, suggesting their greyish-white winter pelage was beneficial to concealment from predators during winter. As we expected, seasonal changes of redness and yellowness were more clearly recognized in S. vulgaris than in P. volans. As S. vulgaris is diurnal and vulnerable to attack by diurnal avian predators, reddish and yellowish pelage patterns may be important for concealment. Because it is nocturnal, P. volans may not need this reddish and yellowish pelage. Sciurus vulgaris also had a remarkably counter-shaded body, indicating that its body may reduce predation risk from daytime visual predators. Differences in seasonal pelage color change of these two arboreal squirrels may be caused by their different circadian rhythms.

• Nyakatura, J. A., Baumgarten, R., Baum, D., Stark, H. & Youlatos, D. (2019) Muscle internal structure revealed by contrast-enhanced µCT and fibre recognition: The hindlimb extensors of an arboreal and a fossorial squirrel. Mammalian Biology.

In individuals of similar body mass representing closely related species with different lifestyles, muscle architectural properties can be assumed to reflect adaptation to differing, lifestyle-related functional demands. We here employ a fibre recognition algorithm on contrast-enhanced micro-computed tomography (µCT) scans of one specimen each of an arboreal (Sciurus vulgaris) and a fossorial (Spermophilus citellus) sciuromorph rodent. The automated approach accounts for potential heterogeneity of architectural properties within a muscle by analysing all fascicles that compose a muscle. Muscle architectural properties (volume, fascicle length and orientation, and force-generating capacity) were quantified in 14 hindlimb (hip, knee, and ankle) extensor muscles and compared between specimens. We expected the arboreal squirrel to exhibit greater force-generating capacity and a greater capacity for length change allowing more powerful hindlimb extension. Generally and mostly matching our expectations, the S. vulgaris specimen had absolutely and relatively larger extensor muscles than the S. citellus specimen which were thus metabolically more expensive and demonstrate the relatively larger investment into powerful hindlimb extension necessary in the arboreal context. We conclude that detailed quantitative data on hindlimb muscle internal structure as was gathered here for a very limited sample further lends support to the notion that muscle architecture reflects adaptation to differential functional demands in closely related species with different locomotor behaviours and lifestyles.

• Nzalak, J. O., Wanmi, N. & Samuel, M. O. (2015) Morphometric Study on the Digestive System of the Wild Gray Squirrel (Sciurus carolinensis). Journal of Veterinary Anatomy 8(2): 59-68.

The present study was conducted on Five apparently healthy wild gray squirrel of different ages and sexes. The various parts of the digestive system of each animal were dissected out and were weighed and measured. The small intestine was observed to have the highest mean weight (5.86 ± 0.8 g) followed by the large intestine (2.98 ± 0.49 g). Of all the intestinal segments, the duodenum was observed to have the highest mean weight (2.49 ± 0.32 g) followed by the jejunum (2.03 ± 0.29 g) The mean length of the gastrointestinal tract (GIT) was observed to be 136.10 ± 10 cm with the small intestine having a mean length value 90.73 ± 10.31 cm and was ob- served to have the longest segment, followed by the large intestine (31.54 ± 2.25 cm). Out of all the intestinal segments (small and large) the jejunum was observed to have the longest segment (32.39 ± 4.19 cm) followed by the ileum (27.93 ± 4.31 cm) and the duodenum (25.53 ± 2.67 cm), respectively. The liver was observed to have the mean weight (18.36 ± 0.77 g) and had no gall bladder. The caecum had the least value of the segments of the large intestine; the colon and rectum had almost the same mean length (14.64 ± 1.84 cm and 14.54 ± 0.60 cm, respectively). The absence of gallbladder will have a direct effect on the emulsification of fats. The short length of the colon and rectum indicates that no digestion takes place in those regions.

• Olson, E. R., Martin, J. G., Anich, P. S. & Kohler, A. M. (2019) Ultraviolet fluorescence discovered in New World flying squirrels (Glaucomys). Journal of Mammalogy.

Abstract: Fluorescence of visible wavelengths under ultraviolet (UV) light has been previously detected in a wide range of birds, reptiles, and amphibians and a few marsupial mammals. Here, we report the discovery of vivid UV fluorescence of the pelage in Glaucomys, the New World flying squirrels. Fluorescence in varying intensities of pink was observed in females and males of all extant species (G. oregonensis, G. sabrinus, and G. volans) across all sampled geographic areas in North and Central America and a temporal range of 130 years. We observed fluorescence in museum specimens (n = 109) and wild individuals (n = 5) on both dorsal and ventral surfaces. Museum specimens of three co-occurring, diurnal sciurid species (Sciurus carolinensis, S. niger, and Tamiasciurus hudsonicus) were also examined but did not fluoresce. The ecological significance of this trait in the nocturnal–crepuscular flying squirrels warrants further investigation.

• Pagès, M., Fischer, A., van der Wal, R. & Lambin, X. (2019) Empowered communities or “cheap labour”? Engaging volunteers in the rationalised management of invasive alien species in Great Britain. Journal of Environmental Management 229: 102-111.

Abstract: Volunteers are increasingly involved in the delivery of nature conservation policies, usually supported by a twofold rationale: volunteering can (a) enhance citizen participation in environmental governance and (b) ensure a workforce is in place to support conservation work in times of budget shortages. Here, we ask how these two rationales correspond to volunteers’ own motivations to engage in a specific nature conservation activity, namely the control of invasive alien species (IAS). We use qualitative interviews with professional project managers, local group leaders, and volunteers to examine the interactions between policies aiming to rationalise the management of IAS and the motivations for and goals of volunteer engagement. Our findings suggest that although volunteering can lead to positive conservation outcomes, satisfying experiences and empowerment, the different interests do not always align in practice. We investigate the implications of strategies that aim to improve the efficiency of invasive species and volunteer management, and discuss organisational arrangements that reconcile different objectives.

• Pal, R., Thakur, S., Bhattacharya, T. & Sathyakumar, S. (2019) Range extension and high elevation record for the endangered woolly flying squirrel Eupetaurus cinereus in Western Himalaya, India. Mammalia 83(4): 410.

The woolly flying squirrel (Eupetaurus cinereus Thomas, 1888) is one of the least-known endangered mammals of the Himalayas and recorded only from few localities at 2400–3600 m in Hindu Kush and North-Western Himalayas. We report first confirmed record of this species from Upper Bhagirathi Basin, Uttarakhand, Western Himalaya. The squirrel was photo-captured twice in camera traps placed in temperate and alpine habitats. The photo-capture at 4800 m is higher than the described upper elevation range limit of any other flying squirrels. Continuous monitoring would reveal the extent of threats to this rare species in its newly described range.

• Pedreschi, D., García-Rodríguez, O., Yannic, G., Cantarello, E., Diaz, A., Golicher, D., Korstjens, A. H., Heckel, G., Searle, J. B., Gillingham, P., Hardouin, E. A. & Stewart, J. R. (2019) Challenging the European southern refugium hypothesis: Species-specific structures versus general patterns of genetic diversity and differentiation among small mammals. Global Ecology and Biogeography 28(2): 262-274.

Abstract: Aim In this study, we conduct a quantitative meta-analysis to investigate broad patterns of genetic variation throughout large geographical regions in order to elucidate concordant geographical patterns across species and identify common historical processes to better inform the ?cryptic refugia? versus the traditional ?southern refugia? hypothesis debate. Location Europe. Time period Late Pleistocene to present day. Major taxa studied Small mammals (Rodentia, Eulipotyphla). Methods A meta-analysis was performed on large-scale patterns of genetic diversity for 19 species from 59 papers. For each species, haplotype and nucleotide diversity were calculated using the mitochondrial D-loop and compared to the species? range. Results No consistent patterns were observed between mitochondrial DNA (mtDNA) diversity indices (nucleotide and haplotype diversity) and any of the indicators of distribution examined [latitude and longitude (max, min, centre, range)]. The patterns of genetic diversity observed in all the 19 species studied appear to be species-specific. Main conclusions In contrast to the traditional southern refugial hypotheses, we found no evidence for a consistent south?north post-glacial expansion. Instead individual species appear to respond to climate oscillations in niche-specific ways. This individual nature of each species? phylogeographical history indicates a complex web of post-glacial recolonization dynamics across Europe.

• Potash, A. D., Conner, L. M. & McCleery, R. A. (2019) Vertical and horizontal vegetation cover synergistically shape prey behaviour. Animal Behaviour 152: 39-44.

Abstract: An animal’s perception of predation risk varies across a heterogeneous landscape. Animals rely on indirect cues in the environment, including availability of protective cover, openness of sightlines and distance to refuge to evaluate potential predation risk. Interactions between the indirect cues that influence an animal’s perception of predation risk are poorly understood, especially for prey at risk of avian and terrestrial predation. We conducted a giving-up density (GUD) study to examine how interactions between terrestrial predator presence, ceiling (canopy) cover, wall (shrub/grass) cover and distance to nearest tree influence fox squirrel, Sciurus niger, risk perception. The GUD indicates an animal’s perceived predation risk, such that increased GUD (decreased foraging) corresponds to greater perceived risk. We found that fox squirrels perceived cues of predation risk in response to a synergistic interaction between ceiling and wall cover. In open canopy areas, fox squirrels increased GUD where there was also increased wall cover. However, fox squirrels reduced GUD in closed canopy areas where there was also increased wall cover. We attribute the differential effects of wall cover depending on ceiling cover to the effectiveness of open sightlines in decreasing predation risk from avian or terrestrial predators. Our study shows that the effect of vegetation structure on prey risk perception and antipredator behaviour depends on the relative risk posed by avian and terrestrial predators that use different hunting strategies.

• Ratliff, C., Kingsley, L., Kusmierczyk, J., Gentry, J., Russell, K. E. & Heatley, J. J. (2019) Hematologic and biochemical values of the juvenile eastern gray squirrel Sciurus carolinensis. Journal of Zoo and Wildlife Medicine 50(3): 644-649, 646.

Venous blood samples were collected from 64 apparently healthy juvenile Eastern gray squirrels (Sciurus carolinensis) after sedation with midazolam at the Wildlife Center of Texas located in Houston, Texas, during 2012. Blood gas (pH, PCO2, PO2, base excess, bicarbonate, oxygen saturation), electrolyte (sodium, potassium), biochemical (total CO2, ionized calcium, glucose), and hematologic parameters (hematocrit, hemoglobin, complete blood count) were determined using the i-STAT point-of-care analyzer. Sex did not affect any analyte. All squirrels recovered uneventfully and were successfully rehabilitated and released. Most values were as expected based on comparison to other young rodent species. These analyte data for healthy juvenile Eastern gray squirrels may be useful in assessment of Eastern gray squirrel population health and management and treatment of individual squirrels presented in need of medical care.

• Reed, A., Pigage, J. C., Pigage, H. K., Glickman, C. & Bono, J. M. (2019) Comparative analysis of microbiota along the length of the gastrointestinal tract of two tree squirrel species (Sciurus aberti and S. niger) living in sympatry. Ecology and Evolution n/a(n/a).

Abstract Microbiota inhabiting the gastrointestinal (GI) tract of animals has important impacts on many host physiological processes. Although host diet is a major factor influencing the composition of the gut micro-organismal community, few comparative studies have considered how differences in diet influence community composition across the length of the GI tract. We used 16S sequencing to compare the microbiota along the length of the GI tract in Abert’s (Sciurus aberti) and fox squirrels (S. niger) living in the same habitat. While fox squirrels are generalist omnivores, the diet of Abert’s squirrels is unusually high in plant fiber, particularly in winter when they extensively consume fiber-rich inner bark of ponderosa pine (Pinus ponderosa). Consistent with previous studies, microbiota of the upper GI tract of both species consisted primarily of facultative anaerobes and was less diverse than that of the lower GI tract, which included mainly obligate anaerobes. While we found relatively little differentiation between the species in the microbiota of the upper GI tract, the community composition of the lower GI tract was clearly delineated. Notably, the Abert’s squirrel lower GI community was more stable in composition and enriched for microbes that play a role in the degradation of plant fiber. In contrast, overall microbial diversity was higher in fox squirrels. We hypothesize that these disparities reflect differences in diet quality and diet breadth between the species.

Sainsbury , A., Cunningham, A., Gurnel, J., Lurz, P. W. W. & McInnes, C. J. (2019) Releasing grey squirrels into the wild. Veterinary Record 184(23): 714.

• Santicchia, F., Romeo, C., Ferrari, N., Matthysen, E., Vanlauwe, L., Wauters, L. A. & Martinoli, A. (2019) The price of being bold? Relationship between personality and endoparasitic infection in a tree squirrel. Mammalian Biology

Abstract: Individual variation in behaviour can contribute to the heterogeneous distribution of parasites among hosts for example by affecting the probability of encountering infective stages (larvae). Here, we investigated the relationship between host boldness/exploration tendency and gastro-intestinal helminth infection in invasive Eastern grey squirrels (Sciurus carolinensis). We used direct helminth counts, data rarely available in host-parasite studies that often used less reliable indirect estimates of infection. We predicted that bolder animals with a strong exploration tendency will have higher parasite load than shy, less explorative hosts. We examined two parameters of parasite infection: infection status and intensity of infection. Individual personality of 207 grey squirrels was assessed by capture-mark-recapture (CMR), calculating the trappability and trap diversity indices as estimates of boldness and exploration, respectively. Since both indices were strongly correlated, we used PCA to derive a single score (first component) which had a high value for bold, exploring animals. At the end of the study, 77 individuals were euthanized and gastro-intestinal helminths were identified and counted. Overall 73% of grey squirrels were infected by Strongyloides robustus with the intensity of infection varying from 1 to 86 worms (mean ± SE = 10.7 ± 2.1 helminths per host). We found that bolder, more explorative animals were more likely to be infected by S. robustus. However, host personality was not related to parasite intensity, which was instead positively associated with host body mass. Our results confirm that differences in personality-related host behaviour can influence the distribution of infections within host populations and stimulate further questions on whether such personality-parasite relationships may affect the invasion process.

• Santicchia, F., Wauters, L. A., Dantzer, B., Westrick, S. E., Ferrari, N., Romeo, C., Palme, R., Preatoni, D. G. & Martinoli, A. (2019) Relationships between personality traits and the physiological stress response in a wild mammal. Current Zoology.

Glucocorticoids are involved in regulation of an animal’s energetic state. Under stressful situations, they are part of the neuroendocrine response to cope with environmental challenges. Animals react to aversive stimuli also through behavioural responses, defined as coping styles. Both in captive and wild populations, individuals differ in their behaviour along a proactive – reactive continuum. Proactive animals exhibit a bold, active-explorative and social personality, while reactive ones are shy, less active-explorative and less social. Here we test the hypothesis that personality traits and physiological responses to stressors co-vary, with more proactive individuals having a less pronounced glucocorticoid stress response. In wild populations of invasive grey squirrels Sciurus carolinensis, we measured faecal glucocorticoid metabolites (FGMs), an integrated measure of circulating glucocorticoids, and three personality traits (activity, sociability, exploration) derived from open field test (OFT) and mirror image stimulation (MIS) test. Grey squirrels had higher FGMs in autumn than in winter and males with scrotal testes had higher FGMs than non-breeding males. Personality varied with body mass and population density. Squirrels expressed more activity-exploration at higher than at lower density and heavier squirrels had higher scores for activity-exploration than animals that weighed less. Variation in FGM concentrations was not correlated with the expression of the three personality traits. Hence, our results do not support a strong association between the behavioural and physiological stress responses but show that in wild populations, where animals experience varying environmental conditions, the glucocorticoid endocrine response and the expression of personality are uncorrelated traits among individuals.

• Scapin, P., Ulbano, M., Ruggiero, C., Balduzzi, A., Marsan, A., Ferrari, N. & Bertolino, S. (2019) Surgical sterilization of male and female grey squirrels (Sciurus carolinensis) of an urban population introduced in Italy. Journal of Veterinary Medical Science.

We report a successful surgical sterilization procedure for population control of 324 male and female free-ranging grey squirrels (Sciurus carolinensis) in Genoa (Italy). We describe the clinical procedure from the trapping of the animals to their surgical sterilization and release in another part of the city. Live-trapped squirrels were transported to the veterinary clinic within 1-2 hr of capture and maintained in a hospitalization room reserved for them. The waiting period before surgery was kept below 12 hr. The developed procedure has resulted in a survival of 94% of trapped squirrels from surgery to animal release. Sterilized squirrels started to feed in a very short time (1.0-1.5 hr), and after 2-3 days, it was possible to release them in a new area. Amoxicillin was used as a long-acting postoperative antibiotic to reduce the period of captivity. The successful surgical procedure described here can provide an important additional tool for the management of introduced populations of squirrels. We showed that the surgical sterilization of some hundred squirrels is clinically possible and could be included in management strategies aimed at removing critical populations of these species. Moreover, the data allow dosages and operational times in order to provide economic viability assessment of future population control measures.

• Schilling, A.-K., van Hooij, A., Corstjens, P., Lurz, P. W. W., DelPozo, J., Stevenson, K., Meredith, A. & Geluk, A. (2019) Detection of humoral immunity to mycobacteria causing leprosy in Eurasian red squirrels (Sciurus vulgaris) using a quantitative rapid test. European Journal of Wildlife Research 65(3): 49.

Abstract: Eurasian red squirrels (Sciurus vulgaris, ERS) in the British Isles are a recently discovered natural host for Mycobacterium leprae and Mycobacterium lepromatosis. Infected squirrels can develop skin lesions or carry the bacteria without showing clinical signs. Until now the clinical diagnosis of leprosy could only be confirmed in squirrels by isolating DNA of leprosy bacilli from carcasses or by establishing the presence of acid-fast bacilli in skin sections of carcasses with clinical signs. In this study, we assessed the performance of a field-friendly diagnostic test for detection of M. leprae/M. lepromatosis infection in ERS. This up-converting phosphor lateral flow assay (UCP-LFA) is well established for detection of M. leprae specific anti-phenolic glycolipid-I antibodies (αPGL-I) IgM antibodies in humans and associated with bacterial load. Assessment was performed on serum and blood drops from live squirrels and body cavity fluid samples from dead squirrels. Clinically diseased squirrels showed significantly higher αPGL-I levels than healthy animals or subclinically infected individuals (p < 0.0001), both in serum and whole blood drop samples. Subclinically, infected animals were identified using molecular methods to detect the presence of leprosy bacilli DNA in punch biopsy tissue samples. In body cavity fluids, αPGL-I levels antibody levels were lower than in serum or blood drops. This study shows that the αPGL-I UCP-LFAs presented here allows a field-friendly serological confirmation of M. leprae infection in clinically diseased live ERS. For surveillance purposes, the combination of clinical assessment, αPGL-I UCP-LFAs, and molecular methods allow the identification of both diseased animals and subclinically infected animals.

• Schilling, A.-K., Avanzi, C., Ulrich, R. G., Busso, P., Pisanu, B., Ferrari, N., Romeo, C., Mazzamuto, M. V., McLuckie, J., Shuttleworth, C. M., Del-Pozo, J., Lurz, P. W. W., Escalante-Fuentes, W. G., Ocampo-Candiani, J., Vera-Cabrera, L., Stevenson, K., Chapuis, J.-L., Meredith, A. L. & Cole, S. T. (2019) British Red Squirrels Remain the Only Known Wild Rodent Host for Leprosy Bacilli. Frontiers in Veterinary Science 6: 8.

Abstract: Eurasian red squirrels (Sciurus vulgaris) in the British Isles are the most recently discovered animal reservoir for the leprosy bacteria Mycobacterium leprae and Mycobacterium lepromatosis. Initial data suggest that prevalence of leprosy infection is variable and often low in different squirrel populations. Nothing is known about the presence of leprosy bacilli in other wild squirrel species despite two others (Siberian chipmunk [Tamias sibiricus], and Thirteen-lined ground squirrel [Ictidomys tridecemlineatus]) having been reported to be susceptible to experimental infection with M. leprae. Rats, a food-source in some countries where human leprosy occurs, have been suggested as potential reservoirs for leprosy bacilli, but no evidence supporting this hypothesis is currently available. We screened 301 squirrel samples covering four species (96 Eurasian red squirrels, 67 Eastern grey squirrels (Sciurus carolinensis), 35 Siberian chipmunks, and 103 Pallas’s squirrels (Callosciurus erythraeus)) from Europe and 72 Mexican white-throated woodrats (Neotoma albigula) for the presence of M. leprae and M. lepromatosis using validated PCR protocols. No DNA from leprosy bacilli was detected in any of the samples tested. Given our sample-size, the pathogen should have been detected if the prevalence and/or bacillary load in the populations investigated were similar to those found for British red squirrels.

• Schilling, A.-K., Del-Pozo, J., Lurz, P. W. W., Stevenson, K., Avanzi, C., Shuttleworth, C. M., Cole, S. T. & Meredith, A. L. (2019) Leprosy in red squirrels in the UK. Veterinary Record 184(13): 416.

The presence of leprosy in Eurasian red squirrels (Sciurus vulgaris) was first described in Scotland in 2014.1Initially, the causative agent identified was Mycobacterium lepromatosis.1, 2 Consecutive studies demonstrated the presence of M lepromatosis in red squirrels in other areas, including the Isle of Arran, Isle of Wight and the Republic of Ireland. Additionally, an alternative causative agent, Mycobacterium leprae, was identified in red squirrels exclusively on Brownsea Island.3While it is currently unknown whether M lepromatosis has ever …

• Sehrsweeney, M., Wilson, D. R., Bain, M., Boutin, S., Lane, J. E., McAdam, A. G. & Dantzer, B. (2019) The effects of stress and glucocorticoids on vocalizations: a test in North American red squirrels. Behavioral Ecology 30(4): 1030-1040.

Abstract: Acoustic signaling is an important means by which animals communicate both stable and labile characteristics. Although it is widely appreciated that vocalizations can convey information on labile state, such as fear and aggression, fewer studies have experimentally examined the acoustic expression of stress state. The transmission of such public information about physiological state could have broad implications, potentially influencing the behavior and life-history traits of neighbors. North American red squirrels (Tamiasciurus hudsonicus) produce vocalizations known as rattles that advertise territorial ownership. We examined the influence of changes in physiological stress state on rattle acoustic structure through the application of a stressor (trapping and handling the squirrels) and by provisioning squirrels with exogenous glucocorticoids (GCs). We characterized the acoustic structure of rattles emitted by these squirrels by measuring rattle duration, mean frequency, and entropy. We found evidence that rattles do indeed exhibit a “stress signature.” When squirrels were trapped and handled, they produced rattles that were longer in duration with a higher frequency and increased entropy. However, squirrels that were administered exogenous GCs had similar rattle duration, frequency, and entropy as squirrels that were fed control treatments and unfed squirrels. Our results indicate that short-term stress does affect the acoustic structure of vocalizations, but elevated circulating GC levels do not mediate such changes.

• Shuai, L.-Y., Zhou, Y., Yang, Y.-X., Xue, Q.-Q., Xie, Z.-Y. & Zhang, F.-S. (2019) Ecological factors affecting flight initiation distance in Daurian ground squirrels (Spermophilus dauricus). Ethology 39(3): 92-95.

Abstract: More often than not, animals forage under predation risk. Foragers, therefore, face a challenge to balance between two conflicting tasks, namely energy intake and safety. Flight initiation distance (FID, defined as the distance between a prey and a predator when the prey starts to flee) has been widely measured in many taxa to study such economic trade-offs. However, FID may also be affected by limitations on the prey’s ability to detect predators, especially when there is visual obstruction caused by surrounding vegetation. Although both vegetation cover and vegetation height may contribute to such obstruction, the effect of vegetation height on FID has not been well studied. In this study, we explored the effects of vegetation height, vegetation cover and distance to refuge on FID in free-living Daurian ground squirrels (Spermophilus dauricus) inhabiting a grassland in Inner Mongolia, China. Multiple linear regressions suggested that both vegetation height and distance to refuge significantly affected FID in S. dauricus. Ground squirrels fled earlier when vegetation was low or when foraging farther away from a refuge. No significant effect of vegetation cover on FID was detected. Our results have implications for ecologically based pest control, and FID may be used as an effective and easy-to-use behavioral indicator in wildlife management.

• Shuttleworth, C. M., Bertolino, S., Gill, R., Gurnell, J., Hayward, M. W., Kenward, R. E., Lawton, C., Lurz, P. W. W., McInnes, C. J., Mill, A., Trotter, S. & Wauters, L. A. (2019) Releasing grey squirrels into the wild. Veterinary Record 184(12): 389-390.

The introduced grey squirrel is a highly invasive and economically damaging speciesWe were surprised to read Maureen Hutchison describe the evidence that North American eastern grey squirrels (Sciurus carolinensis) have negative impacts on European biodiversity as ‘opinions’ that are ‘opposed by many experts on the subject’.1 A recent peer-reviewed monograph,2 with over 50 international scientific contributors, presented overwhelming evidence that the introduced grey squirrel is a highly invasive and economically damaging species.Bark stripping affects many tree species and not only leads to significant …

• Shuttleworth, C. M., Everest, D. J., Halliwell, E. C., Hulme, B., Wilberforce, L. & Clews-Roberts, R. (2019) Detecting viral infection in red squirrels. Veterinary Record 184(16): 507.

Adenovirus (ADV) infection is geographically widespread in both wild red squirrel (Sciurus vulgaris) populations and captive collections.1 ADV is often asymptomatic but can become pathogenic. It has been associated with significant mortality in some conservation translocation programmes, notably on the island of Anglesey, north Wales.2To investigate whether ADV was present on Anglesey before the first captive-bred red squirrel releases in 2004, we obtained archived hair (shaft and follicles) in 2016, collected during historical microsatellite studies.3 These were 50 samples from animals live-trapped in 2002/3 and two …

• Starkey, A. & delBarco-Trillo, J. (2019) Supplementary feeding can attract red squirrels (Sciurus vulgaris) to optimal environments. Mammalian Biology 94: 134-139.

Abstract: A number of conservation approaches are used to manage threatened species. However, some of these approaches require intensive planning and can often be restricted by funding. Supplementary feeding is a non-invasive and cost-effective approach to manage vulnerable populations, but we lack data on its usefulness. Here we investigated the effects of supplementary feeding on a population of red squirrels (Sciurus vulgaris), a UK priority species which faces competition from the non-native grey squirrel (Sciurus carolinensis). The study took place October-December 2015, lasting 8 weeks. Twenty feeders were installed 1 week prior to the beginning of the study in a protected woodland free from grey squirrels, either containing food (full feeders) or no food (empty feeders), and squirrel abundance before and after feeding was recorded at each feeder (for a total of 27 feeding and recording events). Six times more squirrels were seen at full feeders, and numbers increased by 7 fold after feeding. We also observed that the activity of red squirrels in the vicinity of full feeders increased during the course of the study. Eighty-five hair samples were collected during the study, all of which were found at full feeders. Results demonstrate red squirrels can differentiate between full and empty feeders, suggesting their awareness increases when supplementary food is present. Increased abundance of squirrels at full feeders after feeding times not only implies that squirrels are attracted to and can benefit by supplementation, it also shows that food supplementation can be used to regulate the movement of individuals across habitats. Understanding how red squirrel populations are affected by supplementary feeding will contribute towards existing conservation efforts to improve this species future survival.

• Stirkė, V. (2019) Ecological aspects of Red Squirrel (Sciurus vulgaris) dreys in city parks. Acta Zoologica Academiae Scientiarum Hungaricae 65(1): 75-84.

Modified habitats often pose challenges to native fauna. Red Squirrels (Sciurus vulgaris) successfully inhabits territory of the cities. However research on quite abundant Red Squirrel populations in urban parks of Lithuania was lacking. The investigation was carried out in two city parks (area of Giruliai park 152 ha, Pasakų park forest – 31 ha) in Vilnius, Lithuania. Both areas were mainly coniferous, dominated with pine (Pinus sylvestris). Drey (N = 415) counts were done in February and March 2014–2016. Dreys were found in eight tree species. The Red Squirrels’ dreys were mainly built in most widespread tree species: pine (84.8% of all dreys) and birch (8.1%). Most of the found dreys were built on the side branches at the stem (47.7%). 21.8% of the dreys were built on the branches further from the stem, 18.5% on the top branches and 12% in the stem fork-off. Most of the dreys were set in the southern exposition (33.9%). Average number of red squirrels in both parks in 2014–2016 was 0.20±0.01 squirrels/ha, the average number of dreys – 0.75±0.05 dreys/ha. The population of urban red squirrels in Vilnius city parks can be regarded as stable and having medium abundance, which is supported by supplementary feeding.

• Stothart, M. R., Palme, R. & Newman, A. E. M. (2019) It’s what’s on the inside that counts: stress physiology and the bacterial microbiome of a wild urban mammal. Proceedings of the Royal Society B: Biological Sciences 286(1913): 20192111.

The microbiome’s capacity to shape the host phenotype and its mutability underlie theorization that the microbiome might facilitate host acclimation to rapid environmental change. However, when environmental change occurs, it is unclear whether resultant microbiome restructuring is proximately driven by this changing external environment or by the host’s physiological response to this change. We leveraged urbanization to compare the ability of host environment (urban or forest) versus multi-scale biological measures of host hypothalamic–pituitary–adrenal (HPA) axis physiology (neutrophil : lymphocyte ratio, faecal glucocorticoid metabolites, hair cortisol) to explain variation in the eastern grey squirrel (Sciurus carolinensis) faecal microbiome. Urban and forest squirrels differed across all three of the interpretations of HPA axis activity we measured. Direct consideration of these physiological measures better explained greater phylogenetic turnover between squirrels than environment. This pattern was strongly driven by trade-offs between bacteria which specialize on metabolizing digesta versus host-derived nutrient sources. Drawing on ecological theory to explain patterns in intestinal bacterial communities, we conclude that although environmental change can affect the microbiome, it might primarily do so indirectly by altering host physiology. We demonstrate that the inclusion and careful consideration of dynamic, rather than fixed (e.g. sex), dimensions of host physiology are essential for the study of host–microbe symbioses at the micro-evolutionary scale.

• Struhsaker, T. T. (2019) Diet of red-legged sun squirrels (Heliosciurus rufobrachium Waterhouse) in Kibale National Park, Uganda: Implications for seed defence. African Journal of Ecology n/a(n/a).

At end of paper:  “In summary, Hr in Kibale fed on unripe seeds from more species than they did with ripe seeds. Consumption of seeds by Hr in Kibale is most likely limited by a combination of chemical defences and dispersal of ripe fruit by frugivorous birds and primates and not by mechanical defences of fruit or seed. The impact of Hr granivory on tree populations in Kibale is likely minimal compared with that of other species. The observations presented here are intended to encourage research comparing the chemical defenses of unripe versus ripe seeds and how these relate to seed coat hardness. In addition, experimental research would increase our understanding of the physiological adaptations allowing Hr to deal with toxins and anti‐feedants.”

• Suzuki Kei, K. & Ando, M. (2019) Early and efficient detection of an endangered flying squirrel by arboreal camera trapping. Mammalia 83(4): 372.

Abstract: Endangered species management is typically informed by an ecological knowledge of a species. Currently, little is known about the distribution and ecology of the Japanese flying squirrel (Pteromys momonga). To provide an effective rapid survey technique for flying squirrels, we used camera trap surveys and determined what methodology (i.e. camera placement, survey length) was most efficient. We placed 154 cameras in trees for 30 days. We detected flying squirrels at 12% of the camera points. The average suitable distance between camera and targeted tree (DCT) was 130 cm (SE: 15.4, range: 90–220). Moreover, flying squirrels were frequently detected on the trunks of taller trees. We found camera trap surveys were an efficient technique for detecting flying squirrels. Approximately 11% of camera points detected flying squirrels within one survey night. Initial detection of flying squirrels at a site occurred within 10 days at 58% of the points. To efficiently detect flying squirrels, we suggest that it is better to aim the camera towards taller trees at a suitable DCT and to conduct surveys for a minimum of 10 days at each site.

• Thomas, L. S., Teich, E., Dausmann, K., Reher, S. & Turner, J. M. (2018) Degree of urbanisation affects Eurasian red squirrel activity patterns. Hystrix, the Italian Journal of Mammalogy 29(2): 175-180.

With cities growing at a rapid pace, animal species must either retreat to patches of intact natural habitat or adapt to novel conditions in urban areas. While this disturbance causes most species to be in local decline, some show specific behavioural plasticity, facilitating success in a new habitat. The Eurasian red squirrel ( Sciurus vulgaris ) is a common small mammal species which occurs in high numbers in urban environments. To determine which characteristics enable its success, we investigated space use and activity budgets of seven free-ranging individuals living in semi-natural and urban habitats within a large city. We did not find significant differences in animals’ space use between habitat types but tendencies towards smaller home ranges and increased home range overlap existed among individuals in the urban site. Squirrels differed between sites in both overall activity levels and temporal activity patterns: urban animals spent less time active and activity onset was later compared to semi-natural conspecifics. This is likely explained by a combination of dense and reliable supplementary food sources in the urban habitat, reducing foraging effort, and restrictions to movement imposed by higher fragmentation. Flexibility in space use and activity budgets, as well as the ability to exploit anthropogenic food sources and tolerate reduced habitat connectivity, are likely the most important factors contributing to the squirrels’ success in cities. Accordingly, these traits could be used as indicators of low sensitivity towards urbanisation when assessing other species’ potential resilience. However, they do not immunise squirrels against extirpation. Further research on individuals’ foraging ecology and population health may reveal possible threats to urban red squirrels and help predict their future persistence in this challenging habitat.

• Trapp, S. E., Day, C. C., Flaherty, E. A., Zollner, P. A. & Smith, W. P. (2019) Modeling impacts of landscape connectivity on dispersal movements of northern flying squirrels (Glaucomys sabrinus griseifrons). Ecological Modelling 394: 44-52.

Abstract: Landscape connectivity is a key component for successful dispersal of wildlife. Many approaches to quantify landscape connectivity utilize landscape characteristics and wildlife behavior in response to those characteristics, whereas other approaches focus on wildlife dispersal behavior. Combining landscape structure and wildlife behavior (e.g. movement, food acquisition, response to predation risk) provides critical information for conservation and management of many species. Individual-based models (IBM) are useful tools for evaluating the interaction between landscape characteristics and individual behaviors, thereby providing insights into management and conservation. Our objective was to evaluate the effect of reductions in landscape connectivity on northern flying squirrel dispersal movements under alternative timber management scenarios within the modeling framework of a spatially-explicit IBM. We simulated timber harvests of 6% and 9% of old-growth forest, the primary habitat for this species, distributed across 5, 10 or 15 harvest locations. We measured the influence of these scenarios upon landscape connectivity for flying squirrels. Landscapes with greater connectivity exhibited longer flying squirrel dispersal distances, more sinuous dispersal paths, and a greater total area of landscape utilization. However, connectivity was not directly correlated with habitat loss. The harvest scenario with 6% harvest of old growth distributed among 15 harvest locations had the lowest connectivity index value despite other scenarios modeling a greater percentage of old growth loss. The IBM demonstrated the importance of behaviors such as path tortuosity and movement rates in conjunction with landscape configuration in influencing the movement of dispersing individuals. The spatially explicit IBM provided a framework to evaluate connectivity from a fine-scale behavioral rather than structural perspective as well as to evaluate the distribution of new home range locations, which could be a useful management tool when evaluating the influence of landscape heterogeneity and stochastic behavior on wildlife movement and dispersal.

• Travaglia, E., La Morgia, V. & Venturino, E. (2019) Poxvirus, red and grey squirrel dynamics: Is the recovery of a common predator affecting system equilibria? Insights from a predator-prey ecoepidemic model. Discrete and Continuous Dynamical Systems B 22: 1-18.

In Europe, the Eastern grey squirrel is an allochthonous species causing severe impacts on the native red squirrel. The invasive species establishes complex relationships with the native one, out-competing it through resource and disease-mediated competition. However, recent research shed light on the potential role of a predator, the pine marten, in reversing the outcome of the competition between squirrels. Here, we investigate this hypothesis developing a one predator-two prey ecoepidemic model, including disease (squirrel poxvirus) transmission. We assess the equilibria of the dynamical system and investigate their sensitivity to ecosystem parameters changes through numerical simulations.
Our analysis reveals that the system is more likely to evolve toward points where the red squirrels thrive than toward a disease-and-red-squirrels-free point. Although the disease is likely to remain endemic in the system, the introduction of the pine marten destabilizes previous equilibria, favouring the native squirrel and facilitating wildlife managers in their efforts to protect it. Nevertheless, the complete eradication of grey squirrels could be achieved only for specified values of the predation rates and pine marten carrying capacity. The active management of grey squirrel populations remains therefore necessary to try to eradicate the invader from the system.

• Twining, J. P., Montgomery, I., Fitzpatrick, V., Marks, N., Scantlebury, D. M. & Tosh, D. G. (2019) Seasonal, geographical, and habitat effects on the diet of a recovering predator population: the European pine marten (Martes martes) in Ireland. European Journal of Wildlife Research 65(3): 51.

Abstract: Human activity is increasingly altering the natural world. Yet the natural adaptability of most mammal species remains unknown. Seasonal and spatial influences on the diet of temperate carnivores tending towards omnivory are, particularly, poorly understood. The pine marten is one such species which in Ireland and Britain is of additional interest due to the recent recovery in its range and abundance from near collapse. We investigated diet of the pine marten on regional, national, and continental scales and with regard to seasonal and habitat variation. Habitat effects on diet were examined with regard to samples from deciduous woodland, coniferous forestry plantations, heath-coniferous matrices, and mixed habitats. Finally, we discuss the implications of dietary variation in the ecological role of the European pine marten in Ireland and elsewhere and consider how these may be affected by further environmental change. The diet of the pine marten differed significantly amongst all studies across its range, although it maintains the same approximate trophic niche breadth throughout. This plasticity may explain its recovery in an environment where resources are scarce, and underscores its status as an opportunistic species which is likely to be robust to environmental and habitat changes in the future.

• Vitalijus, S. (2019) Ecological aspects of Red Squirrel (Sciurus vulgaris) dreys in city parks. Acta Zoologica Academiae Scientiarum Hungaricae 65(1): 75-84.

Abstract: Modified habitats often pose challenges to native fauna. Red Squirrels (Sciurus vulgaris) successfully inhabits territory of the cities. However research on quite abundant Red Squirrel populations in urban parks of Lithuania was lacking. The investigation was carried out in two city parks (area of Giruliai park 152 ha, Pasakų park forest – 31 ha) in Vilnius, Lithuania. Both areas were mainly coniferous, dominated with pine (Pinus sylvestris). Drey (N = 415) counts were done in February and March 2014–2016. Dreys were found in eight tree species. The Red Squirrels’ dreys were mainly built in most widespread tree species: pine (84.8% of all dreys) and birch (8.1%). Most of the found dreys were built on the side branches at the stem (47.7%). 21.8% of the dreys were built on the branches further from the stem, 18.5% on the top branches and 12% in the stem fork-off. Most of the dreys were set in the southern exposition (33.9%). Average number of red squirrels in both parks in 2014–2016 was 0.20±0.01 squirrels/ha, the average number of dreys – 0.75±0.05 dreys/ha. The population of urban red squirrels in Vilnius city parks can be regarded as stable and having medium abundance, which is supported by supplementary feeding.

• Waterman, J. M. & Archibald, A. J. (2019) Both familiarity and kinship influence odour discrimination by females in a highly social African ground squirrel. Animal Behaviour 148: 145-151.

Abstract: Kin recognition can be important in species where inbreeding avoidance or nepotism (favouritism towards kin) rely on identifying kin, particularly in species with alloparental care. The mechanisms that facilitate kin discrimination, where recognition is determined through cues that correlate with relatedness, usually include either prior association (familiarity) or phenotype matching or both. Odour is an important cue used in a number of mammalian species to discern kin, particularly the ground-dwelling squirrels. Cape ground squirrels, Xerus inauris, are a cooperative-breeding species, living in tight-knit family groups. However, group fission, promiscuity and the large home ranges of breeding males result in a high variance in relatedness both within and among social groups, making them a good species to investigate kin discrimination. We examined whether females are capable of discriminating between the odours of familiar versus unfamiliar females that varied in their relatedness to the focal female using odour experiments. Overall, the average duration of sniffing of the odours of familiar and unfamiliar females did not differ. Similarly, females did not adjust their sniff duration relative to the degree of relatedness of familiar females. However, females appeared to discriminate by the degree of relatedness of unfamiliar (stranger) females, spending longer sniffing odours from females that were not related to them. Thus, females were capable of discriminating the degree of relatedness from odour but they did not do so within their family group. We conclude that Cape ground squirrels are able to discriminate kin. However, whether females in this facultative cooperative breeder use the degree of relatedness in direct social interactions and nepotistic behaviours remains to be investigated.

• Wauters, L. A., Mazzamuto, M. V., Santicchia, F., Van Dongen, S., Preatoni, D. G. & Martinoli, A. (2019) Interspecific competition affects the expression of personality-traits in natural populations. Scientific Reports 9(1): 11189.

Competition between animal species can cause niche partitioning and shape an individual’s phenotype, including its behaviour. However, little is known about effects of interspecific competition on personality, the among-individual variation in behaviour that is consistent across different spatial and temporal contexts. We investigated whether alien grey squirrels (Sciurus carolinensis) influenced the expression of personality traits in native red squirrels (Sciurus vulgaris). In Italy, alien grey squirrels replaced native reds through competition for food resources and space, reducing breeding and recruitment in the native species. We compared personality of red squirrels in red-only (no interspecific competition) and red-grey (with interspecific competition) sites, using arena-tests. The trait activity was measured by Open Field Test while sociability and avoidance were quantified by Mirror Image Stimulation test. Red squirrels co-occurring with the alien species had higher sociability scores and higher between-individual variation in sociability than in red-only sites. Differences in activity and avoidance were not significant. Personality – fitness relationships were not affected by presence or absence of grey squirrels, suggesting that the expression of sociability in red squirrels was not due to short-term selection, but was likely the result of context-related advantages when co-occurring with the competing species.

• Weldy, M., Wilson, T., Lesmeister, D. & Epps, C. (2019) Effects of trapping effort and trap placement on estimating abundance of Humboldt’s flying squirrels. PeerJ 7: e7783.

Live trapping is a common tool used to assess demography of small mammals. However, live-trapping is often expensive and stressful to captured individuals. Thus, assessing the relative tradeoffs among study goals, project expenses, and animal well-being is necessary. Here, we evaluated how apparent bias and precision of estimates for apparent annual survival, abundance, capture probability, and recapture probability of Humboldt’s flying squirrels (Glaucomys oregonensis) varied with the number of secondary trapping occasions. We used data from forested sites trapped on 12 consecutive occasions annually in the HJ Andrews Experimental Forest (9 sites, 6 years) and the Siuslaw National Forest (seven sites, three years) in Oregon. We used Huggins robust design models to estimate parameters of interest for the first 4, 8, and 12 trapping occasions. We also estimated the effect of attaching Tomahawk traps to tree boles on site-and year-specific flying squirrel capture frequencies. Our estimates with 12 occasions were similar to those from previous studies. Abundances and capture probabilities were variable among years on both sites; however, variation was much lower on the Siuslaw sites. Reducing the length of primary trapping occasions from 12 to 8 nights had very little impact on parameter estimates, but further reducing the length of primary trapping occasions to four nights caused substantial apparent bias in parameter estimates and decreased precision. We found that attaching Tomahawk traps to tree boles increased the site-and year-specific capture frequency of flying squirrels. Our results suggest that live-trapping studies targeting Humboldt’s flying squirrels in the Pacific Northwest of the United States could reduce per-site costs and stress to captured individuals without biasing estimates by reducing the length of primary trapping occasions to 8 nights. We encourage similar analyses for other commonly-trapped species in these and other ecosystems.

• Yi, X., Bartlow, A. W., Curtis, R., Agosta, S. J. & Steele, M. A. (2019) Responses of seedling growth and survival to post-germination cotyledon removal: an investigation among seven oak species. Journal of Ecology 0(ja).

Abstract: Rodents regularly rely on emerged epicotyls to locate and remove cotyledons still containing valuable nutrients. However, the extent to which acorn characteristics influence tolerance to post-germination predation has received little attention. Here, we investigated the impact of cotyledon removal following epicotyl emergence on seedling performance and survival of seven oak (Quercus) species. We imitated cotyledon predation at different stages of seedling establishment and development in order to detect effects on seedling height, leaf number, and tissue/component mass. Seedling growth and survival were negatively affected by cotyledon loss regardless of oak species. However, these negative effects decreased as the epicotyl length at which cotyledons were removed increased. We also found that there was a threshold epicotyl length above which seedling survival and performance were relatively unaffected in white oak species compared to red oak species. Following cotyledon removal, early germinating white oak (section Quercus) seedlings survived and/or grew better than the late germinating red oak (section Lobatae) seedlings. This was likely caused by a difference in dependence on cotyledon reserves, which ultimately affected the ability of seedlings to tolerate cotyledon removal. Synthesis. From an evolutionary perspective, this is likely to follow from the early germination in white oaks and the ability of seed consumers to locate young seedlings from the emerging epicotyls. Our study has implications for forest regeneration by suggesting additional opportunities for white oak species to establish following epicotyl emergence. Future studies should consider quantifying the rates of post-germination cotyledon loss.

• Yi, X., Ju, M., Yang, Y. & Zhang, M. (2019) Scatter-hoarding and cache pilfering of rodents in response to seed abundance. Ethology 125(7): 492-499.

Abstract Seed caching and reciprocal cache pilferage play an important role in the coexistence of food-hoarding animals. Understanding what affects seed caching and how cache pilferage occurs is an important question in seed dispersal ecology. However, tracking seed fate and cache pilferage presents substantial practical difficulties. Siberian chipmunks Tamias sibiricus always remove the entire pericarp when scatter-hoarding acorns of Mongolian oak Quercus mongolica, whereas wood mice Apodemus peninsulae often store whole acorns in their caches. These differences in behavior provide an opportunity to investigate unilateral cache pilferage of T. sibiricus from A. peninsulae in response to seed abundance. In this study, tagged acorns were released at the peak and end periods of seed rain from Q. mongolica. This allowed us to investigate seed caching and unilateral cache pilferage at different seed abundances. We found that a higher proportion of acorns were cached at lower level of seed abundance (toward the end of seed rain), mainly because T. sibiricus rather than A. peninsulae scatter-hoarded significantly more acorns at this time. Cache distances decreased with increasing seed abundance, indicating that acorns were cached further away and into smaller caches at lower seed abundance. Unexpectedly, unilateral cache pilferage by T. sibiricus was not significantly influenced by seed abundance?remaining at around 28% during both periods of high and low seed abundance.

2018

• Ancillotto, L., T. Notomista, E. Mori, S. Bertolino and D. Russo (2018) Assessment of Detection Methods and Vegetation Associations for Introduced Finlayson’s Squirrels (Callosciurus finlaysonii) in Italy. Environmental Management.

Abstract: Managing biological invasions requires rapid, cost-effective assessments of introduced species’ occurrence, and a good understanding of the species’ vegetation associations. This is particularly true for species that are elusive or may spread rapidly. Finlayson’s squirrel (Callosciurus finlaysonii) is native to Thailand and southeastern Asia, and two introduced populations occur in peninsular Italy. One of the two introduced populations is rapidly expanding, but neither effective monitoring protocols nor reliable information on vegetation associations are available. To fill this gap, we conducted visual surveys and hair tube sampling in a periurban landscape of southern Italy to compare the effectiveness of these two methods in assessing presence of Finlayson’s squirrel. We also determined the species’ association with vegetation types at detection locations and nesting sites. Both visual and hair tube sampling effectively assessed the species’ presence, but hair tubes resulted in fewer false absences. Moreover, when we controlled for the costs of labor and equipment, hair tubes were 33.1% less expensive than visual sampling. Presence of squirrels and their nests was positively correlated with shrub species richness, indicating that the occurrence of forests with well-developed understory may inhibit the spread of the species.

• Armenta, T. C., Cole, S. W., Geschwind, D. H., Blumstein, D. T. & Wayne, R. K. (2018) Gene expression shifts in yellow-bellied marmots prior to natal dispersal. Behavioral Ecology 30(2): 267-277.

The causes and consequences of vertebrate natal dispersal have been studied extensively, yet little is known about the molecular mechanisms involved. We used RNA-seq to quantify transcriptomic gene expression in blood of wild yellow-bellied marmots (Marmota flaviventer) prior to dispersing from or remaining philopatric to their natal colony. We tested 3 predictions. First, we hypothesized dispersers and residents will differentially express genes and gene networks since dispersal is physiologically demanding. Second, we expected differentially expressed genes to be involved in metabolism, circadian processes, and immune function. Finally, in dispersing individuals, we predicted differentially expressed genes would change as a function of sampling date relative to dispersal date. We detected 150 differentially expressed genes, including genes that have critical roles in lipid metabolism and antigen defense. Gene network analysis revealed a module of 126 coexpressed genes associated with dispersal that was enriched for extracellular immune function. Of the dispersal-associated genes, 22 altered expression as a function of days until dispersal, suggesting that dispersal-associated genes do not initiate transcription on the same time scale. Our results provide novel insights into the fundamental molecular changes required for dispersal and suggest evolutionary conservation of functional pathways during this behavioral process.

• Baldwin, M. K. L., Young, N. A., Matrov, D. & Kaas, J. H. (2018) Cortical projections to the superior colliculus in grey squirrels (Sciurus carolinensis). European Journal of Neuroscience 0(0).

Abstract The superior colliculus is an important midbrain structure involved with integrating information from varying sensory modalities and sending motor signals to produce orienting movements towards environmental stimuli. Because of this role, the superior colliculus receives a multitude of sensory inputs from a wide variety of subcortical and cortical structures. Proportionately, the superior colliculus of grey squirrels is among the largest in size of all studied mammals, suggesting the importance of this structure in the behavioural characteristics of grey squirrels. Yet, our understanding of the connections of the superior colliculus in grey squirrels is lacking, especially with respect to possible cortical influences. In this study, we placed anatomical tracer injections within the medial aspect of the superior colliculus of five grey squirrels (Sciurus carolinensis) and analysed the areal distribution of corticotectal projecting cells in flattened cortex. V1 projections to the superior colliculus were studied in two additional animals. Our results indicate that the superior colliculus receives cortical projections from visual, higher order somatosensory, and higher order auditory regions, as well as limbic, retrosplenial and anterior cingulate cortex. Few, if any, corticotectal projections originate from primary motor, primary somatosensory or parietal cortical regions. This distribution of inputs is similar to the distribution of inputs described in other rodents such as rats and mice, yet the lack of inputs from primary somatosensory and motor cortex is features of corticotectal inputs more similar to those observed in tree shrews and primates, possibly reflecting a behavioural shift from somatosensory (vibrissae) to visual navigation.

• Balmer, A., Zinner, B., Gorrell, J. C., Coltman, D. W., Raveh, S. & Dobson, F. S. (2018) Alternative reproductive tactics and lifetime reproductive success in a polygynandrous mammal. Behavioral Ecology 30(2): 474-482.

The widespread occurrence of alternative reproductive tactics (ARTs) highlights the diverse ways in which sexual selection can operate within a population. We studied ARTs in Columbian ground squirrels (Urocitellus columbianus), evaluating paternity, lifetime reproductive success, and life histories. Reproductively mature male Columbian ground squirrels displayed either a territorial or satellite (nonterritorial) tactic. Territorial males secured a higher proportion of copulations, were more likely to mate at earlier positions in females’ mating sequences, and sired more offspring than satellite males. The tactic males adopted were largely a function of weight, age, and experience, with larger, older, and more experienced males displaying the territorial tactic. At 2 years of age, males adopted the satellite tactic or deferred breeding for the season. Two-year-old satellite males were heavier than males that deferred breeding, but by age 3, there was no difference between their weights. Males that deferred breeding either adopted the satellite or territorial tactic in the following year, with the lighter males displaying the satellite tactic. The ARTs that males adopted at 2 and 3 years of age led to alternative life history pathways. Males that deferred breeding and only adopted the territorial tactic during their lifetime had twice the number of lifetime offspring compared with males that adopted the satellite tactic and then switched to territorial behavior in following years, although this difference was not statistically significant. Our findings show that the satellite tactic results in lower reproductive output both within a single year, except at age 2, and over their lifetime.

• Barbara, F., Morgia, V. L., Parodi, V., Toscano, G. & Venturino, E. (2018) Analysis of the Incidence of Poxvirus on the Dynamics between Red and Grey Squirrels. Mathematics 6(113): 1-21.

Abstract: A model for the interactions of the invasive grey squirrel species as asymptomatic carriers of the poxvirus with the native red squirrel is presented and analyzed. Equilibria of the dynamical system are assessed, and their sensitivity in terms of the ecosystem parameters is investigated through numerical simulations. The findings are in line with both field and theoretical research. The results indicate that mainly the reproduction rate of the alien population should be drastically reduced to repel the invasion, and to achieve disease eradication, actions must be performed to reduce the intraspecific transmission rate; also, the native species mortality plays a role: if grey squirrels are controlled, increasing it may help in the red squirrel preservation, while the invaders vanish; on the contrary, decreasing it in favorable situations, the coexistence of the two species may occur. Preservation or restoration of the native red squirrel requires removal of the grey squirrels or keeping them at low values. Wildlife managers should exert a constant effort to achieve a harsh reduction of the grey squirrel growth rate and to protect the remnant red squirrel population.

Bono, J. M., Pigage, H. K., Wettstein, P. J., Prosser, S. A. & Pigage, J. C. (2018) Genome-wide markers reveal a complex evolutionary history involving divergence and introgression in the Abert’s squirrel (Sciurus aberti) species group. BMC Evolutionary Biology 18(1): 139.

Background
Genetic introgression between divergent lineages is now considered more common than previously appreciated, with potentially important consequences for adaptation and speciation. Introgression is often asymmetric between populations and patterns can vary for different types of loci (nuclear vs. organellar), complicating phylogeographic reconstruction. The taxonomy of the ecologically specialized Abert’s squirrel species group has been controversial, and previous studies based on mitochondrial data have not fully resolved the evolutionary relationships among populations. Moreover, while these studies identified potential areas of secondary contact between divergent lineages, the possibility for introgression has not been tested.

Results
We used RAD-seq to unravel the complex evolutionary history of the Abert’s squirrel species group. Although some of our findings reinforce inferences based on mitochondrial data, we also find significant areas of discordance. Discordant signals generally arise from previously undetected introgression between divergent populations that differentially affected variation at mitochondrial and nuclear loci. Most notably, our results support earlier claims (disputed by mitochondrial data) that S. aberti kaibabensis, found only on the north rim of the Grand Canyon, is highly divergent from other populations. However, we also detected introgression of S. aberti kaibabensis DNA into other S. aberti populations, which likely accounts for the previously inferred close genetic relationship between this population and those south of the Grand Canyon.

Conclusions
Overall, the evolutionary history of Abert’s squirrels appears to be shaped largely by divergence during periods of habitat isolation. However, we also found evidence for interbreeding during periods of secondary contact resulting in introgression, with variable effects on mitochondrial and nuclear markers. Our results support the emerging view that populations often diversify under scenarios involving both divergence in isolation and gene flow during secondary contact, and highlight the value of genome-wide datasets for resolving such complex evolutionary histories.

• Cao, L., Wang, B., Yan, C., Wang, Z., Zhang, H., Geng, Y., Chen, J. & Zhang, Z. (2018) Risk of cache pilferage determines hoarding behavior of rodents and seed fate. Behavioral Ecology 29(4): 984-991.

Cache pilferage by competitors is thought to drive the evolution of hoarding behavior in animals, which plays significant roles in tree regeneration and formation of mutualisms between trees and animals. However, little is known how cache pilferage risk among seeds of different tree species or years affects hoarding behavior and seed dispersal by animals. We hypothesized that scatter-hoarding rodents could adjust hoarding behavior according to variation in cache pilferage risk among seeds and years to minimize cache pilferage, by investigating the relationship between cache pilferage risk and seed dispersal of 7 tree species over 3 years in tropical forest in southwest China. Among years, the high pilferage risk was related to high probability of larder-hoarding and short periods of scatter-hoarding; whereas, the probability of scatter-hoarding was higher in intermediate pilferage year than in both low and high pilferage years. Among seeds, high pilferage risk was related to low probability and short periods of scatter-hoarding. Our results indicated that cache pilferage risk significantly affected hoarding behaviors and seed dispersal by scatter-hoarding rodents as well as seed fates. Cache pilferage risk was a reliable explanatory factor for variation in seed dispersal, and it might be an important driving force in the evolution of rodent hoarding behaviors and seed characteristics.

• Chow, P. K. Y., P. W. W. Lurz and S. E. G. Lea (2018). A battle of wits? Problem-solving abilities in invasive eastern grey squirrels and native Eurasian red squirrels. Animal Behaviour 137: 11-20.

Abstract: Behavioural flexibility has been argued to be an evolutionarily favourable trait that helps invasive species to establish themselves in non-native environments. Few studies, however, have compared the level of flexibility (whether considered as an outcome or as a process) in mammalian invaders and related native species. Here, we tested whether flexibility differs between groups of free-ranging invasive eastern grey squirrels, Sciurus carolinensis, and native Eurasian red squirrels, Sciurus vulgaris, in the U.K., using an easy and a difficult food extraction task. All individuals of both species showed flexibility, at the outcome level, in solving the easy task and solution time was comparable between species across a series of successes. A higher proportion of grey squirrels than red squirrels solved the difficult task. However, for those squirrels that did solve the task, solving efficiency was comparable between species on their first success, and a few red squirrels outperformed the grey squirrels in subsequent successes. Between-species analysis showed that instantaneous flexibility, flexibility at the process level that was measured as the rate of switching between tactics after a failed attempt, was higher in red squirrels than in grey squirrels. Within-species analysis also revealed that red squirrel problem solvers showed higher flexibility at the process level than their nonsolver counterparts. Nonsolvers also failed to make ‘productive’ switches (switching from ineffective to effective tactics). Together, the results suggest that problem-solving ability overlaps in the two species, but is less variable, and on average higher, in grey squirrels than in red squirrels. The superior behavioural flexibility of the grey squirrels, shown here by success at problem solving, may have facilitated their invasion success, but it may also have resulted from selective pressures during the invasion process.

• Davidson, A. D., Hunter, E. A., Erz, J., Lightfoot, D. C., McCarthy, A. M., Mueller, J. K. & Shoemaker, K. T. (2018) Reintroducing a keystone burrowing rodent to restore an arid North American grassland: challenges and successes. Restoration Ecology 26(5): 909-920.

Prairie dogs (Cynomys spp.) are important ecosystem engineers in North America’s central grasslands, and are a key prey base for numerous predators. Prairie dogs have declined dramatically across their former range, prompting reintroduction efforts to restore their populations and ecosystem functions, but the success of these reintroductions is rarely monitored rigorously. Here, we reintroduced 2,400 Gunnison’s prairie dogs (C. gunnisoni) over a period of 6 years to the Sevilleta National Wildlife Refuge, in central New Mexico, U.S.A., a semi-arid grassland ecosystem at the southern edge of their range. We evaluated the population dynamics of prairie dogs following their reintroduction, and their consequent effects on grassland vertebrates. We found postrelease survival of prairie dogs stabilized at levels typical for the species (ca. 50%) after approximately 1 month, while average annual recruitment was ca. 0.35 juveniles per female, well below what was required for a self-sustaining, stable population. Extreme drought conditions during much of the study period may have contributed to low recruitment. However, recruitment increased steadily over time, indicating that the reintroduced colony may simply need more time to establish in this arid system. We also found well-known associates of prairie dog colonies, such as American badgers (Taxidea taxus) and burrowing owls (Athene cunicularia), were significantly more common on the colonies than off. After 7 years, we have yet to meet our goal of establishing a self-sustaining population of Gunnison’s prairie dogs in this semi-arid grassland. But despite the uncertainty and challenges, our work shows that reestablishing keystone species can promote ecosystem restoration.

• Desantis, L. M., Bowman, J., Faught, E., Boonstra, R., Vijayan, M. M. & Burness, G. (2018) Corticosteroid-binding globulin levels in North American sciurids: implications for the flying squirrel stress axis. Canadian Journal of Zoology 96(10): 1090-1096.

Corticosteroid-binding globulin (CBG) helps to regulate tissue bioavailability of circulating glucocorticoids (GCs), and in most vertebrates, ≥80%–90% of GCs bind to this protein. New World flying squirrels have higher plasma total cortisol levels (the primary corticosteroid in sciurids) than most vertebrates. Recent research suggests that flying squirrels have either low amounts of CBG or CBG molecules that have a low binding affinity for cortisol, as this taxon appears to exhibit very low proportions of cortisol bound to CBG. To test whether CBG levels have been adjusted over evolutionary time, we assessed the levels of this protein in the plasma of northern (Glaucomys sabrinus (Shaw, 1801)) and southern (Glaucomys volans (Linnaeus, 1758)) flying squirrels using immunoblotting, and compared the relative levels among three phylogenetically related species of sciurids. We also compared the pattern of CBG levels with cortisol levels for the same individuals. Flying squirrels had higher cortisol levels than the other species, but similar levels of CBG to their closest relatives (tree squirrels). We conclude that CBG levels in flying squirrels have not been adjusted over evolutionary time, and thus, the uncoupling of CBG levels from cortisol concentrations may represent an evolutionary modification in the lineage leading to New World flying squirrels.

• d’Ovidio, D. & Pirrone, F. (2018) A cross-sectional survey to evaluate the pet squirrel population and ownership profiles. Preventive Veterinary Medicine 159: 65-71.

While the presence of squirrels in households is growing, little data is published on their status in captivity. A web-based questionnaire for owners was devised eliciting information about them, their squirrels and their squirrels’ husbandry and health. One hundred owners answered the survey, with most respondents being located in Europe (n = 81). Only data from these respondents were analysed. Twenty-five percents of the owners housed an invasive non-native species of European Union concern (S. carolinensis and T. sibiricus), some of which were younger than three years of age and all but one were sexually intact. This is of particular concern, as the acquisition of these invasive species is illegal since 2015 (European Union Regulation 1143/2014), due to the severe threats they pose to biodiversity. Moreover, escapes derived from improper keeping of intact specimens may augment feral populations or establish new colonies. Among 81 cases, only 5% were neutered, mostly for health reasons. Sixty-three percents of the squirrels had health problems, particularly dermatologic (52%) and intestinal disorders (34%). Most owners reported to visit the veterinarian only if their pet was ill rather than for preventive care. This is the first survey on pet squirrel ownership reported to date. Information that emerges from this study will be useful in implementing rational veterinary strategies for managing pet squirrels properly and, in parallel, meeting the challenges arising from private keeping of alien species.

• Dunham, N. T., McNamara, A., Shapiro, L., Phelps, T., Wolfe, A. N. & Young, J. W. (2019) Locomotor kinematics of tree squirrels (Sciurus carolinensis) in free-ranging and laboratory environments: Implications for primate locomotion and evolution. Journal of Experimental Zoology Part A: Ecological and Integrative Physiology 331(2): 103-119.

Abstract: The grasping capabilities and gait kinematics characteristic of primates are often argued to be adaptations for safely moving on small terminal branches. The goal of this study was to identify whether Eastern gray squirrels (Sciurus carolinensis)?arboreal rodents that frequently move and forage on small branches, lack primate-like grasping and gait patterns, and arguably represent extant analogs of a stem primate ancestor?adjust gait kinematics to narrow and nonhorizontal branches. We studied locomotor kinematics of free-ranging and laboratory-housed squirrels moving over various substrates. We used high-speed video to film (a) a population of free-ranging squirrels moving on natural substrates and (b) laboratory-housed squirrels moving on horizontal poles. Substrates were coded as small, medium, or large relative to squirrel trunk diameter, and as inclined, declined, or horizontal. Free-ranging squirrels used more gallops and half-bounds on small- and medium-sized substrates, and more high-impact bounds, with reduced limb-lead durations, on declined substrates. Laboratory squirrels moved at higher speeds than free-ranging squirrels and responded to decreasing diameter by using more gallops and half-bounds, lowering speed, and?controlling for speed?increasing mean duty factor, mean number of supporting limbs, and relative forelimb lead duration. Our inability to detect substantial diameter or orientation-related gait adjustments in the wild may be due to a limited accounting of confounding influences (e.g., substrate compliance). Ultimately, studies assessing stability measures (e.g., center of mass fluctuations and peak vertical force) are required to assess whether primates’ enhanced grasping and gait patterns engender performance advantages on narrow or oblique substrates.

• Everest DJ, Shuttleworth CM, Grierson SS, Dastjerdi A, Stidworthy MF, Duff JP, et al. (2018) The implications of significant adenovirus infection in UK captive red squirrel (Sciurus vulgaris) collections: How histological screening can aid applied conservation management. Mammalian Biology 88:123-9.

Abstract: Conservation trans-locations using captive bred red squirrels (Sciurus vulgaris) is increasing in the United Kingdom (UK). However, project managers are often unaware of the risk of pathological adenovirus (ADV) infection. In this study we illuminate the viral threat using transmission electron microscopy (TEM) and polymerase chain reaction (PCR) assays. Both techniques were used to screen samples collected from 26 English and Welsh captive red squirrel collections. Of 181 carcasses received between 2002 and 2016, 129 (71%) were suitable for routine surveillance post mortem examination (PME). A range of tissues were examined with ADV identified from a variety of samples by PCR and TEM in 92 (72%) cases encompassing 23 of the 26 study collections (89%). ADV enteritis was histologically confirmed in two deaths (2%) with another 39 (30%) through both laboratory and clinical findings, considered as likely clinically-significant ADV cases, but advanced autolysis precluded accurate assessment and confirmatory histological diagnosis. Other positive cases were more indicative of sub-clinical infection. Clusters of ADV red squirrel deaths were recorded with circumstantial evidence suggesting inter-collection movement of presumed ADV infected donated animals had triggered mortality in recipient collections. During the study, several collections intermittently experienced ADV-associated deaths. Definitive cause of death was not determined in most cases, but a diverse range of diagnoses were recorded in 25 (19%) animals. Implications of these findings for captive United Kingdom (UK) red squirrel husbandry are discussed. It is recommended that protocols be drawn up to minimise potential intra-species ADV infection and highlight the danger of contact with ADV infected wood mice (Apodemus sylvaticus).

• Goldstein, E.A., M.J. Merrick & J.L. Koprowski (2018) Low survival, high predation pressure present conservation challenges for an endangered endemic forest mammal. Biological Conservation 221: 67-77.

Abstract: Knowledge of which population parameters and mortality risks contribute most to population decline and endangerment is necessary to develop informed and actionable conservation plans for threatened and endangered species (Rushton et al., 2006). The federally endangered Mount Graham red squirrel (Tamiasciurus fremonti grahamensis) is restricted to the Pinaleño Mountains, in southeastern Arizona, USA. The population is critically threatened with extensive habitat loss from fire as well as by an introduced non-native squirrel species, the Abert’s squirrel (Sciurus aberti). Recovery is challenged by low survival and poor reproduction, such that the subspecies is functionally semelparous. We calculate survival rates and cause-specific mortality hazards from known-fate individuals to understand the impact of predation on survival and demography in this peripheral population. We document the lowest survival and highest rates of mortality in any population of North American red squirrels in both adult and juvenile age classes (mean annual survival: adults = 0.32, juveniles = 0.26). We attributed the majority of confirmed deaths to avian predation (adults 65%, juveniles 75%), and the daily hazard rate for avian predation was 15 times higher than for mammalian predation and 2 times higher than death from unknown causes. It is likely that the presence of an ecologically similar, non-native tree squirrel subsidizes a diverse avian predator guild, which includes two raptor species of conservation concern. In addition to efforts to remove the non-native Abert’s squirrel, we recommend immediate forest restoration efforts in the long term, and habitat augmentation to increase structural complexity, cover, shelter, and food resources in the short term.

• Hämäläinen, S., Fey, K. & Selonen, V. (2018) Habitat and nest use during natal dispersal of the urban red squirrel (Sciurus vulgaris). Landscape and Urban Planning 169: 269-275.

As urban environments differ from the natural environment, the ability of a species to move in and use variable land composition types determines its fate in the urban environment. In many mammalian species, the selection of home ranges mainly occurs during natal dispersal. Thus, habitat selection of juvenile individuals greatly determines where animals are found in the city. Here, our goal is to understand how an originally forest-dwelling rodent selects components of an urban habitat. We used radio telemetry to record the habitat and nest use of Eurasian red squirrels (Sciurus vulgaris) in the city of Turku in southwestern Finland, with the main focus on habitat use during natal dispersal. We found that although the red squirrels preferred areas with more trees available than in the surrounding urban landscape they nevertheless often used sites with only a few trees. This result was highlighted by the analysis of the home range scale, as the area used did not differ greatly from the habitat composition of the available landscape. Juvenile red squirrels used the deciduous land cover type more often during movements in the natal area than during dispersal and explorative movements; however, they also settled in areas with less deciduous land cover type than in the natal area. Our results show that in urban areas movements of red squirrels are not restricted to their natural habitat type, which is a coniferous forest, but that they seem to be well adapted to urban areas, being able to utilize urban structures.

• Hammond, T. T., Palme, R. & Lacey, E. A. (2018) Ecological specialization, variability in activity patterns and response to environmental change. Biology Letters 14(6).

Differences in temporal patterns of activity can modulate the ambient conditions to which organisms are exposed, providing an important mechanism for responding to environmental change. Such differences may be particularly relevant to ecological generalists, which are expected to encounter a wider range of environmental conditions. Here, we compare temporal patterns of activity for partially sympatric populations of a generalist (the lodgepole chipmunk, Tamias speciosus) and a more specialized congener (the alpine chipmunk, Tamias alpinus) that have displayed divergent responses to the past century of environmental change. Although mean activity budgets were similar between species, analyses of individual-level variation in locomotion revealed that T. alpinus exhibited a narrower range of activity patterns than T. speciosus. Further analyses revealed that T. alpinus was more active earlier in the day, when temperatures were cooler, and that activity patterns for both species changed with increased interspecific co-occurrence. These results are consistent with the greater responsiveness of T. alpinus to changes in environmental conditions. In addition to highlighting the utility of accelerometers for collecting behavioural data, our findings add to a growing body of evidence, suggesting that the greater phenotypic variability displayed by ecological generalists may be critical to in situ responses to environmental change.

• Hanmer, H. J., Thomas, R. L. & Fellowes, M. D. E. (2018) Introduced Grey Squirrels subvert supplementary feeding of suburban wild birds. Landscape and Urban Planning 177: 10-18.

Abstract: Providing food for wild birds is perhaps the most widespread intentional interaction between people and wildlife. In the UK, almost half of households feed wild birds, often as peanuts and seed supplied in hanging feeders. Such food is also taken by the introduced, invasive Grey Squirrel Sciurus carolinensis. Little is known of how Grey Squirrels utilise this resource and how they affect feeder use by wild birds. To assess this we recorded the numbers and time spent by animals visiting experimental feeding stations in suburban gardens, and also asked if exclusionary guards (to prevent Grey Squirrel access), food type (peanut, mixed seed), habitat and weather conditions influenced visits. Using automated cameras, we recorded 24,825 bird and 8577 Grey Squirrel visits. On average >44% of the time feeders were utilised, they were being visited by Grey Squirrels. Grey Squirrel presence prevented birds from feeding at the same time (>99.99%). Feeders where Grey Squirrels were dominant were less likely to be visited by birds, even in their absence. Guards reduced Grey Squirrel use to a minimum on seed feeders, and by approximately half on peanut feeders. Squirrels, food type, guard status, habitat and rainfall all influenced bird activity and timing of feeder visits. Our work suggests that Grey Squirrels reduce the availability of supplementary food to wild birds, while gaining large volumes of food resources with corresponding benefits. Given the ubiquity of supplementary feeding, it is likely that this is an important resource for urban Grey Squirrels; feeder guards mitigate this effect.

• Lane, J., McAdam, A., McFarlane, E., Williams, C., Humphries, M., Coltman, D., Gorrell, J. & Boutin, S. (2018) Phenological shifts in North American red squirrels: disentangling the roles of phenotypic plasticity and microevolution. Journal of Evolutionary Biology

Abstract: Phenological shifts are the most widely reported ecological responses to climate change, but the requirements to distinguish their causes (i.e., phenotypic plasticity versus microevolution) are rarely met. To do so, we analyzed almost two decades of parturition data from a wild population of North American red squirrels (Tamiasciurus hudsonicus). Although an observed advance in parturition date during the first decade provided putative support for climate change‐driven microevolution, a closer look revealed a more complex pattern. Parturition date was heritable (h2 = 0.14 (0.07 to 0.21 (HPD interval)) and under phenotypic selection (β = ‐0.14 ± 0.06 (SE)) across the full study duration. However, the early advance reversed in the second decade. Further, selection did not act on the genetic contribution to variation in parturition date, and observed changes in predicted breeding values did not exceed those expected due to genetic drift. Instead, individuals responded plastically to environmental variation, and high food (white spruce (Picea gluaca) seed) production in the first decade appears to have produced a plastic advance. In addition, there was little evidence of climate change affecting the advance, as there was neither a significant influence of spring temperature on parturition date or evidence of a change in spring temperatures across the study duration. Heritable traits not responding to selection in accordance with quantitative genetic predictions has long presented a puzzle to evolutionary ecologists. Our results on red squirrels provide empirical support for one potential solution: phenotypic selection arising from an environmental, as opposed to genetic, covariance between the phenotypic trait and annual fitness.

McInnes, C. (2018) Why do red squirrels die? Veterinary Record 183(16): 500.

There has been considerable recent interest in why free-living red squirrels (Sciurus vulgaris) (Fig 1) in the British Isles die. The reason for this is that the red squirrel population in Great Britain has been in noticeable decline since the early 20th century, which Middleton reported was almost certainly due to epidemics of disease. Although he was only able to name coccidiosis with any certainty, Middleton felt that other causes, some producing external signs of disease, were also responsible for red squirrel mortalities. He even ventured the suggestion that introduced grey squirrels (Sciurus carolinensis) may carry a disease to which they themselves were immune, but to which the red squirrels were highly susceptible. He did concede, however, that disease epidemics in red squirrels had been reported not only in parts of the country where grey squirrels were found, but also in areas where grey squirrels had yet to be seen. Fig 1: Red squirrel populations in Great Britain have been in noticeable decline for the past 100 years. Although there had been reports of grey squirrels in the Welsh countryside as early as 1830, the best documented introductions of grey squirrels from America occurred between 1876 and 1910. By the 1940s it was realised that grey squirrels had expanded considerably from their points of introduction and that in areas of England and Wales where the grey squirrels had been resident for 15 years or more there were no red squirrels. As the grey squirrel population …

• Mielke, F., Schunke, V., Wölfer, J. & Nyakatura, J. A. (2018) Motion analysis of non-model organisms using a hierarchical model: Influence of setup enclosure dimensions on gait parameters of Swinhoe’s striped squirrels as a test case. Zoology 129: 35-44.

In in-vivo motion analyses, data from a limited number of subjects and trials is used as proxy for locomotion properties of entire populations, yet the inherent hierarchy of the individual and population level is usually not accounted for. Despite the increasing availability of hierarchical model frameworks for statistical analyses, they have not been applied extensively to comparative motion analysis. As a case study for the use of hierarchical models, we analyzed locomotor parameters of four Swinhoe’s striped squirrels. The small-bodied arboreal mammals exhibit brief bouts of rapid asymmetric gaits. Spatio-temporal parameters on runways with experimentally varied dimensions of the setup enclosure were compared to test for their potentially confounding effects. We applied principal component analysis to evaluate changes to the overall locomotor pattern. A common, non-hierarchical, pooled statistical analysis of the data revealed significant differences in some of the parameters depending on enclosure dimensions. In contrast, we used a hierarchical Bayesian generalized linear model (GLM) that considers subject specific differences and population effects to compare the effect of enclosure dimensions on the measured parameters and the principal components. None of the population effects were confirmed by the hierarchical GLM. The confounding effect of a single subject that deviates in its locomotor behavior is potentially bigger than the influence of the experimental variation in enclosure dimensions. Our findings justify the common practice of researchers to intuitively select an enclosure with dimensions assumed as “non-constraining”. Hierarchical models can easily be designed to cope with limited sample size and bias introduced by deviating behavior of individuals. When limited data is available—a typical restriction of in-vivo motion analyses of non-model organisms—density distributions of the Bayesian GLM used here remain reliable and the hierarchical structure of the model optimally exploits all available information. We provide code to be adjusted to other research questions.

• Mori, E., Menchetti, M., Zozzoli, R. & Milanesi, P. (2018) The importance of taxonomy in species distribution models at a global scale: the case of an overlooked alien squirrel facing taxonomic revision. Journal of Zoology 0(0).

Abstract The Siberian chipmunk is native to north-eastern Asia, but alien populations of this squirrel, introduced through the pet trade, occur in many European countries. This rodent has been listed as an invasive species of European concern, being a potential vector of ticks spreading Lyme disease. We aimed to assess its current distribution range and to identify areas of potential invasion. Two sets of species distribution models were conducted, one considering the locations of the species (n = 625 occurrences) and the other with only the occurrences of the Korean subspecies (the invasive one in Europe; n = 255 occurrences), which might be a separate species from the Siberian one. We included 19 uncorrelated predictors (two topographic, nine land cover, five bioclimatic and three anthropogenic variables), which may represent the habitat characteristics of the target species. Most of the northern hemisphere supports the establishment of the Siberian chipmunk, particularly for the invasive Korean subspecies (especially in Europe, where it is already established), mostly in urban areas. Anthropogenic food supply was found to be an important factor promoting the growth of alien populations of chipmunks, whereas the presence of the native red squirrel at the time of introduction may limit it.

• Mori, E., Zozzoli, R. & Menchetti, M. (2018) Global distribution and status of introduced Siberian chipmunks Eutamias sibiricus. Mammal Review 48(2): 139-152.

Abstract: * Among invasive alien species, squirrels are prominent because of their popularity as pets and their positive perception by the general public. In Europe, populations of five alien squirrel species are reported. The pet trade represents a high-risk pathway for the introduction of rodents, which are likely to become invasive because of their reproductive biology and wide native distribution ranges. In the European Union, a trade ban has been imposed on some particularly impacting species, such as the eastern grey squirrel Sciurus carolinensis. Eradication and numerical control is needed for most introduced squirrels in Europe (including the Siberian chipmunk), according to European Union Regulation 1143/2014 on alien species. * We summarise the current distribution of the ground-dwelling Siberian chipmunk Eutamias sibiricus, one of the most common pet squirrels in Europe and Asia, in its invasive range. * Established populations of Siberian chipmunk, in most cases composed of small numbers of individuals, currently occur in France, Switzerland, Italy, Germany, Belgium, the Netherlands, Denmark, and Russia. The species was present in Austria, Sweden, and Hong Kong, and unsuccessful introductions occurred in Spain and the British Isles. The first records from Greece are also reported. Most introductions took place during the 1990s, when individuals were intentionally released (67% of extant populations result from such releases). Population size was correlated with the proximity to urban areas. * Human risk of infection with the agent of Lyme borreliosis seems to be the highest where populations of alien Siberian chipmunk occur. Other impacts have never been reported. Competition with native rodents (e.g. the red squirrel Sciurus vulgaris and dormice, Gliridae) has been speculated on, as well as predation on nests of native warblers (Sylvidae). * Impacts by alien Siberian chipmunks have been overlooked for a long time. Despite this, given that the species may affect native biodiversity and human health, eradication of established populations is recommended.

• Perkins-Taylor, I. E. & Frey, J. K. (2018) Ecological factors associated with site occupancy of an endemic chipmunk. The Journal of Wildlife Management 82(7): 1466-1477.

Abstract: The Oscura Mountains Colorado chipmunk (Neotamias quadrivittatus oscuraensis) is a rare subspecies of the Colorado chipmunk that is listed as threatened by the state of New Mexico, primarily because it is an endemic subspecies with a small, isolated habitat and the potential for continuing habitat loss. Knowledge about its ecology is limited, which has hindered the development of scientifically defensible management plans. Our goal was to better understand the ecological factors related to the distribution and habitat selection of this chipmunk. We deployed baited camera traps in the Oscura Mountains, New Mexico, USA, using an occupancy modeling framework to determine which ecological factors are associated with occupancy and detection probability. We collected microhabitat and landscape-level data for use as covariates in the occupancy models. We detected the Oscura Mountains chipmunk at 26 of the 137 survey sites. Occupancy probability was not influenced by the microhabitat characteristics measured and the final model contained only landscape-level covariates on occupancy. Probability of occupancy was positively associated with proximity to an escarpment, two-needle pinyon (Pinus edulis) woodland vegetation community type, and elevation. Detection probability was positively associated with the presence of mature two-needle pinyons at the site. Habitat loss is a major concern for this taxon, especially because climate change is expected to exacerbate threats to pinyon woodlands. Drought, wildfire, bark beetle (Ips confusus) outbreaks, and other diseases pose a large risk to conifer woodlands throughout the southwestern United States. Conserving the pinyon woodlands on the Oscura Mountains, particularly in areas near escarpments, will help maintain habitat that is important for the Oscura Mountains chipmunk.

• Pero, E. M. & Hare, J. F. (2018) Costs of Franklin’s ground squirrel (Poliocitellus franklinii) ectoparasitism reveal adaptive sex allocation. Canadian Journal of Zoology 96(6): 585-591.

Parasite infestation may impose direct costs of blood, nutrient, and energy depletion, along with indirect costs of increased immune response upon hosts. We investigated how ectoparasitism influences body mass and reproduction in a free-living population of Franklin’s ground squirrels (Poliocitellus franklinii (Sabine, 1822)) located near Delta Marsh, Manitoba, Canada. We experimentally reduced ectoparasite burden by treating seven reproductive females with an insecticide following breeding and contrasted body mass and reproductive performance of those individuals to seven sham-treated control females. Insecticide-treated dams did not differ from sham-treated dams in body mass, litter size, or juvenile mass, and thus, dam growth and reproduction were not compromised by ectoparasite defense at the infestation levels experienced in this study. However, litter sex ratio differed significantly between insecticide-treated and control females, with a higher proportion of male offspring produced among females with reduced ectoparasite load. Our findings are thus consistent with the Trivers–Willard model for adaptive sex allocation, yet they provide novel comparative insight into how sociality may modulate the expression of adaptive sex allocation among small mammals given the differential payoff associated with the production of high-quality female versus male offspring in more social versus less social species.

• Pierre-Henri, F., Marie-Ka, T., Christiane, D., Philippe, G., Violaine, N., Emmanuel JP, D. & Laurent, M. (2018) Flightless scaly-tailed squirrels never learned how to fly: A reappraisal of Anomaluridae phylogeny. Zoologica Scripta 0(0).

Anomaluroidea, commonly known as the “scaly-tailed squirrels,” are an emblematic group of tropical African mammals that includes gliding forms. The family Anomaluridae was until recently represented by three genera: the flying scaly-tailed squirrels (Anomalurus), the flying mouse (Idiurus) and the flightless scaly-tailed squirrels (Zenkerella). Idiurus and Zenkerella have long been grouped into the Zenkerellinae subfamily, and Zenkerella was interpreted as a rare case of evolutionary reversal to non-gliding lifestyle. Recent studies have demonstrated that Zenkerella is sister to all other modern anomalurids, and represents in fact the monogeneric family Zenkerellidae. The Anomalurus genus was split into Anomalurus and Anomalurops, but no study has ever considered all Anomalurus species together in a phylogeny to test the status of Anomalurops. Here, we used mitogenomic next-generation sequencing to infer the phylogenetic relationships among all extant anomalurids and to estimate their divergence ages. We found that the arboreal Zenkerella is the sister group of all extant gliding anomalurids (Idiurus and Anomalurus). We confirmed that Anomaluroidea only evolved the gliding adaptation once. A comparison based on morphological traits indicates that Zenkerella harbours several unique morphological features. We propose new morphological characters for the novel classification of modern Anomaluroidea, which includes the families Zenkerellidae and Anomaluridae. Using different calibration schemes, we demonstrated that classical dating methods relying only on mitogenomes provide rather young Miocene estimates between Zenkerellidae and the Anomaluridae. The use of published nuclear genes, internal calibrations and tip dating converged towards an Eocene split between gliding and non-gliding scaly-tailed squirrels, which is in agreement with the African fossil record. Finally, we provide the first exhaustive species-level molecular phylogenetic inference for the genus Anomalurus. We found that Anomalurus beecrofti is the sister group of all other species of Anomalurus and branched off during the Miocene.

• Romeo, C., McInnes, C. J., Dale, T. D., Shuttleworth, C., Bertolino, S., Wauters, L. A. & Ferrari, N. (2018) Disease, invasions and conservation: no evidence of squirrelpox virus in grey squirrels introduced to Italy. Animal Conservation 0(0).

Abstract Native red squirrels Sciurus vulgaris in Great Britain and Ireland are threatened by alien grey squirrels S. carolinensis through exploitation competition and spillover of squirrelpox virus (SQPV). By accelerating the replacement of red squirrels by the invader, SQPV represents a fundamental factor to consider when planning management and conservation strategies. In mainland Europe, grey squirrels introduced to Italy threaten the survival of the whole continental red squirrel population, but no extensive surveys for SQPV presence have been carried out in the region. We therefore investigated SQPV infection in north Italian grey squirrel populations through a combination of serological and molecular methods. Firstly, we analysed sera from 285 individuals through an enzyme-linked immunosorbent assay (ELISA) to detect antibodies against SQPV. Secondly, a PCR designed to amplify a segment of the G8R SQPV gene was carried out on DNA extracted from swabs and skin tissue samples from a second set of 66 grey squirrels. ELISA tests identified four reactors (1.4%), but the subsequent PCR survey did not detect any SQPV DNA. Based on the low prevalence observed and on PCR results, we believe that the four suspected positives were the result of an ELISA cross-reaction following exposure to another pox virus. Considering sample size and performances of the two methods, confidence of freedom from SQPV resulted above 99.9%. However, because of the severe impact of SQPV on red squirrels, we recommend the implementation of a passive surveillance plan for the early detection of an SQPV emergence in continental Europe.

• Santicchia, F., Dantzer, B., van Kesteren, F., Palme, R., Martinoli, A., Ferrari, N. & Wauters, A. L. (2018) Stress in biological invasions: introduced invasive grey squirrels increase physiological stress in native Eurasian red squirrels. Journal of Animal Ecology .

Abstract: 1.Invasive alien species can cause extinction of native species through processes including predation, interspecific competition for resources, or disease‐mediated competition. Increases in stress hormones in vertebrates may be associated with these processes and contribute to the decline in survival or reproduction of the native species. 2.Eurasian red squirrels (Sciurus vulgaris) have gone extinct across much of the British Isles and parts of Northern Italy following the introduction of North American invasive grey squirrels (Sciurus carolinensis). We extracted glucocorticoid metabolites from faecal samples to measure whether the presence of the invasive species causes an increase in physiological stress in individuals of the native species. 3.We show that native red squirrels in seven sites where they co‐occurred with invasive grey squirrels had glucocorticoid concentrations that were three times higher than those in five sites without the invasive species. Moreover, in a longitudinal study, stress hormones in native red squirrels increased after colonisation by grey squirrels. When we experimentally reduced the abundance of the invasive grey squirrels, the concentration of faecal glucocorticoid metabolites in co‐occurring red squirrels decreased significantly between pre‐ and post‐removal periods. 4.Hence, we found that the invasive species acts as a stressor which significantly increases the concentrations of glucocorticoids in the native species. 5.Given that sustained elevations in glucocorticoids could reduce body growth and reproductive rate, our results are consistent with previous studies where the co‐occurrence of the invasive grey squirrel was associated with smaller size and lower reproductive output in red squirrels.

• Selonen, V. & S. Mäkeläinen (2018) Ecology and protection of a flagship species, the Siberian flying squirrel. Hystrix, the Italian Journal of Mammalogy 2018. 28(2).

Abstract: Having clear ecological knowledge of protected species is essential for being able to successfully take actions towards conservation, but this knowledge is also crucial for managing and preventing conservation conflicts. For example, the Siberian flying squirrel, Pteromys volans , listed in the EU Habitats Directive and inhabiting mature forests that are also the target for logging, has had a major role in political discussions regarding conservation in Finland. This species has also been well-researched during recent decades, providing knowledge on the ecology and management of the animal. Herein, we review knowledge on habitats, demography, community interactions and spatial ecology of this flagship species. We compare the ecology of flying squirrels with that of other arboreal squirrels, and summarize conservation management and policy related to flying squirrels. Reviewed research on the Siberian flying squirrel shows that the species has many similarities in behaviour to other arboreal squirrels. For instance, arboreal squirrels deviate from the general pattern of male-biased sexual size dimorphism in mammals, which perhaps relates to the mating system of arboreal squirrels. Important differences are found in the response of Siberian flying squirrels to tree mast, i.e. pulsed food resource, compared to that of red squirrels, and in communal nesting behaviour compared to that of North American flying squirrels. The extensive knowledge on dispersal behaviour of the flying squirrel, well-studied habitat associations and the proved need for evidence-based conservation may guide researchers and managers working with other similar species. For conservation, the case of the Siberian flying squirrel demonstrates that habitat protection becomes both ineffective and uneconomical if ecological knowledge is not applied in the conservation planning process. The cost-effective conservation of the species requires both landscape-level conservation planning and flexible conservation options to increase the motivation of land owners for conservation.

• Sheehy E, Sutherland C, O’Reilly C, Lambin X. (2018) The enemy of my enemy is my friend: native pine marten recovery reverses the decline of the red squirrel by suppressing grey squirrel populations. Proceedings of the Royal Society B: Biological Sciences 285(1874).

Abstract: Shared enemies may instigate or modify competitive interactions between species. The dis-equilibrium caused by non-native species introductions has revealed that the outcome of such indirect interactions can often be dramatic. However, studies of enemy-mediated competition mostly consider the impact of a single enemy, despite species being embedded in complex networks of interactions. Here, we demonstrate that native red and invasive grey squirrels in Britain, two terrestrial species linked by resource and disease-mediated apparent competition, are also now linked by a second enemy-mediated relationship involving a shared native predator recovering from historical persecution, the European pine marten. Through combining spatial capture–recapture techniques to estimate pine marten density, and squirrel site-occupancy data, we find that the impact of exposure to predation is highly asymmetrical, with non-native grey squirrel occupancy strongly negatively affected by exposure to pine martens. By contrast, exposure to pine marten predation has an indirect positive effect on red squirrel populations. Pine marten predation thus reverses the well-documented outcome of resource and apparent competition between red and grey squirrels.

• Shirkorshidi, M. & Tonkin, M. (2018) Saving Scotland’s Red Squirrels Results of Spring 2017 Squirrel Surveys. Saving Scotland’s Red Squirrels.

Abstract: We present the results of Saving Scotland’s Red Squirrels’ (SSRS) Spring 2017 surveys of red and grey squirrels and compare them with the Spring 2011 and 2012 surveys for the north of Scotland and Spring 2013 for the south of Scotland. We also present the results of additional intensive grey squirrel detection surveys in North East Scotland. Overall, we see a picture of red squirrels maintaining or increasing their occurrence since the beginning of the surveys, but a mixed picture for the occurrence of grey squirrels: a positive change towards decreased grey squirrel range for the northern half of Scotland contrasts with expanding grey squirrel distribution in South Scotland. Taking the regions in turn, the evidence supports a statistically significant increase in red squirrel distribution and a significant decline in grey squirrels in the North East, particularly in areas close to the City of Aberdeen. Across the Highland Line the trend shows a stabilisation of red squirrel occurrence in survey areas, and grey squirrel occurrence limited to the significantly reduced range when compared with the tetrads in 2011 and 2012 Thus we have been successful in preventing incursion of grey squirrels into Scotland’s core red squirrel populations to the north of the Highland Line and protecting. In the south of Scotland, overall, grey squirrel occupancy is shown to have increased since 2014, especially in Berwickshire, where red squirrels are now seldom seen, and, more worryingly, in Dumfriesshire between Thornhill and New Galloway. We believe this to be associated with an exceptional masting year in the autumn of 2014, when there was a superabundance of beech nuts. This led to increased over-winter survival and a peak in production of grey squirrels in 2015, which was evident in our 2015 trapping figures as a surge in grey squirrel numbers right across the country. Despite this, red squirrels appear to have maintained their range and even slightly expanded their occupancy in South Scotland, although the change is not statistically significant.

• Shuttleworth, C. & Halliwell, E. (2018) Red Squirrels In My Garden. European Squirrel Initiative, Stoneleigh Park, Warwickshire, CV8 2LG England: 120 pp.

• Starkey, A. & delBarco-Trillo, J. (2018) Vienna Supplementary feeding can attract red squirrels (Sciurus vulgaris) to optimal environments. Mammalian Biology

Abstract: A number of conservation approaches are used to manage threatened species. However, some of these approaches require intensive planning and can often be restricted by funding. Supplementary feeding is a non-invasive and cost-effective approach to manage vulnerable populations, but we lack data on its usefulness. Here we investigated the effects of supplementary feeding on a population of red squirrels (Sciurus vulgaris), a UK priority species which faces competition from the non-native grey squirrel (Sciurus carolinensis). The study took place October-December 2015, lasting 8 weeks. Twenty feeders were installed 1 week prior to the beginning of the study in a protected woodland free from grey squirrels, either containing food (full feeders) or no food (empty feeders), and squirrel abundance before and after feeding was recorded at each feeder (for a total of 27 feeding and recording events). Six times more squirrels were seen at full feeders, and numbers increased by 7 fold after feeding. We also observed that the activity of red squirrels in the vicinity of full feeders increased during the course of the study. Eighty-five hair samples were collected during the study, all of which were found at full feeders. Results demonstrate red squirrels can differentiate between full and empty feeders, suggesting their awareness increases when supplementary food is present. Increased abundance of squirrels at full feeders after feeding times not only implies that squirrels are attracted to and can benefit by supplementation, it also shows that food supplementation can be used to regulate the movement of individuals across habitats. Understanding how red squirrel populations are affected by supplementary feeding will contribute towards existing conservation efforts to improve this species future survival.

• Stuart, K. D., Vander Haegen, W. M., Jenkins, K. J., Keren, I. N. & West, S. D. (2018) Western gray squirrel resource selection related to fire fuel management. The Journal of Wildlife Management 0(0).

Abstract: One of 3 populations of the state-threatened western gray squirrel (Sciurus griseus) in Washington occurs in the northern Cascade Range (i.e., North Cascades), where long-term fire suppression has increased the risk of catastrophic wildfire. Land management agencies throughout this region have implemented fire fuel reduction programs that alter squirrel habitat and may affect their populations. From April 2008 to September 2011, we investigated resource selection of 38 radio-collared western gray squirrels at 2 study sites in the North Cascades following fire fuel management activities including mechanical thinning and prescribed burning. We developed conditional logistic models to examine resource selection at 3 spatial scales: nest trees, nest sites, and core areas within home ranges. The odds of a squirrel selecting a tree for nesting increased with dwarf mistletoe (Arceuthobium spp.) presence, greater number of surrounding trees with interlocking branches, and tree size. Squirrels selected nest sites that had greater canopy cover, tree connectivity, and presence of dwarf mistletoe than available, unused sites. Core-use areas within home ranges had greater canopy cover, a greater number of tree species, and trees with higher live crowns compared to low-use areas. Our results indicate that fire fuel treatments may negatively affect western gray squirrel habitat across multiple spatial scales. Most variables that were positively related to habitat selection are specifically targeted for reduction in fire fuel management plans and were lower in sampled treated areas compared to untreated areas within the study sites. Key considerations in designing fuel reduction programs that benefit both squirrel habitat conservation and fire fuel management include maintaining forest patches with suitable canopy cover and connectivity, retaining large trees of a mix of species, and allowing for mistletoe infection at a reduced rate. © 2018 The Wildlife Society.

• Tamura, N., Boonkhaw, P., Prayoon, U., Kanchanasaka, B. & Hayashi, F. (2018) Mating calls are a sensitive indicator of phylogenetic relationships in Callosciurus, tropical tree squirrels. Mammalian Biology.

Abstract: Vocal communication plays an important role in the mating behaviour of arboreal squirrels. Callosciurus is a genus of tree squirrels that includes 15 species distributed in Southeast Asia, and congeneric species often inhabit the same forest. As closely related species of Callosciurus have the potential to interbreed, species recognition from mating calls may be a fundamental reproductive barrier. We compared seven acoustic characteristics of male mating calls in six Callosciurus species and estimated whether the species differences were clear enough to function as a cue. Discriminant function analyses (DFA) classified 87.4% of mating calls to the correct species. All of the calls by C. notatus, C. nigrovittatus and C. caniceps, and 88% of the calls by C. prevostii, were assigned to the correct species, while the percentage of correct classifications was lower in C. finlaysonii (71%) and C. erythraeus (63%). We compared these results with the genetic relationships to determine whether interspecific acoustic differences are caused by adaptive selection (habitat selection and body size) or by a stochastic process (drift). The genetic relationships among the six species were coincident with the differences in mating calls, which supports the stochastic divergence. Species-specific mating calls may be a useful cue for species recognition in Callosciurus, and thus these calls could be an effective trait for phylogenetic analysis in Callosciurus.

• Thomas, L. S., Teich, E., Dausmann, K., Reher, S. & Turner, J. M. (2018) Degree of urbanisation affects Eurasian red squirrel activity patterns. Hystrix, the Italian Journal of Mammalogy 29(2): 175-180.

With cities growing at a rapid pace, animal species must either retreat to patches of intact natural habitat or adapt to novel conditions in urban areas. While this disturbance causes most species to be in local decline, some show specific behavioural plasticity, facilitating success in a new habitat. The Eurasian red squirrel ( Sciurus vulgaris ) is a common small mammal species which occurs in high numbers in urban environments. To determine which characteristics enable its success, we investigated space use and activity budgets of seven free-ranging individuals living in semi-natural and urban habitats within a large city. We did not find significant differences in animals’ space use between habitat types but tendencies towards smaller home ranges and increased home range overlap existed among individuals in the urban site. Squirrels differed between sites in both overall activity levels and temporal activity patterns: urban animals spent less time active and activity onset was later compared to semi-natural conspecifics. This is likely explained by a combination of dense and reliable supplementary food sources in the urban habitat, reducing foraging effort, and restrictions to movement imposed by higher fragmentation. Flexibility in space use and activity budgets, as well as the ability to exploit anthropogenic food sources and tolerate reduced habitat connectivity, are likely the most important factors contributing to the squirrels’ success in cities. Accordingly, these traits could be used as indicators of low sensitivity towards urbanisation when assessing other species’ potential resilience. However, they do not immunise squirrels against extirpation. Further research on individuals’ foraging ecology and population health may reveal possible threats to urban red squirrels and help predict their future persistence in this challenging habitat.

• Turkia, T., Selonen, V., Danilov, P., Kurhinen, J., Ovaskainen, O., Rintala, J. & Brommer, J. E. (2018) Red squirrels decline in abundance in the boreal forests of Finland and NW Russia. Ecography 41(8): 1370-1379.

Abstract: Recent global warming and other anthropogenic changes have caused well-documented range shifts and population declines in many species over a large spatial extent. Most large-scale studies focus on birds, large mammals, and threatened species, whereas large-scale population trends of small to medium-sized mammals and species that are currently of least concern remain poorly studied. Large-scale studies are needed because on a smaller scale, important patterns may be masked by local variation and stochastic processes. Here, we utilized snow track census data from Finland and NW Russia to estimate population growth rates of the Eurasian red squirrel Sciurus vulgaris for a period of 17 yr in an area of over 1 000 000 km2. We also studied the effects of changes in summer and winter temperatures, winter precipitation, predator abundance, and canopy cover on estimated red squirrel population growth rates. Our results suggest that red squirrel populations have declined in most parts of the study area, the only remarkable exception being SW Russia. These results are in concordance with previous studies suggesting that species that are still common and of least concern may be declining. However, our findings are in contrast to the common pattern of northern populations of boreal species increasing due to global warming. The estimated population growth rates are in synchrony over vast areas, suggesting that the underlying reasons also operate on a large scale. We indeed find that the population growth rate was lower in regions where winters warmed faster during the study period, suggesting that changes in the environment (or biotic changes associated with it) are linked with the decline of red squirrels.

• Vander Haegen, W.M., et al., (2018) Endemic diseases affect population dynamics of tree squirrels in contrasting landscapes. The Journal of Wildlife Management 82(2):  328-343.

Abstract: Habitat loss and fragmentation can have detrimental effects on wildlife populations and where pervasive can create population isolates that may experience reduced genetic diversity and lower persistence. Diseases that cause epizootics also can reduce wildlife populations and may have disproportionate effects on small populations. We studied survival of radio-marked western gray squirrels (Sciurus griseus) using known-fate models in Program MARK and we quantified annual reproductive success by following females through the breeding season and counting young at natal nests. We used data on survival and productivity to model population growth rate and associated parameters using deterministic and stochastic approaches. Populations of western gray squirrels that we studied in an extensive, forested landscape and in a highly fragmented, urbanizing landscape in Washington, USA differed in their modeled growth rate. Adult survival was similar between populations although both were strongly affected by different endemic diseases with high epizootic potential. The demographic parameters that differed most between these 2 populations were related to productivity; litter size was marginally smaller and reproductive success was significantly lower in the urbanizing Puget Trough compared to rural Klickitat County. Results of our demographic modeling suggest that the larger Klickitat population is robust to immediate threats, whereas the smaller Puget Trough population is at risk because of its small size and low fecundity. Periodic outbreaks of notoedric mange in the Klickitat population reduce adult survival, although our models suggest that these epizootics would need to occur more frequently than observed to be of significant risk to the population. Continued degradation and fragmentation of western gray squirrel habitat in the Klickitat region along with mild winters resulting from climate change could increase the frequency and severity of mange epizootics and further threaten this population. Actions to retain and improve habitat resources may help ameliorate the effects of future mange epizootics and maintaining quality habitat should be a management priority in this region. The insular Puget Trough population experienced mortality due to tularemia each year of our study with infection rates ≥14%. Lack of characteristic histological indications in some affected squirrels may lead to underreporting of tularemia in animals submitted for routine necropsy and could complicate assessment of mortality risks in wildlife population studies. Given its small size and isolation, the Puget Trough population should be monitored closely for indications of decline in number or occupancy; this small population may need periodic augmentation to maintain genetic diversity. Increasing suitable habitat and maintaining connectivity between currently occupied range and potential habitat in the surrounding landscape will be crucial to the long-term viability of this population but will be challenging in the urbanizing landscape of the Puget Trough.

• Wells, C. P. & Van Vuren, D. H. (2018) Developmental and social constraints on early reproduction in an asocial ground squirrel. Journal of Zoology 306(1): 28-35.

Abstract: For short-lived species, selection for early reproduction should be strong, yet females often delay their first reproductive bout. Delay in age of first reproduction due to developmental constraints, such as food availability, or social constraints, such as the inhibitory presence of breeding adults, has been documented for social mammals, but effects on asocial species are less well known. We evaluated the influence of developmental and social factors on early reproduction in a short-lived, asocial species, the golden-mantled ground squirrel (Callospermophilus lateralis). We found that females who reproduced as yearlings had been weaned earlier in their natal summer and experienced early snow melt during their yearling spring, suggesting the importance of access to high-quality food at critical stages. Females were more likely to reproduce as yearlings when there were more adult males present during the breeding season, possibly because exposure to males accelerates reproductive maturity. Maternal presence had no effect on yearling reproduction, but yearlings with a littermate sister present were only 22% as likely to reproduce as females without a sister present, suggesting the effect of sibling competition well past weaning. Furthermore, the negative effect of a sister’s presence, but not the presence of other females, suggests that relatedness affects reproductive competition in this asocial species.

• Wernike, K., Wylezich, C., Höper, D., Schneider, J., Lurz, P. W. W., Meredith, A., Milne, E., Beer, M. & Ulrich, R. G. (2018) Widespread occurrence of squirrel adenovirus 1 in red and grey squirrels in Scotland detected by a novel real-time PCR assay. Virus Research 257: 113-118.

The Eurasian Red Squirrel (Sciurus vulgaris) is distributed throughout large parts of Europe and Asia. However, its distribution in certain regions of Europe is endangered by the invasive, non-native Grey Squirrel (Sciurus carolinensis). Adenoviruses were already described in squirrels in Great Britain almost two decades ago. In 2013, a squirrel adenovirus (SqAdV-1) was additionally found in a red squirrel from Germany, which suffered from acute diffuse catarrhal enteritis, and the complete genome sequence was determined. Here, samples from dead red (n = 25) and grey (n = 12) squirrels collected in Scotland, UK, were analysed for the presence of this squirrel-associated virus. By using a newly developed real-time PCR targeting the adenoviral polymerase gene, viral DNA was detected in at least one of four tissue samples tested per animal in 64.0% of the red squirrels and 41.7% of the grey squirrels. Exceptionally high viral genome loads were detected in the intestine and liver, but SqAdV-1 DNA was also present in lung and kidney samples of affected animals. Almost complete genome sequence determination of a red squirrel-derived SqAdV-1 strain from Scotland indicated a very high degree of identity to the first German strain. Sequence analysis of the hexon gene, which encodes one of the major antigens of the virion, revealed an identity of 100% between viruses found in red and grey squirrels from Scotland. In conclusion, SqAdV-1 appears to be widespread in the Scottish red and grey squirrel population, which highlights the necessity for continuous wildlife surveillance. The novel real-time PCR assay offers a highly sensitive and robust method for SqAdV-1 surveillance.

• Wishart, A. E., Williams, C. T., McAdam, A. G., Boutin, S., Dantzer, B., Humphries, M. M., Coltman, D. W. & Lane, J. E. (2018) Is biasing offspring sex ratio adaptive? A test of Fisher’s principle across multiple generations of a wild mammal in a fluctuating environment. Proceedings of the Royal Society B: Biological Sciences 285(1891).

Fisher’s principle explains that population sex ratio in sexually reproducing organisms is maintained at 1 : 1 owing to negative frequency-dependent selection, such that individuals of the rare sex realize greater reproductive opportunity than individuals of the more common sex until equilibrium is reached. If biasing offspring sex ratio towards the rare sex is adaptive, individuals that do so should have more grandoffspring. In a wild population of North American red squirrels (Tamiasciurus hudsonicus) that experiences fluctuations in resource abundance and population density, we show that overall across 26 years, the secondary sex ratio was 1 : 1; however, stretches of years during which adult sex ratio was biased did not yield offspring sex ratios biased towards the rare sex. Females that had litters biased towards the rare sex did not have more grandoffspring. Critically, the adult sex ratio was not temporally autocorrelated across years, thus the population sex ratio experienced by parents was independent of the population sex ratio experienced by their offspring at their primiparity. Expected fitness benefits of biasing offspring sex ratio may be masked or negated by fluctuating environments across years, which limit the predictive value of the current sex ratio.

• Zarco, A., Benitez, V., Fasola, L., Funes, G. & Guichón, L. (2018) Feeding habits of the Asiatic red-bellied squirrel Callosciurus erythraeus introduced in Argentina. Hystrix, the Italian Journal of Mammalogy 29(2): 223-228.

Knowledge of food habits of invasive species is necessary to predict invasion success and potential interactions in the invaded community. The Asiatic red-bellied squirrel Callosciurus erythraeus has been introduced in Asia, Europe and South America. We studied feeding habits of red-bellied squirrels in Argentina in two sites 600 km apart where wild populations have become established. We used both faecal microhistological analysis and behavioural records to describe diet composition and feeding habits. We also analysed diet selection and the potential role of the squirrels as seed disperser. Squirrels consumed items from 35 species of exotic trees and shrubs and one native tree species; fruits and seeds represented the bulk of the diet in all seasons (faeces analysis: >44%; behavioural observations: >38%). Squirrels also consumed epiphytic and climbing plants, ferns, invertebrates, fungi, lichens, mosses and bird eggs. Diet composition varied throughout the year according to food availability. We did not find evidence of endozoochoric dispersal but we observed squirrels carrying and dropping nuts and acorns during transport. The consumption of a wide range of food items and species, the ability to modify the diet according to food availability, and the capacity to hoard food indicate that feeding habits of red-bellied squirrels favour their invasive potential.

• Zozzoli, R., Menchetti, M. & Mori, E. (2018) Spatial behaviour of an overlooked alien squirrel: The case of Siberian chipmunks Eutamias sibiricus. Behavioural Processes 153: 107-111.

Abstract: Alien species of concern within the European Union have been recently listed and their populations need to be monitored, to plan addressed eradication or control programs. Therefore, the assessment of their presence should be rapidly carried out, particularly for elusive species or for those living at low densities. The Siberian chipmunk Eutamias sibiricus is a ground-dwelling squirrel, naturally distributed in northern and eastern Asia. Many introduced populations occur in Europe and Italy too. This species has been listed within the invasive species concern within the European Union and, thus, monitoring is mandatory to manage its potential range expansion. We carried out a hair-tube survey on 31 wood patches in northern and central Italy, where reproductive populations of Siberian chipmunk have been recorded. Hair tubes provided reliable data in assessing the presence of the Siberian chipmunk, with only 1% pseudo-absence and a high detection probability. The occurrence of Siberian chipmunk was positively influenced by study site and by the distance from release site, confirming low dispersal abilities by this species. Dense understorey also affected the presence of chipmunks, preventing them to search for food on the ground and to dig burrows.

• Zwolak, R. (2018) How intraspecific variation in seed-dispersing animals matters for plants. Biological Reviews 93(2): 897-913.

Abstract: Seed dispersal by animals is a complex phenomenon, characterized by multiple mechanisms and variable outcomes. Most researchers approach this complexity by analysing context-dependency in seed dispersal and investigating extrinsic factors that might influence interactions between plants and seed dispersers. Intrinsic traits of seed dispersers provide an alternative way of making sense of the enormous variation in seed fates. I review causes of intraspecific variability in frugivorous and granivorous animals, discuss their effects on seed dispersal, and outline likely consequences for plant populations and communities. Sources of individual variation in seed-dispersing animals include sexual dimorphism, changes associated with growth and ageing, individual specialization, and animal personalities. Sexual dimorphism of seed-dispersing animals influences seed fate through diverse mechanisms that range from effects caused by sex-specific differences in body size, to influences of male versus female cognitive functions. These differences affect the type of seed treatment (e.g. dispersal versus predation), the number of dispersed seeds, distance of seed dispersal, and likelihood that seeds are left in favourable sites for seeds or seedlings. The best-documented consequences of individual differences associated with growth and ageing involve quantity of dispersed seeds and the quality of seed treatment in the mouth and gut. Individual specialization on different resources affects the number of dispersed plant species, and therefore the connectivity and architecture of seed-dispersal networks. Animal personalities might play an important role in shaping interactions between plants and dispersers of their seeds, yet their potential in this regard remains overlooked. In general, intraspecific variation in seed-dispersing animals often influences plants through effects of these individual differences on the movement ecology of the dispersers. Two conditions are necessary for individual variation to exert a strong influence on seed dispersal. First, the individual differences in traits should translate into differences in crucial characteristics of seed dispersal. Second, individual variation is more likely to be important when the proportions of particular types of individuals fluctuate strongly in a population or vary across space; when proportions are static, it is less likely that intraspecific differences will be responsible for changes in the dynamics and outcomes of plant–animal interactions. In conclusion, focusing on variation among foraging animals rather than on species averages might bring new, mechanistic insights to the phenomenon of seed dispersal. While this shift in perspective is unlikely to replace the traditional approach (based on the assumption that all important variation occurs among species), it provides a complementary alternative to decipher the enormous variation observed in animal-mediated seed dispersal.

2017

• Amasifuén, C. & E. Heymann (2016) Toucan predation attempt on a Neotropical pygmy squirrel. Mammalia 81: 527.

Abstract: Predation on arboreal squirrels is rarely observed directly. Here we report the observation of a predatory attack on a Neotropical pygmy squirrel, Sciurillus pusillus (É. Geoffroy Saint-Hilaire, 1803), by a white-throated toucan, Ramphastos tucanos, from north-eastern Peruvian Amazonia. The toucan plucked the squirrel from a trunk at ca. 8 m height in flight and then slammed the individual against a liana in an apparent attempt to kill the prey item. This is the first field observation on a predation attempt on S. pusillus, and adds further evidence for predatory habits in toucans.

• Boonkhaw, P., Prayoon, U., Kanchanasaka, B., Hayashi, F., & Tamura, N. (2017) Colour polymorphism and genetic relationships among twelve subspecies of Callosciurus finlaysonii in Thailand. Mammalian Biology – Zeitschrift für Säugetierkunde. doi:dx.doi.org/10.1016/j.mambio.2017.02.001.

Abstract: Finlayson’s squirrel (Callosciurus finlaysonii) is distributed in lowland forests in Southeast Asia. The pelage colour is conspicuously polymorphic, and 16 subspecies have been described based on pelage colour patterns. Among them, 12 subspecies are distributed in Thailand, of which 7 are on the mainland and 5 are on islands. In addition, the distribution range of another closely related species, Pallas’s squirrel (Callosciurus erythraeus), overlaps the range of C. finlaysonii in western Thailand. In this study, phylogenetic analysis based on mitochondrial DNA suggested that C. finlaysonii and C. erythraeus did not form separate monophyletic groups. This C. finlaysonii/C. erythraeus complex in Thailand consisted of seven divergent groups, some of which may have arisen from isolation due to large rivers and seas: (1) C. f. nox/cinnamomeus, (2) C. f. finlaysonii/folletti/trotteri, (3) C. f. menamicus, (4) C. erythraeus, (5) C. f. annellatus, (6) C. f. bocourti/boonsongi/floweri, and (7) C. f. frandseni/albivexilli. Pelage colour did not consistently correspond to these genetic groups, suggesting that specific colours may have been acquired multiple times or the genes associated with colour may have variations and polymorphisms within subspecies. Several small populations in the lowlands and on small islands had lower genetic diversity. To conserve the local genetic diversity of C. finlaysonii, it may be necessary to enact legal restrictions on their trade and hunting.

• Bosch, S., P. W. W. Lurz, et al. (2017). “Nachweis eines neuen Squirrel-Adenovirus bei einem Krankheitsausbruch bei Eichhörnchen (Sciurus vulgaris) in Deutschland 2013 – 2016.  Detection of a new Squirrel-Adenovirus in a disease outbreak in German red squirrels 2013 – 2016 (in German).” Mitteilungen aus unserer Säugetierwelt 20:  27-35.

Summary: 2015 und 2016 kam  es bei wildlebenden Eichhörnchen zu einem großflächigen Krankheits- ausbruch mit
gehäuften Todesfällen in elf Bundesländern in Deutschland. Betroffen waren sowohl junge als auch erwachsene Eichhörnchen. Von den in Pflegestationen eingelieferten Tieren verstarben 161 und nur 24 überlebten. Kranke Tiere litten an zwei unterschiedlichen Symptomkomplexen: schwerer Durchfall oder schwere Atemnot. Bei den Tieren des aktuel- len  Ausbruches wurden  Adenoviren  nachgewiesen. Adenovirus-Infektionen  sind  bei  an schwerem Durchfall leidenden Eichhörnchen von Ausbrüchen in Großbritannien bekannt, wo in den Jahren 2005-2008 in Wales örtlich hohe Sterberaten beobachtet wurden.  Darüber hinaus gelang der histologische Nachweis von Darmparasiten und der kulturelle Nachweis mehrerer  bakterieller Erreger aus Lungen-, Leber- und Darmgewebe. Einige dieser Erreger sind in der  Lage, insbesondere bei  geschwächten,  immunkompromittierten  Tieren  eine schwere Infektion mit Todesfolge hervorzurufen. Zur Abklärung  der Frage, inwieweit die Erreger an der Erkrankung der Tiere beteiligt sind, sind weitere Untersuchungen erforderlich. In keinem der bisher
untersuchten Eichhörnchen aus Deutschland und Großbritannien wur- de ein Hinweis auf das zoonotische VSBV-1 gefunden.

• Brady Matthew, J., Koprowski John, L., Gwinn, R. N., Jo, Y.-S., & Young, K. (2017) Eastern fox squirrel (Sciurus niger, Linnaeus 1758) introduction to the Sonoran Desert. Mammalia 81: 221. doi:10.1515/mammalia-2015-0162.

Abstract: The eastern fox squirrel, native to the eastern and midwestern United States, was recently documented in the Sonoran Desert in the vicinity of Yuma, Arizona, constituting the first state record for this species. We surveyedthe people of Yuma to determine when and how the squirrels arrived. The squirrels were first observed in the 1960s, but may have been resident for a longer period. Since the 1960s, squirrels have spread throughout the city limits and extended south ~15 km into Somerton, Arizona. How the squirrels arrived is not clear, but must be the result of an introduction, as no nearby populations exist. The persistence of eastern fox squirrels in this unique habitat is due to synanthropic relationships.

• Brommer, J.E., Wistbacka, R. & Selonen, V.  (2017) Immigration ensures population survival in the Siberian flying squirrel. Ecology and Evolution 7: 1858-1868.

Abstract: Linking dispersal to population growth remains a challenging task and is a major knowledge gap, for example, for conservation management. We studied relative roles of different demographic rates behind population growth in Siberian flying squirrels in two nest-box breeding populations in western Finland. Adults and offspring were captured and individually identifiable. We constructed an integrated population model, which estimated all relevant annual demographic rates (birth, local [apparent] survival, and immigration) as well as population growth rates. One population (studied 2002–2014) fluctuated around a steady-state equilibrium, whereas the other (studied 1995–2014) showed a numerical decline. Immigration was the demographic rate which showed clear correlations to annual population growth rates in both populations. Population growth rate was density dependent in both populations. None of the demographic rates nor the population growth rate correlated across the two study populations, despite their proximity suggesting that factors regulating the dynamics are determined locally. We conclude that flying squirrels may persist in a network of uncoupled subpopulations, where movement between subpopulations is of critical importance. Our study supports the view that dispersal has the key role in population survival of a small forest rodent.

• Chavel, E. E., Mazerolle, M. J., Imbeau, L., & Drapeau, P. (2017) Comparative evaluation of three sampling methods to estimate detection probability of American red squirrels (Tamiasciurus hudsonicus). Mammalian Biology – Zeitschrift für Säugetierkunde 83: 1-9. doi:doi.org/10.1016/j.mambio.2016.11.003.

Abstract:  Measuring changes in species distribution and understanding factors influencing site occupancy are recurring goals in wildlife studies. Imperfect detection of species hinders such studies, resulting in the underestimation of the number of sites occupied by the species of interest. American red squirrels (Tamiasciurus hudsonicus) are sampled traditionally with live-traps that require substantial resources to deploy and monitor. Here, we assessed whether auditory methods yield similar detection probabilities. We compared the detection probability of American red squirrels in boreal forest using point counts, playback counts, and live-trapping. Over the summer of 2014, we conducted three trapping sessions in 60 sites within black spruce forests of northwestern Quebec, Canada. We also conducted 10 min point counts in the same sites, together with playback counts using recordings of American red squirrel alarm and territorial calls. Using dynamic occupancy models to analyse three primary periods, all composed of three secondary periods, we found that the detection probability of squirrels from point counts was as high as with live-trapping. Our results thus highlight the value of the point count method in measuring American red squirrel occupancy.

• Cooper, Eve B., Taylor, Ryan W., Kelley, Amanda D., Martinig, April R., Boutin, S., Humphries, Murray M., Dantzer, B., Lane, Jeffrey E. & McAdam, Andrew G. (2017) Personality is correlated with natal dispersal in North American red squirrels. Behaviour 154(9-10): 939-961.

Individual natal dispersal behaviour is often difficult to predict as it can be influenced by multiple extrinsic and intrinsic factors. Individual differences in personality have been shown to be an important correlate of dispersal behaviour. However, the relationships between personality traits and dispersal are often inconsistent within and across studies and the causes of these discrepancies are often unknown. Here we sought to determine how individual differences in activity and aggression, as measured in an open-field trial, were related to natal dispersal distance in a wild population of North American red squirrels (Tamiasciurus hudsonicus). For 14 cohorts, while individual aggression consistently had no association with dispersal distance, the association between activity and dispersal fluctuated through time, mediated by population density. The environmental-dependence of the relationship between personality and dispersal in this population is indicative of the importance of considering external conditions when predicting dispersal behaviour.

• Everest, D., Floyd, T., Donnachie, B., Irvine, R., Holmes, J., & Shuttleworth, C. (2017). Confirmation of squirrelpox in Welsh red squirrels. Veterinary Record 181: 514-515.

 

• J. Everest, D., M. Shuttleworth, C., Grierson, S., Dastjerdi, A., Stidworthy, M., Duff, J., J. Higgins, R., Mill, A. & Chantrey, J. (2017) The implications of significant adenovirus infection in UK captive red squirrel (Sciurus vulgaris) collections: How histological screening can aid applied conservation management. Mammalian Biology – Zeitschrift für Säugetierkunde 88: 123-129.Abstract: Conservation trans-locations using captive bred red squirrels (Sciurus vulgaris) is increasing in the United Kingdom (UK). However, project managers are often unaware of the risk of pathological adenovirus (ADV) infection. In this study we illuminate the viral threat using transmission electron microscopy (TEM) and polymerase chain reaction (PCR) assays. Both techniques were used to screen samples collected from 26 English and Welsh captive red squirrel collections. Of 181 carcasses received between 2002 and 2016, 129 (71%) were suitable for routine surveillance post mortem examination (PME). A range of tissues were examined with ADV identified from a variety of samples by PCR and TEM in 92 (72%) cases encompassing 23 of the 26 study collections (89%). ADV enteritis was histologically confirmed in two deaths (2%) with another 39 (30%) through both laboratory and clinical findings, considered as likely clinically-significant ADV cases, but advanced autolysis precluded accurate assessment and confirmatory histological diagnosis. Other positive cases were more indicative of sub-clinical infection. Clusters of ADV red squirrel deaths were recorded with circumstantial evidence suggesting inter-collection movement of presumed ADV infected donated animals had triggered mortality in recipient collections. During the study, several collections intermittently experienced ADV-associated deaths. Definitive cause of death was not determined in most cases, but a diverse range of diagnoses were recorded in 25 (19%) animals. Implications of these findings for captive United Kingdom (UK) red squirrel husbandry are discussed. It is recommended that protocols be drawn up to minimise potential intra-species ADV infection and highlight the danger of contact with ADV infected wood mice (Apodemus sylvaticus).

• Fisher, D. N., Boutin, S., Dantzer, B., Humphries, M. M., Lane, J. E., & McAdam, A. G. Multilevel and sex-specific selection on competitive traits in North American red squirrels. Evolution n/a-n/a. doi:10.1111/evo.13270.

Abstract: Individuals often interact more closely with some members of the population (e.g. offspring, siblings or group members) than they do with other individuals. This structuring of interactions can lead to multilevel natural selection, where traits expressed at the group-level influence fitness alongside individual-level traits. Such multilevel selection can alter evolutionary trajectories, yet is rarely quantified in the wild, especially for species that do not interact in clearly demarcated groups. We quantified multilevel natural selection on two traits, postnatal growth rate and birth date, in a population of North American red squirrels (Tamiasciurus hudsonicus). The strongest level of selection was typically within-acoustic social neighbourhoods (within 130m of the nest), where growing faster and being born earlier than nearby litters was key, while selection on growth rate was also apparent both within-litters and within-study areas. Higher population densities increased the strength of selection for earlier breeding, but did not influence selection on growth rates. Females experienced especially strong selection on growth rate at the within-litter level, possibly linked to the biased bequeathal of the maternal territory to daughters. Our results demonstrate the importance of considering multilevel and sex-specific selection in wild species, including those that are territorial and sexually monomorphic.

• Haigh A, Butler F, O’Riordan R, Palme R  (2017) Managed parks as a refuge for the threatened red squirrel (Sciurus vulgaris) in light of human disturbance. Biological Conservation 211 Part A: 29-36.

Abstract: As the invasive grey squirrel continues to spread, red squirrels are dying out. The result may be isolated populations in managed parks, where access can be controlled. However, recreation can often have a negative effect on wildlife, reducing the conservation potential of parks. Fota Wildlife Park receives over 300,000 visitors each year and is located on an island that is currently free of grey squirrels. We examined the effect of visitors on the existing red squirrel population. Sampling was conducted in the presence and absence of the public. Ten trapping sessions took place from March 2013 to 2014 and faeces were collected to examine stress levels. Squirrels were observed to concentrate their activity in non-public areas and move into public areas when the park was closed. Radio tracked squirrels, from the adjacent gardens (intermediate disturbance), also used habitats in the wildlife park (high disturbance) when it was closed but returned when the park had opened. When squirrels were observed in public areas, visitors were only visible on 15% of occasions. Levels of faecal cortisol metabolites (FCM) were highest in areas where human disturbance was greatest. However, there was no correlation between visitor numbers and the stress levels of squirrels. FCM levels were however, positively correlated with density of squirrels. The fact that high numbers of squirrels continued to utilise the wildlife park demonstrates that managed parks could provide an important reserve for the maintenance of the species, as long as non-public areas are accessible.

• Hoset, K. S., Villers, A., Wistbacka, R., & Selonen, V. (2017) Pulsed food resources, but not forest cover, determines lifetime reproductive success in a forest-dwelling rodent. Journal of Animal Ecology n/a-n/a. doi:10.1111/1365-2656.12715.

Abstract: 1.The relative contributions of habitat and food availability on fitness may provide evidence for key habitat features needed to safeguard population persistence. However, defining habitat quality for a species can be a complex task, especially if knowledge on the relationship between individual performance and habitat quality is lacking. 2.Here, we determined the relative importance of availability of suitable forest habitat, body mass, and food from masting tree species on female lifetime reproductive success (LRS) of Siberian flying squirrels (Pteromys volans). 3.We calculated LRS of 500 female flying squirrels based on a 22 year-long longitudinal data set of two populations from western Finland. We assessed with generalised additive models the potential effects of availability of suitable habitat and cumulative lifetime availability of food from masting tree species on female LRS, longevity and fecundity. On a reduced dataset, we evaluated the importance of female winter body mass and conducted a piecewise path analysis to determine how variables were connected. 4.According to generalised additive models female longevity, fecundity and LRS were mainly determined by variation in cumulative lifetime availability of food from masting alder and birch. Instead, habitat and body mass had smaller role. The path analysis indicated that lifetime food availability had direct effect on longevity and fecundity, and these had equal effect on LRS at both study sites. 5.Our results on LRS shows that the occurrence of tree masting events during a flying squirrel female’s lifetime have profoundly larger effect on lifetime reproductive success than the cover of suitable forest habitat. Furthermore, this study emphasises the importance of both fecundity and longevity, and the indirect effects of food availability via those components, as determinants of lifetime fitness of female flying squirrels.

• Inskip, S., Taylor, G. M., Anderson, S., & Stewart, G. (2017). Leprosy in pre-Norman Suffolk, UK: biomolecular and geochemical analysis of the woman from Hoxne. Journal of Medical Microbiology 66(11): 1640-1649. doi:doi:10.1099/jmm.0.000606

Abstract: PURPOSE: A woman’s skull, exhibiting features of lepromatous leprosy (LL), was recovered from a garden in Hoxne, Suffolk. The absence of post crania and lack of formal excavation meant that diagnosis and dating was uncertain. The aim of this research was to confirm the diagnosis using biomolecular means and second, to place it in context with other British leprosy cases using SNP genotyping and radiocarbon dating. METHODOLOGY: Bone from the skull was analysed by ancient DNA (aDNA) methods and subjected to radiocarbon dating. As a result, stable carbon and nitrogen isotope values were produced, both useful for assessing aspects of the woman’s diet.Results/Key findings. aDNA confirmed the presence of mycobacterium leprae and genotyping demonstrated an ancestral variant of subtype 3I, the same lineage recently identified in living squirrels in the south of England. Radiocarbon dating revealed the woman lived approximately between 885-1015 AD, providing evidence for endurance of this subtype in East Anglia, having been previously identified as early as the fifth-sixth century (Great Chesterford) and as late as the thirteenth century (Ipswich). CONCLUSIONS: The confirmation of a new pre-Norman leprosy case in East Anglia is of interest as this is where a high proportion of cases are located. Possible factors for this may include preservation and excavation biases, population density, but also connection and trade, possibly of fur, with the continent. Future research on other British LL cases should focus on exploring these aspects to advance understanding of the disease’s history, here and on the continent.

• Jokimäki, J., Selonen, V., Lehikoinen, A. & Kaisanlahti-Jokimäki, M.-L. (2017) The role of urban habitats in the abundance of red squirrels (Sciurus vulgaris, L.) in Finland. Urban Forestry & Urban Greening 27: 100-108.

Abstract: Because the amount of urban areas has increased, it is important to investigate the abundance of wildlife species in relation to urban environments. Analyzing the impact of urbanization on the presence of forest-dwelling mammals is of interest due to the possible effects of urbanization on human-wildlife relationships and urban biodiversity. The Eurasian red squirrel (Sciurus vulgaris) is a declining forest species, and its occurrence in urban environments has been inadequately studied. The loss and fragmentation of forests due to urbanization may be detrimental for squirrels, whereas the abundant and predictable food resources and the low number of natural predators in urban areas may encourage squirrels to invade towns. We used large-scale data collected by volunteer bird watchers along a 950km south-north gradient to study whether the winter abundance of squirrels in Finland is dependent on urbanization, while controlling for effects of habitat type, food abundance (spruce cone crop; number of winter feeding sites), predator abundance (northern goshawk, Accipiter gentilis; feral cat Felis catus), season and latitude. We found that squirrel abundance increased with human population density, number of feeding sites and spruce cone crop and decreased with latitude and season. Feral cats showed weak negative connection with squirrel numbers, but there were no effect of goshawks. Relative squirrel abundance was approximately twice as high in urban habitats than in forests. Artificial feeding rather than a low number of predators may attract squirrels in urban environments. Planting spruce trees in urban environments will also benefit squirrels. Our results indicate that urban areas are an important habitat for the red squirrel even along the northern edge of their distribution range, where natural forest areas are still widespread. We conclude also that a citizen science −based bird survey protocol associated with mammal surveys seems to be a good large-scale monitoring method to study the urbanization of squirrels.

• Jones, H., White, A., Lurz, P., & Shuttleworth, C. (2017) Mathematical models for invasive species management: Grey squirrel control on Anglesey. Ecological Modelling 359: 276-284. doi:doi.org/10.1016/j.ecolmodel.2017.05.020.

Abstract: The control of invasive species and protection of threatened native species require well-developed policy and species management strategies. Mathematical models provide a key tool that can be used to test, develop and optimise strategies to manage invasive species. We use the native red squirrel and invasive grey squirrel system on the Island of Anglesey, UK, as a case study system in which to parameterise a mathematical model that includes the control of grey squirrels. We develop a stochastic, spatial model that represents the real habitat structure, distribution and linkage on Anglesey and the neighbouring mainland and includes the key population and epidemiological dynamics of the red-grey-squirrelpox system. The model also includes a representation of the trapping and removal of grey squirrels which is parameterised from field data on Anglesey in which grey squirrel were removed and red squirrels reintroduced between 1998–2013. The model is used to assess different management procedures to protect red squirrels from island re-invasion by grey squirrels, including the threat of squirrelpox spread posed by endemic mainland grey populations. The findings have important implications for the conservation of threatened red squirrels throughout the UK and in Europe. Moreover, the modelling framework is based on well-understood, classical models of competitive and epidemiological interactions and therefore the techniques can be adapted and applied more generally to manage the threat of invasive species in a wide range of natural systems.

• Koprowski, J., Nieto-Montes de Oca, A., Palmer, G., Ramos-Lara, N. & Timm, R. (2017) Sciurus aureogaster (Rodentia: Sciuridae). Mammalian Species, 49, 81.

• Lichti, N. I., Steele, M. A., & Swihart, R. K. (2017) Seed fate and decision-making processes in scatter-hoarding rodents. Biological Reviews 92: 474-504. doi:10.1111/brv.12240.

Abstract: A mechanistic understanding of seed movement and survival is important both for the development of theoretical models of plant population dynamics, spatial spread, and community assembly, and for the conservation and management of plant communities under global change. While models of wind-borne seed dispersal have advanced rapidly over the past two decades, models for animal-mediated dispersal have failed to make similar progress due to their dependence on interspecific interactions and complex, context-dependent behaviours. In this review, we synthesize the literature on seed dispersal and consumption by scatter-hoarding, granivorous rodents and outline a strategy for development of a general mechanistic seed-fate model in these systems. Our review decomposes seed dispersal and survival into six distinct sub-processes (exposure, harvest, allocation, preparation, placement, and recovery), and identifies nine intermediate (latent) variables that link physical state variables (e.g. seed and animal traits, habitat structure) to decisions regarding seed allocation to hoarding or consumption, cache placement and management, and deployment of radicle-pruning or embryo excision behaviours. We also highlight specific areas where research on these intermediate relationships is needed to improve our mechanistic understanding of scatter-hoarder behaviour. Finally, we outline a strategy to combine detailed studies on individual functional relationships with seed-tracking experiments in an iterative, hierarchical Bayesian framework to construct, refine, and test mechanistic models for context-dependent, scatter-hoarder-mediated seed fate.

• Marvel-Coen, J. (2017). Finding new homes: Multilevel selection on birth timing and growth in North American red squirrels. Evolution 71(7): 1917-1918. doi:10.1111/evo.13293

Abstract: Individuals often interact more closely with some members of the population (e.g., offspring, siblings, or group members) than they do with other individuals. This structuring of interactions can lead to multilevel natural selection, where traits expressed at the group-level influence fitness alongside individual-level traits. Such multilevel selection can alter evolutionary trajectories, yet is rarely quantified in the wild, especially for species that do not interact in clearly demarcated groups. We quantified multilevel natural selection on two traits, postnatal growth rate and birth date, in a population of North American red squirrels (Tamiasciurus hudsonicus). The strongest level of selection was typically within-acoustic social neighborhoods (within 130 m of the nest), where growing faster and being born earlier than nearby litters was key, while selection on growth rate was also apparent both within-litters and within-study areas. Higher population densities increased the strength of selection for earlier breeding, but did not influence selection on growth rates. Females experienced especially strong selection on growth rate at the within-litter level, possibly linked to the biased bequeathal of the maternal territory to daughters. Our results demonstrate the importance of considering multilevel and sex-specific selection in wild species, including those that are territorial and sexually monomorphic.

• Mazzamuto, M. V., Morandini, M., Panzeri, M., Wauters, L. A., Preatoni, D. G., & Martinoli, A. (2017) Space invaders: effects of invasive alien Pallas’s squirrel on home range and body mass of native red squirrel. Biological Invasions  n/a-n/a doi:10.1007/s10530-017-1396-2.

Abstract: 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.

• Merrick, M. J., & Koprowski, J. L. (2017) Circuit theory to estimate natal dispersal routes and functional landscape connectivity for an endangered small mammal. Landscape Ecology 32: 1163-1179. doi:10.1007/s10980-017-0521-z.

Abstract: Natal dispersal links population dynamics to landscape connectivity. Understanding how organisms perceive barriers to movement, or landscape resistance, during natal dispersal is important to conserve and manage populations threatened by fragmentation and habitat loss.

• Millins, C., Gilbert, L., Medlock, J., Hansford, K., Thompson, D. B., & Biek, R. (2017) Effects of conservation management of landscapes and vertebrate communities on Lyme borreliosis risk in the United Kingdom. Philosophical Transactions of the Royal Society B: Biological Sciences 372. doi:10.1098/rstb.2016.0123.

Abstract: Landscape change and altered host abundance are major drivers of zoonotic pathogen emergence. Conservation and biodiversity management of landscapes and vertebrate communities can have secondary effects on vector-borne pathogen transmission that are important to assess. Here we review the potential implications of these activities on the risk of Lyme borreliosis in the United Kingdom. Conservation management activities include woodland expansion, management and restoration, deer management, urban greening and the release and culling of non-native species. Available evidence suggests that increasing woodland extent, implementing biodiversity policies that encourage ecotonal habitat and urban greening can increase the risk of Lyme borreliosis by increasing suitable habitat for hosts and the tick vectors. However, this can depend on whether deer population management is carried out as part of these conservation activities. Exclusion fencing or culling deer to low densities can decrease tick abundance and Lyme borreliosis risk. As management actions often constitute large-scale perturbation experiments, these hold great potential to understand underlying drivers of tick and pathogen dynamics. We recommend integrating monitoring of ticks and the risk of tick-borne pathogens with conservation management activities. This would help fill knowledge gaps and the production of best practice guidelines to reduce risks.

• Oberosler, V., Groff, C., Iemma, A., Pedrini, P., & Rovero, F. (2017) The influence of human disturbance on occupancy and activity patterns of mammals in the Italian Alps from systematic camera trapping. Mammalian Biology – Zeitschrift für Säugetierkunde. doi:doi.org/10.1016/j.mambio.2017.05.005.

Abstract: As human activities increase in natural areas, so do threats to wildlife, potentially leading to immediate and long-term impacts on species’ distribution, activity, reproduction and survival. This is particularly relevant for large-bodied vertebrates that are especially sensitive to human presence and human-driven habitat changes. Assessing the impact of anthropogenic disturbance requires data on species distribution and activity patterns of target species in relation to human presence and infrastructures. Here, we used camera trap data to study the influence of anthropogenic disturbance on the community of medium-to-large mammals in a mountainous area in the eastern Italian Alps, with emphasis on the local population of brown bear (Ursus arctos). In 2015, we sampled a study area of 220 km2 with 60 camera trap locations adopting a systematic grid. Such design was inspired by the terrestrial vertebrate monitoring protocol developed by the TEAM Network, a pan-tropical biodiversity programme. Camera traps run for 30 days in each site and cumulated 1,978 camera trapping days, yielding 1,514 detection events of 12 species of mammals. For the 8 most recorded species, we used detection/non-detection data to model estimated occupancy and detection probability in relation to a suite of environmental and disturbance covariates. Our analysis revealed that human disturbance plays a significant role in influencing species-specific detection probability, while we found little evidence of significant relationship between occupancy and anthropogenic disturbance. For example, we found that brown bear’s detectability was negatively correlated with capture rate of humans at sampling sites, and positively correlated with distance from settlements. We also assessed species-specific daily activity patterns and found that, for all species, the overlap with human diel pattern decreases significantly at sites with higher human presence. We also discuss the potential of our approach for building cost-efficient and long-term monitoring of mammals.

• Sadeghi, M., & Malekian, M. (2017) The Persian squirrel of Kurdistan Province, western Iran: what determines its geographic distribution? Mammalia 81: 309-314.

Abstract: Here, we used the maximum entropy (MAXENT) method to predict habitat distribution of the Persian squirrel in oak forests of Kurdistan Province, western Iran. We used 70 points with known occurrence of the species and 17 environmental variables (climatic variables represented annual trends in temperature and precipitation, seasonality and extreme or limiting environmental factors) to map the species distribution. The MAXENT model showed high performance. Using a 0.5 logistic probability threshold, the models suggested about 16,783.5 ha of the study area to have high suitability for the Persian squirrel. These areas were thus estimated as “good” habitats. Amongst the environmental variables, land cover had the greatest role in the Persian squirrel’s distribution. Precipitation and temperature were the two major climatic factors that affected the Persian squirrel’s distribution. Gap analysis showed that many parts of the species habitat have remained unprotected what can threaten the survival of the studied species in the region. These findings can be used to develop conservation management plans and boost the network of protected areas in the region.

• Selonen, V. & Wistbacka, R.  (2017) Role of breeding and natal movements in lifetime dispersal of a forest-dwelling rodent. Ecology and Evolution 7: 2204-2213.

Abstract: The lifetime movements of an individual determine the gene flow and invasion potential of the species. However, sex dependence of dispersal and selective pressures driving dispersal have gained much more attention than dispersal at different life and age stages. Natal dispersal is more common than dispersal between breeding attempts, but breeding dispersal may be promoted by resource availability and competition. Here, we utilize mark–recapture data on the nest-box population of Siberian flying squirrels to analyze lifetime dispersal patterns. Natal dispersal means the distance between the natal nest and the nest used the following year, whereas breeding movements refer to the nest site changes between breeding attempts. The movement distances observed here were comparable to distances reported earlier from radio-telemetry studies. Breeding movements did not contribute to lifetime dispersal distance and were not related to variation in food abundance or habitat patch size. Breeding movements of males were negatively, albeit not strongly, related to male population size. In females, breeding movement activity was low and was not related to previous breeding success or to competition between females for territories. Natal philopatry was linked to apparent death of a mother; that is, we did not find evidence for mothers bequeathing territories for offspring, like observed in some other rodent species. Our results give an example of a species in which breeding movements are not driven by environmental variability or nest site quality. Different evolutionary forces often operate in natal and breeding movements, and our study supports the view that juveniles are responsible for redistributing individuals within and between populations. This emphasizes the importance of knowledge on natal dispersal, if we want to understand consequences of movement ecology of the species at the population level.

• Shonfield, J., Gorrell, J. C., Coltman, D. W., Boutin, S., Humphries, M. M., Wilson, D. R., et al. (2017) Using playback of territorial calls to investigate mechanisms of kin discrimination in red squirrels. Behavioral Ecology 28: 382-390.

Abstract: Kin recognition can facilitate kin selection and may have played a role in the evolution of sociality. Red squirrels (Tamiasciurus hudsonicus) defend territories using vocalizations known as rattles. They use rattles to discriminate kin, though the mechanism underlying this ability is unknown. Our objective was to distinguish between the mechanisms of prior association, where animals learn the phenotypes of kin they associate with early in life, and phenotype matching/recognition alleles, where animals use a template to match phenotypes, thereby allowing them to recognize kin without an association early in life. We used audio playbacks to measure the responses of squirrels to rattles from familiar kin, unfamiliar kin, and non-kin. Initial analyses revealed that red squirrels did not discriminate between familiar and unfamiliar kin, but also did not discriminate between kin and non-kin, despite previous evidence indicating this capability. Post hoc analyses showed that a squirrel’s propensity to rattle in response to playback depended on an interaction between relatedness and how the playback stimuli had been recorded. Red squirrels discriminated between rattles from close kin (r = 0.5) and rattles from non-kin (r &lt; 0.125) when the rattles were recorded from provoked squirrels. Squirrels did not exhibit kin discrimination in response to unsolicited rattles. Once we accounted for how the stimuli had been recorded, we found no difference in the responses to familiar and unfamiliar kin. Our study suggests that kin discrimination by red squirrels may be context dependent.

• Shuttleworth, C. (2017). Grey squirrels are bad for the British countryside – full stop. The Conservation (April 6 2017).

• Siracusa, E., Boutin, S., Humphries, M. M., Gorrell, J. C., Coltman, D. W., Dantzer, B., . . . McAdam, A. G. (2017). Familiarity with neighbours affects intrusion risk in territorial red squirrels. Animal Behaviour 133 (Supplement C), 11-20. doi:doi.org/10.1016/j.anbehav.2017.08.024

Abstract: Interactions with conspecifics are an important aspect of an individual’s environment. Although it is well known that the presence of conspecifics can have important effects on behaviour, in general it is also now acknowledged that the composition of the social environment can vary, and that this variation may have profound effects on individual behaviour and fitness. Using a wild population of North American red squirrels, Tamiasciurus hudsonicus, we investigated the importance of the composition of the social environment in a territorial species by assessing whether the risk of intrusion faced by territory owners varied with the degree of relatedness and familiarity in their social neighbourhoods. To test this, we conducted temporary removals of territory owners and observed the time until intrusion and the identity of intruding individuals. We found that individuals in neighbourhoods with low average familiarity faced a higher risk of intrusion and that unfamiliar neighbours were more likely to intrude. Surprisingly, we found that related neighbours also posed a higher risk of intrusion. The results from our study suggest that familiarity with neighbours may be an ecologically and evolutionarily relevant measure of the social environment, even in a species considered to be ‘asocial’. Future studies should consider the potential importance of the social environment, which has heretofore been mostly overlooked, as a relevant selective pressure in asocial, territorial species.

• Steen, R., & Barmoen, M. (2017) Diel activity of foraging Eurasian red squirrels ( Sciurus vulgaris ) in the winter revealed by camera traps. Hystrix, the Italian Journal of Mammalogy 28  . doi:10.4404/hystrix-28.1-11997.

Abstract: Different animals are adapted to a range of activity patterns, from diurnal to nocturnal. Under normal conditions, the 24-hour rhythm is entrained by changes in light intensity during twilight, i.e. the light and dark cycle. Many rodent species exhibit predator avoidance by adjusting their activity in relation to space or time, which is weighed against the need for food. Hence, an individual’s foraging activity consists of a compromise between foraging and predation risk, in addition to competition. Our study species, the Eurasian red squirrel, is diurnal and utilises dense forest habitats. In the present study, we used non-invasive camera traps to model diel activity during the winter months in 2014-2015 (December to February 24-hour day). We found that the Eurasian red squirrels exhibited a strictly diurnal feeding activity pattern and the shape of the modelled activity curve was unimodal and concentrated in the daylight hours, starting at sunrise and peaking in the morning.

• Sullivan, T. P., Ransome, D. B., Sullivan, D. S., Lindgren, P. M. F., & Klenner, W. (2017) Tree squirrel abundance and demography in managed coniferous forests of British Columbia are within the range of natural fluctuations of old-growth stands. Canadian Journal of Forest Research 47: 565-582. doi:10.1139/cjfr-2016-0458.

Abstract: The American red squirrel (Tamiasciurus hudsonicus Exrleben) and northern flying squirrel (Glaucomys sabrinus Shaw) are important mammal species in coniferous forests that are widely distributed across temperate and boreal ecological zones. Although T. hudsonicus and G. sabrinus apparently prefer late-successional forests, their population dynamics show no clear pattern in young second-growth and old-growth conifer forests. We used a compilation of study results that had standardized methodology and sampling effort to compare tree squirrel responses across a range of forest conditions. We tested the hypotheses (H) that abundance, reproduction, recruitment, and survival would be higher in (H1) old-growth than second-growth stands, (H2) unthinned than thinned second-growth stands, and (H3) lightly than heavily thinned stands. Tree squirrel populations in old-growth stands were considered the “standard” treatment to which other treatments in second-growth stands (unthinned and variously thinned) were compared. Thinned stands were grouped into low, medium, and high densities of trees. Datasets from seven published studies included 2804 Tamiasciurus and 837 G. sabrinus individuals, 25 study years, and 158 trapping periods. Mean abundance of Tamiasciurus was similar among treatment stands, ranging from 0.62 to 1.29·ha−1. Mean numbers of G. sabrinus were highest in old-growth stands at 0.65·ha−1, followed by the low- and medium-density stands at 0.50·ha−1, the high-density stands at 0.25·ha−1, and finally the unthinned second-growth stands at a low of 0.12·ha−1. In terms of effect size, mean proportional change in abundance, relative to that within old-growth stands, of both squirrel species in the three thinned stands were within the range of natural fluctuations in old-growth stands. Populations of Tamiasciurus in the unthinned stands were not within the range of fluctuations in the old-growth stands. Mean number of successful pregnancies and total recruits were similar among stands for Tamiasciurus, with variability in female breeding and winter survival. Mean number of recruits of G. sabrinus was highest in old-growth stands with no other differences in demographic attributes among stands. The biological significance of these differences in demographic variables was small in terms of effect sizes. Tamiasciurus and G. sabrinus seem to persist in a relatively broad range of young managed forest habitats, as well as in old-growth stands.

• Turkia, T., Selonen, V., Danilov, P., Kurhinen, J., Ovaskainen, O., Rintala, J., & Brommer, J. (2017). Red squirrels decline in abundance in the boreal forests of Finland and NW Russia. Ecography  n/a-n/a. doi:10.1111/ecog.03093

Abstract: Recent global warming and other anthropogenic changes have caused well-documented range shifts and population declines in many species over a large spatial extent. Most large-scale studies focus on birds, large mammals, and threatened species, whereas large-scale population trends of small to medium-sized mammals and species that are currently of least concern remain poorly studied. Large-scale studies are needed because on a smaller scale, important patterns may be masked by local variation and stochastic processes. Here, we utilized snow track census data from Finland and NW Russia to estimate population growth rates of the Eurasian red squirrel (Sciurus vulgaris L) for a period of 17 years in an area of over 1 000 000 km2. We also studied the effects of changes in summer and winter temperatures, winter precipitation, predator abundance, and canopy cover on estimated red squirrel population growth rates. Our results suggest that red squirrel populations have declined in most parts of the study area, the only remarkable exception being SW Russia. These results are in concordance with previous studies suggesting that species that are still common and of least concern may be declining. However,our findings are in contrast to the common pattern of northern populations of boreal species increasing due to global warming. The estimated population growth rates are in synchrony over vast areas, suggesting that the underlying reasons also operate on a large scale. We indeed find that the population growth rate was lower in regions where winters warmed faster during the study period, suggesting that changes in the environment (or biotic changes associated with it) are linked with the decline of red squirrels.

• Turner, J. M., Reher, S., Warnecke, L. & Dausmann, K. H. (2017) Eurasian Red Squirrels Show Little Seasonal Variation in Metabolism in Food-Enriched Habitat. Physiological and Biochemical Zoology 90(6): 655-662.

Abstract: Energy expenditure and ambient temperature (Ta) are intrinsically linked through changes in an animal’s metabolic rate. While the nature of this relationship is stable, the breadth of change in thermoregulatory cost varies with body size and physiological acclimatization to season. To explore seasonal metabolic changes of small mammals, we studied a population of Eurasian red squirrels (Sciurus vulgaris) in a seminatural environment with a year-round supply of natural and supplemented food. In each season we measured the metabolic rate of wild-caught red squirrels, using open-flow respirometry, and hypothesized that individuals would make adjustments to contend with seasonal weather conditions. In comparison to summer animals, we predicted that winter squirrels would show (1) an increase in metabolic rate within the thermoneutral zone, (2) a decrease in the lower critical temperature of the thermoneutral zone, (3) a shallower slope of resting metabolic rate with decreasing Ta, and (4) lower thermal conductance. Surprisingly, we observed only minor changes in resting metabolic rate, and energetic modeling suggested that the scope of change was unlikely to be of ecological consequence. Hair area density was higher in winter than in summer, corresponding to a slightly elevated thermal conductance in summer, while body mass was reasonably constant year-round. We conclude that the scope of physiological seasonal adaptation is minimal when food is abundant and that squirrels instead rely on adjustments in activity to reduce exposure to low Ta. We suggest that this may explain the squirrel’s success in a wide range of habitats, including urban areas, which require a rapid and flexible response to environmental changes and may indicate the capacity of other small mammal species to cope with environmental disturbance.

• Tye, C. A., McCleery, R. A., Fletcher, R. J., Greene, D. U., & Butryn, R. S. (2017) Evaluating citizen vs. professional data for modelling distributions of a rare squirrel. Journal of Applied Ecology 54: 628-637. doi:10.1111/1365-2664.12682.

Abstract: * To realize the potential of citizens to contribute to conservation efforts through the acquisition of data for broad-scale species distribution models, scientists need to understand and minimize the influences of commonly observed sample selection bias on model performance. Yet evaluating these data with independent, planned surveys is rare, even though such evaluation is necessary for understanding and applying data to conservation decisions. * We used the state-listed fox squirrel Sciurus niger in Florida, USA, to interpret the performance of models created with opportunistic observations from citizens and professionals by validating models with independent, planned surveys. * Data from both citizens and professionals showed sample selection bias with more observations within 50 m of a road. While these groups showed similar sample selection bias in reference to roads, there were clear differences in the spatial coverage of the groups, with citizens observing fox squirrels more frequently in developed areas. * Based on predictions at planned field surveys sites, models developed from citizens generally performed similarly to those developed with data collected by professionals. Accounting for potential sample selection bias in models, either through the use of covariates or via aggregating data into home range size grids, provided only slight increases in model performance. * Synthesis and applications. Despite sample selection biases, over a broad spatial scale opportunistic citizen data provided reliable predictions and estimates of habitat relationships needed to advance conservation efforts. Our results suggest that the use of professionals may not be needed in volunteer programmes used to determine the distribution of species of conservation interest across broad spatial scales.

• Wauters, L. A., Amori, G., Aloise, G., Gippoliti, S., Agnelli, P., Galimberti, A., et al. (2017) New endemic mammal species for Europe: Sciurus meridionalis (Rodentia, Sciuridae). 2017, 28.  doi:10.4404/hystrix-28.1-12015.

Abstract: Combining genetic, morphological and geographical data, we re-evaluate Sciurus meridionalis, Lucifero 1907 as a tree squirrel species. The species, previously considered a subspecies of the Eurasian red squirrel, Sciurus vulgaris, is endemic to South Italy with a disjunct distribution with respect to S. vulgaris. The new species has a typical, monomorphic coat colour characterized by a white ventral fur and a very dark-brown to blackish fur on the back, sides and tail. Specimens of S. meridionalis have a larger hind foot length and weigh about 35% more than live-caught S. vulgaris from northern Italy. S. meridionalis is larger than S. vulgaris specimens from three other regions in Italy for mandible length, skull width and skull (condylobasal) length, and principal component scores indicate significant shape differences of specimens from the Calabria population ( S. meridionalis ) compared to all other specimens ( S. vulgaris ). These morphological differences are further supported by genetic evidence at three mitochondrial markers (D-loop, cytochrome b and the DNA barcoding region COI) using the widest molecular dataset ever assembled for Sciurus vulgaris and S. meridionalis . All the investigated markers revealed exclusive haplotypes for S. meridionalis well separated from those of S. vulgaris and previously published results based on nuclear markers further support our taxonomic hypothesis. We suggest Calabrian black squirrel as common name for this new taxon.

• Wibbelt G, T. S., Dabrowski PW, Kershaw O, Nitsche A, Schrick L. (2017) Berlin squirrelpox virus, a new poxvirus in red squirrels, Berlin, Germany. Emerging Infectious Diseases 23. doi.org/10.3201/eid2310.171008. (2017)

Abstract: Near Berlin, Germany, several juvenile red squirrels (Sciurus vulgaris) were found with moist, crusty skin lesions. Histology, electron microscopy, and cell culture isolation revealed an orthopoxvirus-like infection. Subsequent PCR and genome analysis identified a new poxvirus (Berlin squirrelpox virus) that could not be assigned to any known poxvirus genera.

• Zelditch, M. L., Ye, J., Mitchell, J. S., & Swiderski, D. L. (2017) Rare ecomorphological convergence on a complex adaptive landscape: Body size and diet mediate evolution of jaw shape in squirrels (Sciuridae). Evolution, n/a-n/a. doi:10.1111/evo.13168.

Abstract: Convergence is widely regarded as compelling evidence for adaptation, often being portrayed as evidence that phenotypic outcomes are predictable from ecology, overriding contingencies of history. However, repeated outcomes may be very rare unless adaptive landscapes are simple, structured by strong ecological and functional constraints. One such constraint may be a limitation on body size because performance often scales with size, allowing species to adapt to challenging functions by modifying only size. When size is constrained, species might adapt by changing shape; convergent shapes may therefore be common when size is limiting and functions are challenging. We examine the roles of size and diet as determinants of jaw shape in Sciuridae. As expected, size and diet have significant interdependent effects on jaw shape and ecomorphological convergence is rare, typically involving demanding diets and limiting sizes. More surprising is morphological without ecological convergence, which is equally common between and within dietary classes. Those cases, like rare ecomorphological convergence, may be consequences of evolving on an adaptive landscape shaped by many-to-many relationships between ecology and function, many-to-one relationships between form and performance, and one-to-many relationships between functionally versatile morphologies and ecology. On complex adaptive landscapes, ecological selection can yield different outcomes.

2016

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Book

Shuttleworth, C., Lurz, P., & Gurnell, J. (2016) The Grey Squirrel: Ecology & Management of an Invasive Species in Europe, 532 pp. European Squirrel Initiative, Stoneleigh Park, Warwickshire CV8 2LG UK.

Chapters

• Gurnell, J. (2016). Foreword. The Grey Squirrel: Ecology & Management of an Invasive Species in Europe. C. Shuttleworth, P. Lurz and J. Gurnell, European Squirrel Initiative, Stoneleigh Park, Warwickshire CV8 2LG UK: viii-ix.
•  Bertolino, S., Lurz, P., Shuttleworth, C., Martinoli, A., & Wauters, L ((2016). The management of grey squirrel populations in Europe: evolving best practice. The Grey Squirrel: Ecology & Management of an Invasive Species in Europe. C. Shuttleworth, P. Lurz and J. Gurnell, European Squirrel Initiative, Stoneleigh Park, Warwickshire CV8 2LG UK: 495-514.

Abstract: Grey squirrel management is conducted in Great Britain, Ireland and Italy for red squirrel conservation and timber protection. However, management plans differ hugely with respect to aims, geographical scales, methodologies and costs. A key universal element linked to squirrel control is population monitoring. However, in the United Kingdom (UK) it took a lot of time to convince volunteer squirrel groups to accurately record control effort and success in order to allow subsequent analysis of population trends. In Italy, after decades of inaction, during which time grey squirrels colonized five regions due to translocations within the country, new management actions finally started. With the aid of two European co-funded projects, eradication and control activities have been initiated in the Lombardy, Piedmont, Liguria and Umbria regions. The removal of the grey squirrel from the island of Anglesey was the first eradication project which led to the complete removal of the species from a defined European area, though the bridge connections with the mainland necessitate continued interventions. More  recently, an urban  population was eradicated in Italy through live-trapping and surgical sterilization with subsequent release of the animals into other isolated parks. These two projects are key steps toward a widespread management of this American squirrel species, with the aims to spatially contain or even eradicate grey squirrel populations over large areas. While  the present legal framework ensures the possibility to manage alien squirrels and the available techniques are effective, there is still a need to work on communication skills to increase the consensus for alien species control among citizens, especially at a local scale

• Cowan, D., Mill, A., Everest, D., Gomm, M., Mcinnes, C., Rushton, S., Shirley, M.D.F., Start, C. & Shuttleworth, C.M.(2016). The potential role of ectoparasites in the epidemiology of squirrelpox virus: a possible novel means of intervention to reduce the impact of the disease? . The Grey Squirrel: Ecology & Management of an Invasive Species in Europe. C. Shuttleworth, P. Lurz and J. Gurnell, European Squirrel Initiative, Stoneleigh Park, Warwickshire CV8 2LG UK: 255-274.

Abstract: The grey squirrel (Sciurus carolinensis) is considered a reservoir host of squirrelpox virus (SQPV) which poses a substantial epizootic threat to the native red squirrel (Sciurus vulgaris) in the United Kingdom (UK). Here we evaluate the potential contribution of ectoparasite vectored transmission of SQPV to the epidemiology of the disease. This includes information on ectoparasite abundance on grey squirrels (ectoparasites were recovered from 73% of grey squirrels examined) and in nest boxes (two species of flea were recovered from both grey and red squirrel nest boxes), along with SQPV sero-prevalence (69% of grey squirrels were sero-positive). These data were used, together with information from the literature, to model SQPV scenarios with and without ectoparasite vectored transmission in grey only, red only and mixed squirrel populations. The model predicted that SQPV cannot be maintained, particularly in grey squirrel populations, without ectoparasite vectored transmission. This hypothesis is testable by experimental manipulation. Such an experiment would involve monitoring changes in disease status in grey squirrel populations following administration of an appropriate insecticide. Such an experiment offers the prospect of developing an innovative means of disease intervention that could potentially eliminate the disease from grey squirrel populations considered to pose particular risks to red squirrel strongholds.

• Dale, T. & J. Chantrey (2016). A subtle endemic virus in grey squirrels; squirrelpox virus. The Grey Squirrel: Ecology & Management of an Invasive Species in Europe. C. Shuttleworth, P. Lurz and J. Gurnell, European Squirrel Initiative, Stoneleigh Park, Warwickshire CV8 2LG UK: 211-233.

Abstract: Alien invasive species are well documented in posing a threat to indigenous species. Disease is increasingly being recognized as a mechanism by which an invasive species can have a negative outcome on native fauna. Where an alien species e.g. grey squirrel (Sciurus carolinensis), acts as a reservoir to introduce a novel pathogen e.g. squirrelpox virus (SQPV) to a naive native species e.g. red squirrel (S. vulgaris), it is important to understand the infection dynamics in the reservoir host. Polymerase chain reaction (PCR) analysis was used to identify SQPV infection in grey squirrels from an opportunistically sampled population in the United Kingdom (UK). This showed the virus to be a localised cutaneous infection, with samples from the lip and forelimb identifying the most infected individuals. Only slight epidemiological trends are observed, supporting SQPV as an endemic infection in grey squirrels in the UK. With only tenuous infection patterns being identified, methods for reducing infection in grey squirrels may prove ineffective in controlling the disease in their red counterpart. The study identifies areas for further investigation, while highlighting the need for large sample sizes when investigating pathogens with only subtle trends.

• Derbridge, J., H. Pepper & J.L. Koprowski (2016). Economic damage by invasive grey squirrels in Europe. The Grey Squirrel: Ecology & Management of an Invasive Species in Europe. C. Shuttleworth, P. Lurz and J. Gurnell, European Squirrel Initiative, Woodbridge, Suffolk UK.: 393-405.

Abstract: Invasive alien species (IAS) comprise a global threat to biodiversity and may also cause economic harm by damaging natural resources and property. Indirect costs associated with IAS control and protection of native species add further economic burdens. The eastern grey squirrel (Sciurus carolinensis) is a successful invader in Europe, and this chapter reviews the current knowledge on the types and scale of damage it has inflicted on the continent’s economies.

• Duff, J. & A. Meredith (2016). Disease and mortality of the grey squirrel The Grey Squirrel: Ecology & Management of an Invasive Species in Europe. C. Shuttleworth, P. Lurz and J. Gurnell, European Squirrel Initiative, Stoneleigh Park, Warwickshire CV8 2LG UK: 153-171.

Abstract: This chapter provides a timely, if non-exhaustive, list of pathogens, parasites and causes of disease and death recorded and published in grey squirrels (Sciurus carolinensis) both in North America (NA) and in Europe (EU). The viruses, bacteria and parasites that have been recorded in grey squirrels are listed, largely from extracts of published reports. Those which cause disease or are considered to be significant for other reasons are described. Mortality as a direct result of control operations, to protect timber crops, property or regional red squirrel (Sciurus vulgaris) populations, are excluded from this review and can be found described in Chapters elsewhere.

• Flaherty, M., E. Goldstein & C. Lawton (2016). Predicting grey squirrel spread in Ireland; spatial tools for 37 future management of an invasive species The Grey Squirrel: Ecology & Management of an Invasive Species in Europe. C. Shuttleworth, P. Lurz and J. Gurnell, European Squirrel Initiative, Stoneleigh Park, Warwickshire CV8 2LG UK: 37-54.

Abstract: The invasive grey squirrel (Sciurus carolinensis) has successfully spread to cover the eastern half of Ireland since its introduction to the Irish midlands in 1911, but it has so far failed to spread west of the River Shannon. The species has declined in parts of its former range but the south, southwest and northwest of the island remain vulnerable to grey squirrel invasion. Managers  and ecologists can use spatial tools to devise a strategic approach to invasive species  management to minimise costs and maximise success. We demonstrate how these tools can be employed to devise a management approach to control the continuing expansion of this non-native species. Suitable habitat beyond the current grey squirrel range has been identified using a species distribution model. Predicted grey squirrel dispersal routes were highlighted using least cost pathway analysis. A spatially explicit population model examined grey squirrel expansion in the south and southwest of the island, where the species continues to spread. Results from discriminant analysis suggested the failure of grey squirrels to expand into certain areas and the recent decline or disappearance in part of their former range were correlated with pine marten (Martes martes) presence and habitat fragmentation. Through these spatial analyses, we identified future monitoring locations and effective control strategies to inform a more strategic management approach to limit the spread of the grey squirrel in Ireland.

• Gurnell, J., P. Lurz & C.M. Shuttleworth (2016). Ecosystem impacts of an alien invader in Europe, the grey squirrel Sciurus carolinensis. The Grey Squirrel: Ecology & Management of an Invasive Species in Europe. C. Shuttleworth, P. W. W. Lurz and J. Gurnell, The Grey Squirrel: Ecology & Management of an Invasive Species in Europe: 307-326.

Abstract: Grey squirrels (Sciurus carolinensis) are native to the eastern part of North America and were first introduced to Great Britain in 1876 and into northern Italy in 1948. From their centres of introduction they have expanded their range resulting in two widely recorded impacts: they outcompete and replace the native red squirrel (S. vulgaris) and damage trees with economic and aesthetic consequences. However, there may be other less apparent or less studied impacts of the alien invader in its new environment, some which may disrupt ecosystems but others that may have positive ecological or socio-economic function, and some that may only become apparent over time. Although invasive grey squirrels fill a similar niche to the usurped red squirrels, they are larger, live at higher densities in broadleaf woodlands, preferentially exploit the seed crops of tree species differently and are more terrestrial. The forest and woodlands they have moved into also differ to their homeland, for example, in terms of tree species richness and diversity. These factors together with the uncertainties of, for example, the effects of climate change on habitats, make it difficult to predict the importance of some of the less studied impacts. These may become more apparent in time and particularly if grey squirrels continue to expand their range in Europe. In this article, we overview the range of ecosystem impacts of invasive grey squirrels in Europe and, based on our current knowledge, provide a classification of the impact mechanism, the outcome (whether positive or negative) and whether the impact is widespread or occurs locally within the new range of grey squirrels. In many cases, it is clear that we do not have enough information to assess the level of importance of a particular impact and we acknowledge that the classification is provisional and likely to change. Nevertheless, we hope it offers a framework to direct further study to fill in the gaps in our knowledge

• Gurnell, J. & H. Pepper (2016). The control and management of grey squirrel populations in Britain. The Grey Squirrel: Ecology & Management of an Invasive Species in Europe. C. Shuttleworth, P. Lurz and J. Gurnell, European Squirrel Initiative, Stoneleigh Park, Warwickshire CV8 2LG UK: 407-436.

Abstract: Grey squirrels (Sciurus carolinensis) were first introduced from North America to Britain in 1876 and further introductions and translocations within the country were recorded until 1929. Grey squirrels spread from these early points of introduction and are now found throughout much of the country. As early as 1917, concerns were expressed about the possible impact of grey squirrels on native red squirrels (S. vulgaris), to be followed in the 1920s by interest in impacts of grey squirrels on forestry. By the 1930s, the concerns of many landowners had translated into how to manage or control grey squirrels on their land. At the time, the British Government was sympathetic but did not campaign to eradicate grey squirrels; a Grey Squirrel (Prohibition of Importation and Keeping) Order came in to effect in 1937. After the Second World War the control and management of grey squirrels took on greater importance, particularly with respect to tree damage prevention. Extensive control of grey squirrels for red squirrel conservation started later, in the 1990s. In this paper, we discuss the management of grey squirrels with particular reference to Britain since the Second World War. First, we consider management strategies, including the importance of monitoring and bounty schemes that took place in the 1950s. We then discuss different control techniques (shooting, cage trapping, kill trapping, habitat management) and their effectiveness using examples from the literature that provide data on control effort. Now that warfarin poison can no longer be used as a plant protection product, grey squirrel control for both tree damage prevention and red squirrel conservation will depend on trapping and shooting for the foreseeable future. Cage trapping is the preferred method for control at the landscape level, backed up where possible by opportunistic shooting or shooting at feeders. To this end, we provide information on cage trapping best practice and consider the costs and the sociological dimension to the management of grey squirrels, particularly now that large numbers of volunteers are involved in controlling grey squirrels for red squirrel conservation

• Halliwell, E., C. Shuttleworth, S. Cartmel, I. Lloyd & R. Jenkins (2016). Experiences of grey squirrel management in an upland conifer forest. The Grey Squirrel: Ecology & Management of an Invasive Species in Europe. C. Shuttleworth, P. Lurz and J. Gurnell, European Squirrel Initiative, Stoneleigh Park, Warwickshire CV8 2LG UK: 453-472.

Abstract: The introduced grey squirrel (Sciurus carolinensis) has replaced the native red squirrel (Sciurus vulgaris) across large parts of its range in the British Isles. As this replacement took place, red squirrels appeared to have a competitive advantage in upland Sitka spruce dominated forests. Consequently in mainland Britain large areas of upland coniferous plantation were strategically designated as red squirrel strongholds in what has been described as a ‘conifer forest management strategy’. Clocaenog forest in north Wales was one such site where habitat management, in conjunction with grey squirrel control, was used as the approach to safeguard the nationally important red squirrel population. In this paper we review twenty years of data gathered during research and control operations to examine the pattern of grey squirrel incursion and population establishment in this upland spruce plantation. Our findings suggest that grey squirrels may be able to utilise such habitats to a greater extent than previously considered and we examine the implications for red squirrel conservation strategies

• Hayssen, V. (2016). Reproduction in grey squirrels: from anatomy to conservation The Grey Squirrel: Ecology & Management of an Invasive Species in Europe. C. Shuttleworth, P. Lurz and J. Gurnell, European Squirrel Initiative, Stoneleigh Park, Warwickshire CV8 2LG UK: 115-180.

Abstract: Grey squirrels (Sciurus carolinensis) are familiar denizens of many temperate woodlands. When introduced to new areas their reproductive productivity threatens native species, such as the Eurasian red squirrel (Sciurus vulgaris). Thus their reproductive biology is important to conservation efforts. Anatomy and physiology are two components of reproductive biology. Knowledge of reproductive anatomy lets scientists identify reproductive condition while understanding the reproductive cycle helps managers know when to time efforts for population control. This chapter reviews what is currently known about the reproductive biology of grey squirrels from their anatomy and physiology to the timing of their reproductive patterns. It ends with a brief review of current efforts to use contraception to control squirrel numbers.

• Jones, H., A. White, P.W.W. Lurz, M. Boots, & C.M. Shuttleworth (2016). Mathematical models of grey squirrel invasion: a case study on Anglesey. The Grey Squirrel: Ecology & Management of an Invasive Species in Europe. C. Shuttleworth, P. Lurz and J. Gurnell, European Squirrel Initiative, Stoneleigh Park, Warwickshire CV8 2LG UK: 235-252.

Abstract: A spatial, stochastic model to represent the dynamics of red squirrels (Sciurus vulgaris) and grey squirrels (Sciurus carolinensis) and squirrelpox infection is developed to understand the documented replacement of red squirrels by grey squirrels which occurred on the Isle of Anglesey between approximately 1966 and 1998. The model results compare well with the observed historical field data and indicate that competition was the key process responsible for red squirrel decline. Squirrelpox virus (SQPV), which was prevalent in grey squirrels both on the mainland and latterly as they colonized Anglesey, failed to spread extensively through the resident red squirrel populations on the island. Model findings showed that disease outbreaks may occur in relatively high density red squirrel populations at the local level where reds were sympatric to greys, but at the larger scale, red squirrel densities were too low and/or too fragmented to maintain continual intra-specific spread of pathological infection. This finding has important consequences for applied conservation management and suggests that pathological squirrelpox outbreaks may be localised and the risk of extensive squirrelpox spread through low density red squirrel populations may be low

• Koprowski, J., K. Munroe & A. Edelman (2016). Gray not Grey: Ecology of Sciurus carolinensis in native range in North America The Grey Squirrel: Ecology & Management of an Invasive Species in Europe. C. Shuttleworth, P. Lurz and J. Gurnell, European Squirrel Initiative, Stoneleigh Park, Warwickshire CV8 2LG UK: 1-17.

Abstract: Eastern grey squirrels (Sciurus carolinensis) have been introduced to numerous locations around the globe and are considered to be one of the world’s worst invasive species. The species has become notorious for its role as a biological invasive and a pest outside of its natural distribution. The basic ecology of the species does not appear to differ dramatically between native and naturalized lands. However, in deciduous and mixed forests of eastern North America to which they are native, eastern grey squirrels are a welcome species that are rarely  considered problematic. Herein, we examine the ecology of eastern grey squirrels in their native habitats where problems of overpopulation, damage or ecological dominance are rarely encountered.

• Lawton, C., C. Shuttleworth &n R.E. Kenward (2016). Ranging Behaviour, Density And Social Structure In Grey Squirrels The Grey Squirrel: Ecology & Management of an Invasive Species in Europe. C. Shuttleworth, P. Lurz and J. Gurnell, European Squirrel Initiative, Stoneleigh Park, Warwickshire CV8 2LG UK: 133-150.

Abstract: The home range is a settled area in which an animal conducts its daily activities such as feeding, resting and breeding. Within the home range, the animal will have core areas; small  regions where it spends a disproportionate amount of time. There is considerable variation in grey squirrel home range size, as ranging behaviour is influenced by a number of factors. These may be environmental in nature (e.g. habitat type, food availability, degree of fragmentation), or related to population demographics and social interactions. Seasonal variation in home range use is also evident, and can be related to changes in food availability, or intra-specific interactions during breeding or dispersal. Grey squirrels (Sciurus carolinensis) display a hierarchical social  system, with males usually dominant over females, and adults over subadults. Females will  defend exclusive core areas from unrelated females, but overlap in home range core area is tolerated by males. Information on ranging behaviour gained through telemetry studies has proved to be essential in evolving grey squirrel management. It has helped to optimise control programmes and resource use.Everest, D., Floyd, T., Donnachie, B., Irvine, R., Holmes, J., & Shuttleworth, C. (2017). Confirmation of squirrelpox in Welsh red squirrels. Veterinary Record, 181, 514-515.

• Leaver, L. A., Jayne, K., & Lea, S. E. G. (2016) Behavioral flexibility versus rules of thumb: how do grey squirrels deal with conflicting risks? Behavioral Ecology 28: 186-192.

Abstract: In order to test how flexibly animals are able to behave when making trade-offs that involve assessing constantly changing risks, we examined whether wild Eastern grey squirrels showed flexibility of behavioral responses in the face of variation in 2 conflicting risks, cache pilferage, and predation. We established that cache pilferage risk decreased with distance from cover and was thus negatively correlated with long-term predation risk. We then measured changes in foraging and food-caching behavior in the face of changes in the risk of predation and food theft over a short time-scale. We found that, overall, squirrels move further away from the safety of cover when they cache, compared to when they forage, as predicted by pilferage risk. However, there was no effect of immediate pilferage or predation risk (i.e., the presence of potential predators or pilferers) on the distance from cover at which they cached, and only a slight increase in forage distance when predation risk increased. These results suggest that “rules of thumb” based on static cues may be more cost-effective for assessing risk than closely tracking changes over time in the way suggested by a number of models of risk assessment.

• Macpherson, J., H. Denman, D. Tosh, C. McNicol & E.C. Halliwell (2016). A review of the current evidence for impacts of the pine marten (Martes martes) on non-native and native squirrel populations. The Grey Squirrel: Ecology & Management of an Invasive Species in Europe. C. Shuttleworth, P. Lurz and J. Gurnell, European Squirrel Initiative, Stoneleigh Park, Warwickshire CV8 2LG UK: 289-304.

Abstract: The interaction between the European pine marten (Martes martes) and the American eastern grey squirrel (Sciurus carolinensis) has recently become the subject of much public and media interest in the United Kingdom (UK) and Ireland. The range of the grey squirrel has only coincided with the range of the European pine marten in recent years in both Ireland and Scotland. In both countries, native pine marten populations have begun to recover following historical persecution and grey squirrels are an introduced, invasive species. Distributional evidence from a study in Ireland has provided a first attempt to substantiate the potential suppressant effect pine martens could have on grey squirrel populations, with a subsequent positive impact on the native red squirrel (Sciurus vulgaris). Studies in the UK and from elsewhere show that the interactions between martens and squirrels are clearly complex and influenced by a number of different factors. These are reviewed and the implications for both red and grey squirrels discussed, along with the need for further research

• Merrick, M., K. Evans & S. Bertolino (2016). Urban grey squirrel ecology, associated impacts, and management challenges The Grey Squirrel: Ecology & Management of an Invasive Species in Europe. C. Shuttleworth, P. Lurz and J. Gurnell, European Squirrel Initiative, Stoneleigh Park, Warwickshire CV8 2LG UK: 57-77.

Abstract: Grey squirrels (Sciurus carolinensis) are common inhabitants of wooded urban and suburban parks throughout their native and introduced range. The ecology of grey squirrels in rural environments has been the focus of considerable research, yet the ecology, behaviours, economic impact, and conservation implications of urban grey squirrels continue to gain increasing attention. In this chapter, we summarise key ecological characteristics of grey squirrels within an urban/suburban environment and how these differ from those observed in rural environments, the ecological role grey squirrels play in an urban ecosystem, and associated management challenges. Whilst urban and rural grey squirrels select similar habitats, urban  populations can occur at much higher densities than their rural counterparts, from which they  exhibit behavioural differences. Urban grey squirrels provide a wide range of ecosystem services, including cultural services that result in many urban dwellers having positive attitudes towards grey squirrels. This presents challenges for managing conflicts that can arise due to the damage that grey squirrels can cause to infrastructure, vegetation and other wildlife. Understanding basic  ecology and population dynamics of urban grey squirrels, particularly in their introduced range, is essential for predicting their risk of invasion and spread, impacts on native wildlife, and for designing control programmes.

• Merrick, M. J., & Koprowski, J. L. (2016) Altered natal dispersal at the range periphery: The role of behavior, resources, and maternal condition. Ecology and Evolution n/a-n/a. doi:10.1002/ece3.2612.

Abstract: Natal dispersal outcomes are an interplay between environmental conditions and individual phenotypes. Peripheral, isolated populations may experience altered environmental conditions and natal dispersal patterns that differ from populations in contiguous landscapes. We document nonphilopatric, sex-biased natal dispersal in an endangered small mammal, the Mt. Graham red squirrel (Tamiasciurus hudsonicus grahamensis), restricted to a single mountain. Other North American red squirrel populations are shown to have sex-unbiased, philopatric natal dispersal. We ask what environmental and intrinsic factors may be driving this atypical natal dispersal pattern. We test for the influence of proximate factors and ultimate drivers of natal dispersal: habitat fragmentation, local population density, individual behavior traits, inbreeding avoidance, competition for mates, and competition for resources, allowing us to better understand altered natal dispersal patterns at the periphery of a species’ range. A juvenile squirrel’s body condition and its mother’s mass in spring (a reflection of her intrinsic quality and territory quality) contribute to individual behavioral tendencies for movement and exploration. Resources, behavior, and body condition have the strongest influence on natal dispersal distance, but affect males and females differently. Male natal dispersal distance is positively influenced by its mother’s spring body mass and individual tendency for movement; female natal dispersal distance is negatively influenced by its mother’s spring body mass and positively influenced by individual tendency for movement. An apparent feedback between environmental variables and subsequent juvenile behavioral state contributes to an altered natal dispersal pattern in a peripheral population, highlighting the importance of studying ecological processes at the both range center and periphery of species’ distributions.

• Millens, C., D. Everest & A. Brereton (2016). The role of invasive grey squirrels as hosts of selected viruses, bacteria and parasites; implications for conservation and public health. The Grey Squirrel: Ecology & Management of an Invasive Species in Europ. C. Shuttleworth, P. Lurz and J. Gurnell, European Squirrel Initiative, Stoneleigh Park, Warwickshire CV8 2LG UK: 173-190.

Abstract: The introduction of invasive vertebrate hosts can result in changed parasite dynamics, with implications for native species conservation and human health. While the association of invasive grey squirrels (Sciurus carolinensis) and squirrelpox virus (SQPV) with red squirrel (S. vulgaris) decline in the United Kingdom (UK) is well documented, the effect of grey squirrel introduction on the dynamics of other infections and parasites has been less intensively studied. Here, we review current knowledge of invasive grey squirrels as hosts of selected viruses (adenovirus and rotavirus), bacteria (Borrelia burgdorferi sensu lato and Bartonella) and protozoa (Hepatozoon) to assess potential impacts on red squirrels and public health. We describe the known distribution of these organisms in grey and red squirrels in Europe, pathogenicity and methods of detection. We found that knowledge of each pathogen’s distribution and assessment of the risk of cross-species transmission are limited by the opportunistic nature of sampling and limited molecular strain typing. For all organisms, sampling from sympatric grey and red squirrel populations and applying molecular strain typing will improve knowledge for potential crossspecies transmission and reservoir host associations. In the context of theselected viruses, bacteria and protozoa considered in this chapter (SQPV is considered separately in this book), adenovirus is associated with the most significant pathogenic effects and mortality in red squirrels and should therefore be a priority for further research

• Nichols, C. & R. Gill (2016). Bark stripping behaviour by the grey squirrel, Sciurus carolinensis. The Grey Squirrel: Ecology & Management of an Invasive Species in Europe. C. Shuttleworth, P. Lurz and J. Gurnell, European Squirrel Initiative, Stoneleigh Park, Warwickshire CV8 2LG UK: 369-390.

Abstract:Grey squirrels (Sciurus carolinensis) damage trees by stripping bark and ingesting the underlying phloem. Damage has negative impacts on forest regeneration, and reduces timber value, imposing an economic toll on forestry in the United Kingdom (UK). A literature review was conducted to explore bark stripping behaviour. Papers were categorized according to their focus to analyse shifting research attitudes. Factors affecting damage likelihood are well known. Tree physiology, e.g. phloem width, age, species and habitat characteristics can affect the probability of damage, and the main bark stripping season is April to July. The motivations for grey squirrels to strip bark are still unclear. Putative motivators for damage include a fondness for sugar, phloem as a food source, agonistic encounters, and calcium deficiency. Over time, the bark stripping literature has shifted focus from research into predictive factors and causes, to monitoring impacts and improving squirrel control. Returning research focus to underlying causes of bark stripping may aid the production of humane, low-cost, low-effort preventive methods. Current and future research will focus on exploring the calcium hypothesis and investigating the effects of the pine marten (Martes martes) on grey squirrels. The potential impact of grey squirrels in Europe makes research into preventative measures urgent

• Romeo, C., L. Wauters & N. Ferrari (2016). Parasites of grey squirrels: an additional threat to red squirrels in Italy? The Grey Squirrel: Ecology & Management of an Invasive Species in Europe. C. Shuttleworth, P. W. Lurz and J. Gurnell, European Squirrel Initiative, Stoneleigh Park, Warwickshire CV8 2LG UK: 193-209.

Abstract: Parasitic infections in north Italian populations of alien grey squirrels (Sciurus carolinensis) and native red squirrels (S. vulgaris) were investigated to (i) verify whether the invader lost, introduced or acquired any parasite species following its introduction into the new range and (ii) detect any alteration to native hosts’ parasite community induced by grey squirrels. Overall, results of field studies show that introduced grey squirrels harbour an impoverished parasite community compared to their native range and this reduction in parasite pressure may have facilitated their establishment and spread, as postulated by the enemy-release hypothesis. However, the invader introduced to Italy the North American nematode Strongyloides robustus which successfully spilled over to potentially naive red squirrels, as revealed by subsequent analysis of native hosts co-inhabiting with the alien congener. In addition, where grey squirrels are present, a significant increase in prevalence of infection by a local parasite Trypanoxyuris sciuri has been observed, suggesting that competition with invaders may indirectly reduce red squirrels’ ability to cope with parasitic infections. These findings highlight the need to further investigate the role of parasites in the red-grey squirrel system, both in the context of enemy-release facilitating invasion and as an added threat to red squirrel conservation

• Scheibe, J. & N. Moncrief (2016). Morphometric divergence and functional similarity in Sciurus vulgaris and Sciurus carolinensis The Grey Squirrel: Ecology & Management of an Invasive Species in Europe. C. Shuttleworth, P. Lurz and J. Gurnell, European Squirrel Initiative, Stoneleigh Park, Warwickshire CV8 2LG UK: 79-95.

Abstract: We used geometric morphometric techniques to explore and compare th shapes of dentaries and skulls in four species of tree squirrels: Eurasian red squirre (Sciurus vulgaris), eastern grey  squirrel (S. carolinensis), eastern fox squirre (S. niger) and western grey squirrel (S.  griseus). These species were chose because of current competitive interactions amongst the species, and becaus of their  phylogenetic affinities. A canonical variates analysis of  Procruste shape coordinates revealed significant shape differences between the skull and  dentaries of S. carolinensis and S. vulgaris. We compared biomechanica properties of the dentaries for the four species, and used discriminant functio analysis to discriminate between the species in a jaw-function space. Here there was extensive functional overlap between S. carolinensis and S. vulgaris but not between S. carolinensis and S. niger. Although the skulls and dentarie of S. carolinensis and S. vulgaris differ morphologically, they are functionall similar.

• Shuttleworth, C., E. Halliwell & P.A. Robertson (2016). Identifying incursion pathways, early detection responses and management actions to prevent grey squirrel range expansion: an island case study in Wales. The Grey Squirrel: Ecology & Management of an Invasive Species in Europe. C. Shuttleworth, P. Lurz and J. Gurnell, European Squirrel Initiative, Stoneleigh Park, Warwickshire CV8 2LG UK: 475-492.

Abstract: Grey squirrels (Sciurus carolinensis) are a non-indigenous species in the United Kingdom (UK) and were eradicated from the island of Anglesey in 2013 through a live-trapping based control programme. In parallel with the eradication effort, native red squirrel (Sciurus vulgaris) population restoration occurred. Red squirrels are a charismatic species, popular with the general public, and consequently people frequently provide supplemental foods for animals on garden bird tables or within squirrel feeding hoppers. The existence of these feeding stations, along with online social media platforms encouraging the reporting of squirrel sightings, resulted in the progressive evolution of a ‘community based’ network for monitoring squirrels. It was anticipated that this level of surveillance would lead to the early detection of grey squirrel incursion onto the island. Incursion contingency plans were drafted in 2008 and annual proactive surveillance trapping was established within woodland near the railway bridge linking Anglesey with the mainland. Against this background, a series of grey squirrel sightings were reported by the public during the autumn of 2015. This information led to the capture of three individuals. However, reporting by the public also included the misidentification of red squirrels as grey squirrels (false positives), unnecessarily increasing trapping efforts and highlighting a confounding factor in landscapes with sympatric squirrel populations. In this Chapter, we critically review the efficiency of reporting mechanisms, responses to sightings and factors affecting interventions. Decisions with regards to the partitioning of resources in response to grey squirrel sightings were influenced by several factors. A targeted response had to balance an array of available sighting evidence, information that often related to geographically discrete areas which,  on Anglesey, precluded simultaneous action within separate locations. This ultimately meant that managers had to implement a sequential prioritisation of removal measures

• Shuttleworth, C., V. Selonen & J.L. Koprowski (2016). Grey squirrel nesting ecology and the use of nest sites in European population management. The Grey Squirrel: Ecology & Management of an Invasive Species in Europe. C. Shuttleworth, P. Lurz and J. Gurnell, European Squirrel Initiative, Stoneleigh Park, Warwickshire CV8 2LG UK: 349-367.

Abstract: Grey squirrels (Sciurus carolinensis) are an arboreal species that construct leaf nests (dreys) high in the woodland canopy. They will also den within tree cavities and inside artificial den sites such as wooden boxes. Occasionally, individuals will build a nest within the eaves or attic roof spaces of buildings. In this paper, we review the nesting behaviour of grey squirrels, examine nest site occupancy as a means of monitoring population abundance, review the control of grey squirrels whilst using different nest sites, and consider how an understanding of nesting behaviour might help evolve grey squirrel control programmes in Europe. Although there are several detailed studies of nest box use in North American populations, across Europe there remain only limited data with respect to artificial dens. We report nest box use by grey squirrels in Britain before highlighting opportunities for future ecological study, which may assist in managing this invasive pest species. In this context, we make reference to findings of long-term nest box studies of European red squirrel (S. vulgaris) and flying squirrels that require replication.

• Signorile, A. & C. M. Shuttleworth (2016). Genetic evidence of the effectiveness of grey squirrel control operations: lessons from the Isle of Anglesey. The Grey Squirrel: Ecology & Management of an Invasive Species in Europe. C. Shuttleworth, P. Lurz and J. Gurnell, European Squirrel Initiative, Stoneleigh Park, Warwickshire CV8 2LG UK: 439-451.

Abstract: Eradication and control of Invasive Alien Species (IAS) populations presents a major challenge to the preservation of natural environments. Issues relating to the costs, logistics and social acceptance of eradications can undermine the success of management interventions and lead to failures. In this study, we examine grey squirrel genetics over a seven year period during which an island population was subjected to an eradication programme. The aim was to assess temporal changes in genetic diversity, population structure and inbreeding levels and also, to detect gene flow from the mainland. Our data show that the reduction in the number of individuals present on the island led to a marked genetic disequilibrium, reduced genetic diversity, high levels of inbreeding and clustering of individuals in fragmented populations. A very moderate gene flow was detected from the mainland to the island. These results indicate that intensive and continuous culling operations can disrupt the genetic patterns of a thriving invasive population and accelerate eradication even in non-isolated populations. The results also suggest that population genetics can be applied as an effective tool in investigating the ongoing success of an eradication process

• Steele, M. & L.A. Wauters (2016). Diet and food hoarding in eastern grey squirrels (Sciurus carolinensis): implications for an invasive advantage The Grey Squirrel: Ecology & Management of an Invasive Species in Europe. C. Shuttleworth, P. Lurz and J. Gurnell, European Squirrel Initiative, Stoneleigh Park, Warwickshire CV8 2LG UK: 97-113.

Abstract: The invasive advantage of the eastern grey squirrel (Sciurus carolinensis) in its introduced range of Europe likely follows from its broad, opportunistic diet, and its ability to scatter-hoard seeds of many hardwood species such as oaks (Quercus spp.) and to manage these resources through periods of food scarcity, even in the face of intense competition with other rodents. We review the dietary patterns reported for this species, the seasonal and geographic variation in the diet, and the limited information on diet breadth of grey squirrels in Europe. We also describe the behaviour of scatter-hoarding by grey squirrels, and how seed characteristics such as seed perishability (due to germination schedules or insect infestation), seed size and seed chemistry influence scatter-hoarding decisions. We discuss the close evolutionary relationship between grey squirrels and the oaks, as evidenced by the innate basis of embryo removal, which significantly extends storage time. We also highlight recent studies that demonstrate that grey squirrels can maintain a hoarder’s advantage—in part, by deceiving potential pilferers about cache locations—but also by storing seeds in open habitats where pilferage is reduced. We conclude that many of the adaptive strategies this species exhibits for storing seeds may provide it with a competitive advantage in an introduced setting

• Sullivan, S., A. Sullivan & J. S. Brown (2016). Citizen Scientists: fundamental partners in squirrel monitoring. The Grey Squirrel: Ecology & Management of an Invasive Species in Europe. C. Shuttleworth, P. Lurz and J. Gurnell, European Squirrel Initiative, Stoneleigh Park, Warwickshire CV8 2LG UK: 329-346.

Abstract: Monitoring squirrel populations across broad areas can be challenging especially in the patchwork of anthropogenic environments that range from urban neighbourhoods to rural tree plantations. The rapid and unpredictable change of such habitats, along with legal, logistical, and physical barriers to workers can hinder traditional data collection. Project Squirrel was designed as a way to overcome these barriers while also making connections between non-scientists and their local ecology. Fluid and rapidly changing populations in human-dominated habitats are best tracked by citizen scientists. These enthusiastic volunteers can provide accurate and timely observations from a greater number of locations than could ever be visited by professional scientists or covered by autonomous data collection devices. Repeated observations from the same location provide a timely, low-cost understanding of species occurrence and change in distribution. Sciurids are particularly amenable to observation by the general public because they are primarily diurnal, charismatic and are readily identified. Citizen scientists are self-selecting individuals and generally well aware of their own degree of environmental knowledge (70% vs 56% for reference group). However, they are a diverse group, so data collection methodologies must accommodate non-specialist abilities. Formative evaluation and periodic review of data collection tools can facilitate their accurate use by participants. Focused training to develop skill sets beyond the common knowledge of the general public may be necessary. Similarly, evaluation of participant skills allows confidence in the accuracy of submitted data. Intense and repeated participant recruitment is necessary to ensure an adequate number of data submissions over time. Through careful study design and standardized data collection practices, citizen scientists can be powerful partners to effectively monitor the ecological change occurring in their neighborhoods that, in turn, may impact a region or even a country.

• Shuttleworth, C., P. Lurz & J. Gurnell (2016). Grey squirrel management in Europe – the future. The Grey Squirrel: Ecology & Management of an Invasive Species in Europe. C. Shuttleworth, P. Lurz and J. Gurnell, European Squirrel Initiative, Stoneleigh Park, Warwickshire CV8 2LG UK: 517-519.

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Journal references

• Avanzi C, del-Pozo J, Benjak A, Stevenson K, Simpson VR, Busso P, et al. (2016) Red squirrels in the British Isles are infected with leprosy bacilli. Science. 354(6313): 744-7.

Summary: With the exception of armadillos in the Americas, leprosy infections are considered almost exclusively restricted to humans. Avanzi et al. examined warty growths on the faces and extremities of red squirrels in the British Isles and found that two species of leprosy-causing organisms were to blame. Mycobacterium leprae in the southern population of Brownsea Island squirrels originated from a medieval human strain. M. lepromatosis was found in red squirrels from elsewhere in the United Kingdom and Ireland. Human leprosy is proving hard to eradicate, despite available drugs. Perhaps other wildlife species are also reservoirs for this stubborn disease. Leprosy, caused by infection with Mycobacterium leprae or the recently discovered Mycobacterium lepromatosis, was once endemic in humans in the British Isles. Red squirrels in Great Britain (Sciurus vulgaris) have increasingly been observed with leprosy-like lesions on the head and limbs. Using genomics, histopathology, and serology, we found M. lepromatosis in squirrels from England, Ireland, and Scotland, and M. leprae in squirrels from Brownsea Island, England. Infection was detected in overtly diseased and seemingly healthy animals. Phylogenetic comparisons of British and Irish M. lepromatosis with two Mexican strains from humans show that they diverged from a common ancestor around 27,000 years ago, whereas the M. leprae strain is closest to one that circulated in Medieval England. Red squirrels are thus a reservoir for leprosy in the British Isles.

Biancardi, C., & Gnoli, C. (2016) A review of Sciurus Group studies on the red squirrel (Sciurus vulgaris): presence, population density and colour phases in Lombardy (Italy). Natural History Sciences. Atti Soc. it. Sci. nat. Museo civ. Stor. nat. Milano 3: 27-34.

Abstract: During the nineties of the last Century, under the inspiration  of  Luigi  Cagnolaro,  it  had  been  founded  a  “Sciurus  group”, within the Research Centres of the Società Italiana di Scienze Naturali. Aim of the group was to start a campaign of researches on red squirrel populations in Lombardy, Italy promoting field master thesis on various topics: colour  phases and  indirect estimations of population densities. The researches  were carried out  during 10 years  in 5 study  areas. An investigation and census  with questionnaires  were started in  parallel, with  the  collaboration  of  Forest  Guards  (CFS),  Ecological  Guards (GEV) and students of biological and natural sciences. The results were published in Italian or exposed in national and international congresses. This review has the purpose  to show and  critically discuss the overall results, and make them available to a broader audience. The dark colour phase, in  the red  squirrel, resulted  associated to  elevation and  conifer woodlands, according  to the thermoregulation theory, which  consider the dark  fur thicker and more  suitable in  cold environments. Squirrel population  densities  are  subject  to  periodical  fluctuations,  in  conifer woodlands, due  to cones availability. Cone  crops are  subject to cycle with years of great production followed by years of lean. The recorded squirrel densities are  middle-lows, but  in line with  data from  similar environments

• Bisi, F., von Hardenberg, J., Bertolino, S., Wauters, L. A., Imperio, S., Preatoni, D. G., et al. (2016) Current and future conifer seed production in the Alps: testing weather factors as cues behind masting. European Journal of Forest Research, 1-12. doi:10.1007/s10342-016-0969-4.

Abstract: Temporal patterns of masting in conifer species are intriguing phenomena that have cascading effects on different trophic levels in ecosystems. Many studies suggest that meteorological cues (changes in temperature and precipitation) affect variation in seed-crop size over years. We monitored cone crops of six conifer species in the Italian Alps (1999–2013) and analysed which seasonal weather factors affected annual variation in cone production at forest community level. Larch, Norway spruce and silver fir showed masting while temporal patterns in Pinus sp. were less pronounced. We found limited support for the temperature difference model proposed by Kelly et al. Both seasonal (mainly spring and summer) temperatures and precipitations of 1 and 2 years prior to seed maturation affected cone-crop size, with no significant effect of previous year’s cone crop. Next, we estimated future forest cone production until 2100, applying climate projection (using RCP 8.5 scenario) to the weather model that best predicted variation in measured cone crops. We found no evidence of long-term changes in average cone production over the twenty-first century, despite increase in average temperature and decrease in precipitation. The amplitude of predicted annual fluctuations in cone production varies over time, depending on study area. The opposite signs of temperature effects 1 and 2 years prior to seed set show that temperature differences are indeed a relevant cue. Hence, predicted patterns of masting followed by 1 or more years of poor-medium cone production suggest a high degree of resilience of alpine conifer forests under global warming scenario.

• Brady Matthew, J., Koprowski John, L., Gwinn, R. N., Jo, Y.-S., & Young, K. (2016) Eastern fox squirrel (Sciurus niger, Linnaeus 1758) introduction to the Sonoran Desert Mammalia.

Abstract: The eastern fox squirrel, native to the eastern and midwestern United States, was recently documented in the Sonoran Desert in the vicinity of Yuma, Arizona, constituting the first state record for this species. We surveyed the people of Yuma to determine when and how the squirrels arrived. The squirrels were first observed in the 1960s, but may have been resident for a longer period. Since the 1960s, squirrels have spread throughout the city limits and extended south ~15 km into Somerton, Arizona. How the squirrels arrived is not clear, but must be the result of an introduction, as no nearby populations exist. The persistence of eastern fox squirrels in this unique habitat is due to synanthropic relationships.

• Broome, A., Summers, R., & Vanhala, T. (2016) Understanding the provision of conifer seed for woodland species. Forestry Commission Research Note 023. 12 pp.

Abstract: Conifer seed provides an important food resource for many woodland mammals, birds and insects, including some of Britain’s rarest species. This Research Note brings together information from a number of sources on cone and seed production by the main conifers planted in Britain. This information can help managers assess the seed resources of their woodlands and manage the woods for the objective of seed production, whether for food or to encourage natural regeneration. Cone and seed crops fluctuate annually and the amount of seed available in good compared with bad seed years, as well as the frequency of good years, depends on a range of factors which include tree species, age of the crop and climatic conditions. Some species such as Scots pine produce moderate but consistent crops of seed every year, whereas others are much more variable. For example, in a good year Japanese larch can provide the greatest amount of seed and energy per area of woodland of any conifer species grown in Britain, whereas in a poor year production is almost negligible. The time of year when seed is released differs between conifer species. Woodland management can influence the continuity of seed supply as well as the quantities of cones and seed produced. Managing to provide a continuous and abundant seed resource involves consideration of woodland age structure and species composition as well as choice of appropriate interventions.

• Chavel, E. E., Mazerolle, M. J., Imbeau, L., & Drapeau, P. Comparative evaluation of three sampling methods to estimate detection probability of American red squirrels (Tamiasciurus hudsonicus). Mammalian Biology – Zeitschrift für Säugetierkunde.   doi:dx.doi.org/10.1016/j.mambio.2016.11.003.

Abstract: Measuring changes in species distribution and understanding factors influencing site occupancy are recurring goals in wildlife studies. Imperfect detection of species hinders such studies, resulting in the underestimation of the number of sites occupied by the species of interest. American red squirrels (Tamiasciurus hudsonicus) are sampled traditionally with live-traps that require substantial resources to deploy and monitor. Here, we assessed whether auditory methods yield similar detection probabilities. We compared the detection probability of American red squirrels in boreal forest using point counts, playback counts, and live-trapping. Over the summer of 2014, we conducted three trapping sessions in 60 sites within black spruce forests of northwestern Quebec, Canada. We also conducted 10 min point counts in the same sites, together with playback counts using recordings of American red squirrel alarm and territorial calls. Using dynamic occupancy models to analyse three primary periods, all composed of three secondary periods, we found that the detection probability of squirrels from point counts was as high as with live-trapping. Our results thus highlight the value of the point count method in measuring American red squirrel occupancy.

• Dylewski, Ł., Przyborowski, T., & Myczko, Ł. (2016) Winter habitat choice by foraging the red squirrel (Sciurus vulgaris). Annales Zoologica Fennici 53: 194-200.

Abstract: The red squirrel (Sciurus vulgaris) is an arboreal species, relatively common in mixed, deciduous and coniferous forests and in urban parks. From autumn to early spring the main diet of red squirrels is seeds in closed conifer cones. In this study, we investigated characteristics of a habitat in western Poland where red squirrels were feeding on Scots pine (Pinus sylvestris). Understory cover, number of tree species, distance to an open area, distance to the nearest great spotted woodpecker (Dendrocopos major) anvil, and size of ten randomly selected Scots pine trees were measured on 70 transects. We used binary logistic regression to test which forest habitat parameters affected the presence of feeding signs of red squirrels. Feeding signs of red squirrels were found from sites close to forest edges with less understory cover, higher tree-species richness and larger size of trees. Red squirrels did not avoid sites close to great spotted woodpecker anvils. We conclude that forest stand structure is important for red squirrel feeding site occurrence, but red squirrels do not avoid close contact to open areas.

• Fey, K., Hämäläinen, S., & Selonen, V. (2016) Roads are no barrier for dispersing red squirrels in an urban environment. Behavioral Ecology 27: 741-747.

Abstract: In urban environments, roads are one of the major threats for moving animals. Roads can act as barriers to movement either through mortality during crossing attempts or through behavioral avoidance. This can have severe population-level consequences such as population fragmentation and demographic or genetic isolation. A major limitation for determining the effects of roads, however, is the lack of studies on responses of animals to roads during dispersal. In a radiotelemetry study, we investigated the responses of dispersing juvenile red squirrels (Sciurus vulgaris) to roads in an urban environment in Finland. We observed that, during routine movements within their home range, squirrels were located further from roads and crossed them less frequently than simulated random walk paths, while they did not avoid roads when performing explorative and dispersal movements. Moreover, during routine movements squirrels rarely crossed roads with high traffic volume, whereas during dispersal, they crossed both big and small roads. Traffic did not seem to be a major cause of mortality for juvenile squirrels, based on our observations. Our study provides striking evidence that movement behavior during dispersal differs from that of nondispersers. For road ecology, this implies that the evaluation of the role of roads as barriers for, for example, gene flow cannot be based on the interpretation of movements of nondispersers.

• Goldstein, E., Butler, F. & Lawton, C. (2016) Modeling future range expansion and management strategies for an invasive squirrel species. Biological Invasions.

Abstract: Successful management of an invasive species requires in depth knowledge of the invader, the invaded ecosystem, and their interactions. The complexity of the species-system interactions can be reduced and represented in ecological models for better comprehension. In this study, a spatially explicit population model was created using the RAMAS software package to simulate the past and future  invasion dynamics of the eastern grey squirrel (Sciurus carolinensis) in the fragmented habitat in case study areas in Ireland. This invasive squirrel species causes economic damage by bark stripping forest crops and is associated with the decline of its native congener (S. vulgaris). Three combinations of demographic and dispersal parameters, which best matched the distribution of the species shortly after introduction, were used to simulate invasion dynamics. Future population expansion was modeled under scenarios of no control and two different management strategies: fatal culls and immunocontraceptive vaccination programmes. In the absence of control, the grey squirrel range is predicted to expand to the south and southwest of Ireland endangering internationally important habitats, vulnerable forest crops, and the native red squirrel. The model revealed that region-wide intensive and coordinated culls would have the greatest impact on grey squirrel populations. Control strategies consisting solely of immunocontraceptive vaccines, often preferred by public interest groups, are predicted to be less effective. Complete eradication of the grey squirrel from Ireland is not economically feasible and strategic evidence-based management is required to limit further range expansion. Ecological models can be used to choose between informed management strategies based on predicted outcomes.

• Gwinn, R., & Koprowski, J. (2016) Differential response to fire by introduced and endemic species complicates endangered species conservation. Hystrix, the Italian Journal of Mammalogy 27(2): n/a-n/a. DOI: dx.doi.org/10.4404/hystrix-27.2-11447

Abstract: Fire is a natural component of, and serves as a tool for, the restoration of forested ecosystems worldwide; however, disturbance due to fire also has been implicated in the proliferation of invasive species. How these fires affect occupancy and use of the forest by wildlife is of great concern, in particular, the differential response of non-native and native species. In the North American Southwest, prior to European settlement, frequent wildfires helped to maintain forest structure. We examined the effect of a large wildfire on an introduced population of the Abert’s squirrel (Sciurus aberti) that has invaded the high elevation forests inhabited by the critically endangered Mt. Graham red squirrel (Tamiasciurus fremonti grahamensis). We found that introduced Abert’s squirrels were more common than native red squirrels in burned areas. Abert’s squirrels did not abandon burned areas but nested, foraged, and did not adjust their home range size in burned areas. This suggests that invasive Abert’s squirrels are better able to exploit burned areas than native red squirrels and that fire can favor non-native species. This interaction between non-native species, native species, and fire adds new insight into the complexities of conservation and restoration of ecosystems and helps to inform conservation activities worldwide.

• Hyslop, L. (2016) Like rabbit, but sweeter’: the verdict on eating squirrel The Telegraph, 04 April 2016.
• Jessen, T. G., Kilanowski, A. L., Gwinn, R. N., Merrick, M. J., & Koprowski, J. L. (2016) Microsciurus flaviventer (Rodentia: Sciuridae). Mammalian Species 48: 59-65.

Abstract: Microsciurus flaviventer (Gray, 1867) is a Neotropical tree squirrel commonly known as the Amazon dwarf squirrel. Small bodied with dark brown dorsal pelage contrasted with a gray or yellowish gray venter, and a faintly banded tail. M. flaviventer is 1 of 4 species in the genus Microsciurus. The geographic range of M. flaviventer extends from the Amazon basin of South America throughout western and southeastern Colombia, Ecuador, southern Peru, Brazil west of Río Negro, and Madeira. It is most commonly associated with evergreen lowland tropical rainforest. Status of M. flaviventer is “Data Deficient” under the International Union for Conservation of Nature and Natural Resources Red List of Threatened Species; however, loss of habitat is a major concern.

• Koprowski, J., Gavish, L., & Doumas, S. (2016) Sciurus anomalus (Rodentia: Sciuridae). Mammalian Species 48: 48-58.

Abstract: Sciurus anomalus Güldenstädt, 1785 is a rodent commonly called the Caucasian squirrel.  S. anomalus is a medium-sized  squirrel with chestnut gray to grizzled buff dorsum, buff eye rings, and chestnut to buff-yellow underparts.  S. anomalus is 1 of 28  species in the genus  Sciurus and is found in forests of the Middle East and extreme southwestern Asia. The International Union for  Conservation of Nature and Natural Resources considers  S. anomalus to be a species of “Least Concern;” habitat destruction is the  main threat.

• Koprowski, J., Goldstein, E., Bennett, K., & Pereira, C. (2016) Family Sciuridae (Tree, Flying and Ground Squirrels, Chipmunks, Prairie Dogs and Marmots). In D. WIlson, T. Lacher & R. Mittermeier (eds) Handbook of the Mammals of the World – Volume 6: Lagomorphs and Rodents 1. Lynx Edicions in association with Conservation International and IUCN.
• Lioy, S., Mori, E., Wauters, L. A., & Bertolino, S. (2016) Weight operated see-saw feeding hoppers are not selective for red squirrels when greys are present. Mammalian Biology – Zeitschrift für Säugetierkunde  doi:dx.doi.org/10.1016/j.mambio.2016.03.008.

Abstract: The competition for food resources between the native red squirrel and the introduced American Eastern grey squirrel is well known, and can lead to the extinction of the native species. Providing supplementary food resources for the red squirrel, by adopting selective feeding hoppers, has been proposed as a possible support for the short term conservation of native populations, but studies that investigate its effectiveness have not yet been performed. In this study we evaluate the effectiveness of the feeding hoppers, in terms of selectivity towards the smaller native species, quantifying their utilization by the two species in sympatry and allopatry. Feeding hoppers were not selective toward the native species. The success in the attempts to enter the hoppers was 95% for red and 86% for grey squirrels. The 50 hazelnuts provided during each feeding session, covering the energetic requirements of an individual for 6 (reds) or 3.5 (greys) days, were consumed in 45:43 ± 38:26 hh:mm by red and 31:07 ± 37:18 hh:mm by grey squirrels. The average weight of grey squirrels that entered the feeding hopper (490 ± 47 g) was higher than the calibration weight of the see-saw floor (400 g). This highlights that weight operated see-saw feeding hoppers are poorly selective. Structural modification of the feeding hoppers should be considered to obtain a real selectivity according to species before their implementation in the conservation of red squirrel populations. However feeding hoppers were selective in feeding squirrels when using hazelnuts, excluding the access to food supplies by other species. Therefore they can be used with success in supplementary feeding studies or behavioural and ecological studies, especially if combined with camera traps and individually marked animals.

• Krishna, M. C., Kumar, A., Tripathi, O. P., & Koprowski, J. L. (2016) Diversity, Distribution and Status of Gliding Squirrels in Protected and Non-protected Areas of Eastern Himalayas in India. 2016, 27. doi:10.4404/hystrix-27.2-11688.

Abstract: The tropical forests of South and Southeast Asia hold the highest gliding squirrel diversity but our knowledge of species diversity, ecology and major threats is limited. The present study was undertaken in Arunachal Pradesh, Northeast India between June 2011 and March 2015 to address the paucity of data available on gliding squirrels. Based on field and literature surveys, 14 species of gliding squirrels were detected in the state of Arunachal Pradesh. However, species such as Biswamoyopterus biswasi , which is reported as endemic to Namdapha National Park, were not detected. The high gliding squirrel diversity in this region could be related to a diversity of forest types and its location between the Himalayas and the Indomalayan region. Encounter rates with four different species revealed that Petaurista petaurista was most frequently detected in Namdapha National Park. Major threats include hunting for traditional medicine, cultural purposes or bushmeat, and habitat loss due to forest degradation caused by shifting cultivation. In addition, more intensive studies on population, ecology and conservation status are needed in order to design species and site specific conservation action plans in this region which represents the highest diversity of gliding squirrels globally.

• Lurz, P., Shuttleworth, C. & Gurnell, J. (2016) Which trees help red squirrels? Westmorland Red Squirrel Society Newsletter, 6-8.
• Mazzamuto, M. V., Bisi, F., Wauters, L. A., Preatoni, D. G., & Martinoli, A. (2016) Interspecific competition between alien Pallas’s squirrels and Eurasian red squirrels reduces density of the native species. Biological Invasions  1-13. doi:10.1007/s10530-016-1310-3.

Abstract: When alien species introduced into a new environment have a strong niche overlap with ecologically similar native species, interspecific competition can cause a decrease in abundance and distribution of native species. Pallas’s squirrel (Callosciurus erythraeus) was introduced in Northern Italy where it currently co-occurs with native Eurasian red squirrels (Sciurus vulgaris). The alien species is known for its invasiveness but so far negative effects of Pallas’s squirrels on native tree squirrels have not been demonstrated. Here, we compare demographic parameters of red squirrel populations between sites without (red-only sites) and with (red-Pallas’s sites) C. erythraeus and present results of trapping and removal of Pallas’s squirrel and its effects on red squirrel population dynamics. The native species was patchily distributed and absent in many trapping sites occupied by the Pallas’s squirrel. Red squirrels occurred at much lower densities and showed reduced adult survival in areas of co-occurrence than in red-only sites, but there were no differences in reproductive rate. Removing invasive squirrels throughout the study period resulted in re-colonisation by the native species only in some trapping sites, and several alternatives to explain the lack of a marked increase in population size are discussed. This study is the first to provide evidence that presence of Pallas’s squirrel reduces viability of local red squirrel populations.

• Merrick, M. J., & Koprowski, J. L. (2016) Evidence of natal habitat preference induction within one habitat type. Proceedings of the Royal Society B: Biological Sciences 283.

Abstract: Natal habitat preference induction (NHPI) is a mechanism for habitat selection by individuals during natal dispersal. NHPI occurs in wild animal populations, and evidence suggests it may be a common, although little studied, mechanism for post-dispersal habitat selection. Most tests of NHPI examine the influence of distinct, contrasting natal habitat types on post-dispersal habitat selection. We test the hypothesis that NHPI can occur within a single habitat type, an important consideration for habitat specialists. The Mount Graham red squirrel (Tamiasciurus hudsonicus grahamensis) is an endangered forest obligate restricted to a single mountain primarily within mixed-conifer forest. We test for NHPI by comparing intra-individual differences in natal and settlement habitat structure and composition to expected random pairwise differences. Dispersing juveniles appear to select settlement locations that are more similar to natal areas than expected in several forest structure and composition variables that include canopy cover and live basal area. Our results provide support for NHPI as a mechanism for post-dispersal habitat selection in habitat specialists that occupy a single vegetation community type.

• Rocha, R. G., Leite, Y. L. R., Costa, L. P., & Rojas, D. (2016) Independent reversals to terrestriality in squirrels (Rodentia: Sciuridae) support ecologically mediated modes of adaptation. Journal of Evolutionary Biology 29: 2471-2479. doi:10.1111/jeb.12975.

Abstract: The family Sciuridae is one of the most widespread and ecologically diverse lineages of rodents and represents an ideal model for investigating the evolution of locomotion modes and the historical biogeography of terrestrial mammals. We used a comprehensive database on locomotion modes, an updated phylogeny and novel biogeographic comparative methods to reassess the evolution of locomotion of squirrels and to investigate whether these locomotion modes evolved convergently in different continents. We found that locomotion changes occurred in different independent lineages of the family, including four reversals to terrestriality and one evolution of gliding. We also found evidence for Eurasia as the centre of origin of Sciuridae, challenging the classification of the oldest squirrel fossil records from the early Oligocene in North America. Additionally, Eurasia is also the possible centre of origin for most of squirrel subfamilies and tribes, and where locomotion changes have occurred. Parallel locomotion shifts could be explained by the adaptation towards different ecological niches followed by colonization of new continents.

• Selonen, V., Varjonen, R., & Korpimäki, E. (2016) Predator presence, but not food supplementation, affects forest red squirrels in winter. Annales Zoologica Fennici 53: 183-193.

Abstract: We studied the responses of red squirrels, Sciurus vulgaris, to food supplementation and avian predation risk during two winters in Finland, when the level of natural food production, spruce seeds, varied markedly. We performed an open experiment with 6 replicates in the landscape where squirrel signs were counted near a feeding station and/or active predator nest and control site. Within each site, we counted squirrel snow-tracks and feeding-signs and checked the usage of two nest-boxes. We observed that during the winter when natural food was less plentiful, red squirrel snow-tracks decreased at sites with predator. The same was observed, only less clearly, during the winter with more natural food. Food supplementation had no obvious effect. We conclude that predators have a clear effect on red squirrels in winter. Furthermore, in situations where a species is specialized in using fluctuating amounts of natural food resources, the role of artificial food supplementation may remain unclear.

• Selonen, V. & Wistbacka, R. (2016) Siberian flying squirrels do not anticipate future resource abundance. BMC Ecology 16:51. doi:10.1186/s12898-016-0107-7.

Abstract: One way to cope with irregularly occurring resources is to adjust reproduction according to the anticipated future resource availability. In support of this hypothesis, few rodent species have been observed to produce, after the first litter born in spring, summer litters in anticipation of autumn’s seed mast. This kind of behaviour could eliminate or decrease the lag in population density normally present in consumer dynamics. We focus on possible anticipation of future food availability in Siberian flying squirrels, Pteromys volans. We utilise long-term data set on flying squirrel reproduction spanning over 20 years with individuals living in nest-boxes in two study areas located in western Finland. In winter and early spring, flying squirrels depend on catkin mast of deciduous trees. Thus, the temporal availability of food resource for Siberian flying squirrels is similar to other mast-dependent rodent species in which anticipatory reproduction has been observed.

• Selonen, V., Wistbacka, R., & Korpimäki, E. (2016) Food abundance and weather modify reproduction of two arboreal squirrel species. Journal of Mammalogy 97: 1376-1384.

Abstract: The importance of weather in relation to food resources, in determining reproduction, remains poorly understood for mammals, particularly for species that do not depend on food resources limited by spring weather conditions. We studied the effects that weather and food supply had on timing of spring reproduction and observed litter size in 2 squirrel species, the Siberian flying squirrel and the European red squirrel, using long-term data sets spanning 20–30 years. Both species subsist on foods from tree masting, and these are available for squirrels from autumn until early spring. Good food conditions in winter and spring before reproduction had positive effects on spring reproduction in both species by advancing the onset of reproduction, and in flying squirrels, slightly increasing litter size. Higher temperature in late winter and, surprisingly, increased precipitation in late winter resulted in early reproduction in flying squirrels and red squirrels, respectively. In addition, higher early spring temperature was positively related to litter size in red squirrels, likely reflecting low survival of small juveniles in cold weather. Our study supports the view that spring reproduction in these species is determined by food supply before breeding. Our results also highlight the fact that reproduction is also dependent on weather.

• Selonen, V., Wistbacka, R., & Santangeli, A. (2016) Sex-specific patterns in body mass and mating system in the Siberian flying squirrel. BMC Zoology 1:9.  doi:10.1186/s40850-016-0009-3.

Abstract: Reproductive strategies and evolutionary pressures differ between males and females. This often results in size differences between the sexes, and also in sex-specific seasonal variation in body mass. Seasonal variation in body mass is also affected by other factors, such as weather. Studies on sex-specific body mass patterns may contribute to better understand the mating system of a species. Here we quantify patterns underlying sex-specific body mass variation using a long-term dataset on body mass in the Siberian flying squirrel, Pteromys volans.

• Signorile, A. L., Lurz, P. W. W., Wang, J., Reuman, D. C., & Carbone, C. (2016) Mixture or mosaic? Genetic patterns in UK grey squirrels support a human-mediated ‘long-jump’ invasion mechanism. Diversity and Distributions 22: 1-12. doi:10.1111/ddi.12424.

Abstract: Aim: Clarifying whether multiple introductions of a species remain relatively isolated or merge and interbreed is essential for understanding the dynamics of invasion processes. Multiple introductions from different sources can result in a mixture of genetically distinct populations, increasing the total genetic diversity. This mixing can resolve the ‘genetic paradox’, whereby in spite of the relatively small numbers of introduced individuals, the augmented diversity due to this mixing increases adaptability and the ability of the species to spread in new environments. Here, we aim to assess whether the expansion of a successful invader, the Eastern grey squirrel, was partly driven by the merger of multiple introductions and the effects of such a merger on diversity. Location: UK, Ireland. Methods: We analysed the genetic variation at 12 microsatellite loci of 381 individuals sampled from one historical and 14 modern populations of grey squirrels. Results: Our data revealed that current UK population structure resembles a mosaic, with minimal interpopulation mixing and each element reflecting the genetic make-up of historic introductions. The genetic diversity of each examined population was lower than a US population or a historical UK population. Numbers of releases in a county did not correlate with county-level genetic diversity. Inbreeding coefficients remain high, and effective population sizes remain small. Main conclusions: Our results support the conclusion that rapid and large-scale expansion in this species in the UK was not driven by a genetic mixing of multiple introduced populations with a single expansion front, but was promoted by repeated translocations of small propagules. Our results have implications for the management of grey squirrels and other invasive species and also demonstrate how invaders can overcome the genetic paradox, if spread is facilitated by human-mediated dispersal.

• Tagliacozzo, A., Fiore, I., Rolfo, M., & Salari, L. (2016) New data on Late Pleistocene and Holocene red squirrel, Sciurus vulgaris L., 1758, in Italy. Revue de Paleobiologie 35: 417-445.

Abstract: The genus Sciurus is known since Late Miocene in the European fossil record, but it is quite rare. Fossil remains of squirrels occur in relatively few sites and generally with very few specimens, sometimes only one or two teeth. Recent finds of a Sciurus vuslgaris mandible from Grotta Mora Cavorso (Latium), and the reanalysis of the red squirrel remains from the Caverna delle Arene Candide (Liguria), the Riparo Soman (Veneto) and the Grotta del Santuario della Madonna (Calabria) provide new data and insights on the change in size of the rodent and on its geographic and ecological distribution in the Pleistocene and Holocene of Italy. The study of food preferences of the current red squirrel predators provides solid comparative data to measure the relative rarity of the bone remains found in the Late Pleistocene and Holocene paleontological deposits and archaeological contexts. Taphonomic analysis, particularly on the latest Pleistocene remains from the Caverna delle Arene Candide, sheds light on the alimentary and cultural use of the rodent made by prehistoric man, allowing to say that so far the human contribution to the accumulation of his bones has been underestimated.

• Turkia, T., Selonen, V. & Brommer, J.E.  (2016) Large-scale spatial synchrony in red squirrel ( Sciurus vulgaris ) sex ratios. Journal of Mammalogy 97: 744-752.

Abstract: Large-scale studies on population-level sex ratios are few, even though sex ratio is an important determinant of population viability and dynamics. Mechanisms driving large-scale sex ratio variation include spatially autocorrelated resources and scale differences in local versus global Fisherian feedback of the operational sex ratio. In this study, we reanalyze historic data on sex ratios based on 187,404 hunted subadult and adult red squirrels ( Sciurus vulgaris ) capturing spatial sex ratio variation in 50×50 km squares throughout Finland over a period of 8 years. Overall, sex ratio was slightly male biased (50.9%) and relatively more 50×50 km squares showed a bias toward males (19% of squares) compared to females (8% of squares). Sex ratio was spatially autocorrelated at distances up to 200 km and in some years showed a U-shaped pattern: regions that were in close proximity and those that were far apart had similar sex ratios, but regions in-between had opposite sex ratios. We found no evidence that food supply (spruce cone crop) drives regional red squirrel sex ratio. Our findings add to the scarce evidence that vertebrate sex ratios show spatial patterns over large scales.

• Tye, C. A., McCleery, R. A., Fletcher, R. J., Greene, D. U., & Butryn, R. S. (2016) Evaluating citizen vs. professional data for modelling distributions of a rare squirrel. Journal of Applied Ecology n/a-n/a. doi:10.1111/1365-2664.12682.

Abstract: * To realize the potential of citizens to contribute to conservation efforts through the acquisition of data for broad-scale species distribution models, scientists need to understand and minimize the influences of commonly observed sample selection bias on model performance. Yet evaluating these data with independent, planned surveys is rare, even though such evaluation is necessary for understanding and applying data to conservation decisions. * We used the state-listed fox squirrel Sciurus niger in Florida, USA, to interpret the performance of models created with opportunistic observations from citizens and professionals by validating models with independent, planned surveys. * Data from both citizens and professionals showed sample selection bias with more observations within 50 m of a road. While these groups showed similar sample selection bias in reference to roads, there were clear differences in the spatial coverage of the groups, with citizens observing fox squirrels more frequently in developed areas. * Based on predictions at planned field surveys sites, models developed from citizens generally performed similarly to those developed with data collected by professionals. Accounting for potential sample selection bias in models, either through the use of covariates or via aggregating data into home range size grids, provided only slight increases in model performance. * Synthesis and applications. Despite sample selection biases, over a broad spatial scale opportunistic citizen data provided reliable predictions and estimates of habitat relationships needed to advance conservation efforts. Our results suggest that the use of professionals may not be needed in volunteer programmes used to determine the distribution of species of conservation interest across broad spatial scales.

• Uchida, K., Suzuki, K., Shimamoto, T., Yanagawa, H. & Koizumi, I. (2016) Seasonal variation of flight initiation distance in Eurasian red squirrels in urban versus rural habitat. Journal of Zoology 298: 225-231.

Abstract: Urbanization has caused significant behavioural modifications in wild animals. Change in anti-predator behaviour is the most widespread example across different taxa in urban areas, which is probably due to a decrease in predation pressure and habituation towards humans. Seasonality or phenology has also been modified by urbanization since some resources in urban environments are highly controlled, for example, artificial feeding. Under natural conditions, anti-predator responses vary with seasonal variability in environmental and individual conditions. However, resource stability possibly reduces the seasonality of anti-predator behaviours in urban animals. Here, we compare the seasonal difference of flight initiation distance (FID), a measurement of anti-predator response, in Eurasian red squirrels Sciurus vulgaris between urban and rural areas in the Tokachi region, Hokkaido, Japan. Rural squirrels possessed FIDs two to three times longer than those of urban squirrels. We also found squirrels in rural areas lowered FID in autumn, but no seasonal difference was observed in urban squirrels. Our results suggest that continuous supplementary feeding may have buffered the seasonality in anti-predator response. In addition, strong habituation to humans may allow urban red squirrels to correctly assess human activity as benign rather than reacting unnecessarily.

• Xiao, Z., & Zhang, Z. (2016) Contrasting patterns of short-term indirect seed–seed interactions mediated by scatter-hoarding rodents. Journal of Animal Ecology n/a-n/a. doi:10.1111/1365-2656.12542.

Abstract: * It is well known that direct effects of seed predators or dispersers can have strong effects on seedling establishment. However, we have limited knowledge about the indirect species interactions between seeds of different species that are mediated by shared seed predators and/or dispersers and their consequences for plant demography and diversity. Because scatter-hoarding rodents as seed dispersers may leave some hoarded seeds uneaten, scatter hoarding may serve to increase seed survival and dispersal. Consequently, the presence of heterospecific seeds could alter whether the indirect interactions mediated by scatter-hoarding rodents have a net positive effect, creating apparent mutualism between seed species, or a net negative effect, creating apparent competition between seed species. * We present a testable framework to measure short-term indirect effects between co-occurring plant species mediated by seed scatter-hoarding rodents. We tested this framework in a subtropical forest in south-west China using a replacement design and tracked the fate of individually tagged seeds in experimental patches. We manipulated the benefits to rodents by using low-tannin dormant chestnuts as palatable food and high-tannin non-dormant acorns as unpalatable food. * We found that seed palatability changed the amount of scatter hoarding that occurred when seeds co-occurred either among or within patches. Consistent with our predictions, scatter-hoarding rodents created apparent mutualism through increasing seed removal and seed caching, and enhancing survival, of both plant species in mixed patches compared with monospecific patches. However, if we ignore scatter hoarding and treat all seed harvest as seed predation (and not dispersal), then apparent competition between palatable chestnuts and unpalatable acorns was also observed. * This study is the first to demonstrate that foraging decisions by scatter-hoarding animals to scatter hoard seeds for later consumption (or loss) or consume them can influence indirect effects among co-occurring seeds, and rodent-mediated indirect effects vary depending on whether the harvested seeds are hoarded or eaten.

2015

• Blythe, R. M., Lichti, N. I., Smyser, T. J., & Swihart, R. K. (2015) Selection, caching, and consumption of hardwood seeds by forest rodents: implications for restoration of American chestnut. Restoration Ecology 23: 473-481.

Abstract: ecent field trials on blight-resistant hybrids (BC3F3) of American chestnut (Castanea dentata) and Chinese chestnut (C. mollissima) have intensified planning for widespread restoration of Castanea to eastern U.S. forests. Restoration will likely rely on natural seed dispersal from sites planted with chestnut; however, we do not know how dispersal agents such as granivorous rodents will respond to hybrid chestnuts. At one extreme, excessive seed consumption may impede restoration. Alternatively, scatter-hoarding rodents might facilitate the spread of chestnut by dispersal of seeds from restoration plantings. We conducted trials with five rodent species to quantify foraging preferences and to evaluate the potential role of granivores in chestnut restoration. Specifically, we presented seeds from American and hybrid chestnuts (BC3F2) with other common mast species and recorded the proportion of seeds removed and the fates of tagged seeds. Mice, chipmunks, and flying squirrels harvested both chestnut types preferentially over larger, tougher black walnut, hickory, and red oak seeds, but fox squirrels and eastern gray squirrels preferred larger seeds to chestnuts. All rodents consumed a greater proportion of the chestnuts than other seed types. American and hybrid chestnut also differed in important ways: except for fox squirrels, rodents preferentially removed American chestnuts over hybrid chestnuts, but we estimated that fox squirrels carried a greater proportion of hybrid chestnuts beyond our tag search area, suggesting that hybrids may be dispersed farther and cached more often than American chestnut. These differences indicate that hybrid chestnut may not be functionally equivalent to American chestnut with regard to seed–granivore interactions.

• Bosch, S., Spiessl, M., Müller, M., Lurz, P. W. W., & Haalboom, T. (2015) Mechatronics meets biology: experiences and first results with a multipurpose small mammal monitoring unit used in red squirrel habitats.  Hystrix 26:

Monitoring is a fundamental aspect of species conservation and research. Technological advances, especially with respect to camera trap technologies, have allowed glimpses into unknown aspects of species behaviour and have the potential to greatly assist species distribution monitoring. Here we present the findings of a pilot study combining existing biological monitoring techniques with mechatronics to advance monitoring technologies and develop a multi-purpose, species specific, automated monitoring system. We developed a Small Mammal Monitoring Unit (SMMU) that integrates automated video, and sound recording, carries out body weight measurements and takes hairs samples with a bait station in a portable perspex box. The unit has the potential for use with a range of small mammal species, but has been field-tested here on red squirrels, Sciurus vulgaris , in Germany, Scotland and Switzerland. We successfully collected hair-samples, body mass data as well as video and sound recordings. Preliminary data analyses also revealed behavioural information. Heavier individuals first gained access to the feeder in the morning and have longer feeding bouts. Our prototype demonstrated that the collaboration between mechatronic and biology offers novel, integrated monitoring techniques for a range of research application. The development of units for other mammal species is planned. Future developments will explore the possibilities for wireless data transmission, built-in collection of weather data and collection of images from inside the unit for the recognition of individuals.

• Chen, H. & Koprowsk, J. (2015) Animal occurrence and space use change in the landscape of anthropogenic noise. Biologcal Conservation 192: 315-322.

Habitat fragmentation, destruction, and disturbance are major threats to biodiversity. Global road networks represent one of the most significant human impacts on ecosystems, and a spatially extensive source of anthropogenic disturbance and noise. We developed a novel approach by combining traffic monitoring with noise mapping on the basis of a standardized traffic-noise stimulus generated by controlled vehicle operation to investigate temporal and spatial heterogeneity of traffic noise. We used animal presence or absence, radio-telemetric monitoring of space use, and remotely sensed habitat characteristics with occupancy modeling and spatial analysis to assess influences of distance from roads, habitat characteristics, and traffic noise level on site occupancy and space use of Mt. Graham red squirrels (Tamiasciurus hudsonicus grahamensis). Traffic noise had spatially extensive and negative effects on site occupancy. Animal occurrence decreased as traffic noise increased after accounting for distance from roads. Traffic noise levels in animal core home ranges were lower than noise levels within total home ranges. Our study disentangled effects of traffic noise from confounding environmental characteristics and demonstrated the chronic impacts of traffic noise on animal distribution. We highlight the importance of incorporating spatial and temporal heterogeneity of traffic noise at a local scale when investigating effects of anthropogenic noise on wildlife.

• Dozières, A., Pisanu, B., Kamenova, S., Bastelica, F., Gerriet, O. & Chapuis, J-L. (2015) Range expansion of Pallas’s squirrel (Callosciurus erythraeus) introduced in southern France: habitat suitability and space use. Mammalian Biology – Zeitschrift für Säugetierkunde.

Abstract: The study of space use and population density in different habitats is an important step in understanding the expansion process of an introduced species and in gathering useful knowledge for management actions. Pallas’s squirrel (Callosciurus erythraeus) was introduced on the Cap d’Antibes (southeastern France) at the end of the 1960s. We used direct observations from a grid map centered onto the known historical distribution to document the expanding range of Pallas’s squirrel. We assessed habitat suitability in the invaded area through distance sampling and nest counts and examined space use by quantifying the size of the home range and intra- and inter-sexual overlap based on radio-tracking in a suitable habitat. Our results confirm that Pallas’s squirrel has expanded its range exponentially over the past two decades, although with low diffusion coefficients (0.08–0.20 km/yr). Squirrels reached the highest relative densities in suburban woodlands, in which females showed no territorial behaviour. Our results indicate that Pallas’s squirrel was able to establish in various habitats in Antibes, preferentially in woodlands and gardens. Densely urbanised areas and the presence of the highway A8 (E80) acted as barriers that slowed range expansion. Such information is crucial to improve the control program started in 2012 to limit the spread of this potentially invasive squirrel.

• Goldstein, E., Butler, F., & Lawton, C. (2015) Frontier population dynamics of an invasive squirrel species: Do introduced populations function differently than those in the native range? Biological Invasions 17: 1181-1197. doi:10.1007/s10530-014-0787-x.

Abstract: Several squirrel species are biological invaders and their establishment in an area is often marked by ecological and economic costs to native species and forest crops. The eastern grey squirrel (Sciurus carolinensis Gmelin 1788) has been intentionally introduced multiple times outside of its native range but its success in establishing and spreading has not been consistent. An intensive live-trapping programme was designed to investigate the demography and population dynamics of populations of this species on the invasion frontier in the Republic of Ireland, a region marked by the slow but steady invasion of the grey squirrel. Low densities and high breeding rates distinguished these frontier populations. These results were placed in context with other frontier and established grey squirrel populations throughout their introduced and native ranges. As expected, variations in invasion speed and impact severity between regions were reflected in population demography. The highest densities, survival rates and breeding rates were recorded in Britain where the grey squirrel invasion has been most damaging. Careful comparative demographic study of invading populations could improve management outcomes, indicate differential invasibility of invaded communities, and offer clues to enhance the design of conservation reintroduction projects

• Guichón, M. L., Benetiz, V., V, Gozzi, A. C., Hertzriken, M., & Borgnia, M. (2015) From a lag in vector activity to a constant increase of translocations: invasion of Callosciurus squirrels in Argentina. Biological Invasions 17: 2597-2604.

Abstract: Arboreal squirrels of the Asiatic genus Callosciurus have shown high likelihood of establishment from few released animals, in particular, C. erythraeus has established wild populations in Argentina, Belgium, France, Hong Kong, Japan, and The Netherlands. We report the invasion process of C. erythraeus in Argentina in the last four decades and suggest management actions for each foci. Between February 2011 and November 2014 we conducted field surveys and interviews in nine sites in central Argentina to confirm the presence of C. erythraeus, describe their history of introduction, and estimate range expansion and squirrel relative abundance. We report a two decades lag-phase until the onset of translocations of C. erythraeus within national boundaries that resulted in a constant increase of the cumulative number of releases. We confirm nine new release events between 1995 and 2012 and six new invasion foci that yields a total of 13 deliberate releases and 10 invasion foci established in rural and urban areas of Argentina. Spread rate ranged from 0.12 to 0.66 km/year. An intermediate relative density of squirrels (2–7 ind/ha) was found close to release sites except in one case. All introduction events involved squirrels translocated from the first, 40 years old invasion focus, occasionally involving illegal trade. The rate of introduction events in the last decades and the translocation-lag phase described in this study should call the attention in all countries dealing with charismatic, introduced species. Translocation disruption requires urgent attention to slow down the invasion of this and other species.

• Haigh, A.,O’Riordan, R. & Butler, F. (2015) The preference for yew (Taxus baccata) by a red (Sciurus vulgaris) only squirrel population. Wildlife Research 42: 426-436.

Abstract: Context: As invasive grey squirrels continue to spread, red only areas are becoming rarer. It has been reported that red squirrels can outcompete greys only in pure coniferous woodland. In areas of sympatry with grey squirrels, there are concerns about red squirrels’ dependence on certain coniferous tree species in light of recent tree diseases.
Aims: This study aimed to investigate tree selection by red squirrels in an area vulnerable to the spread of the grey squirrel, but currently free of this species.
Methods: During 2013–14, squirrels were trapped and monitored on a 315-ha managed island, with a woodland characterised by a mixture of deciduous and coniferous species. Radio-tracking revealed that squirrels clustered their activity along a network of yew trees, a preference they showed throughout the year. Trap success was also higher in traps placed on yew trees. Yew and beech were selected most commonly, but squirrels were also observed foraging on other items, such as sycamore flowers and lichen. Squirrels spent 35% of their time foraging, utilising the greatest number of tree species in June (n = 13). In spring, squirrels foraged to a greater extent on the ground than in the trees, and exploited a lower number of tree species.
Conclusions: There has been little previous data on the use of yew trees by red squirrels, but they have previously been listed as a species that is preferred by red squirrels rather than greys. This study has further emphasised the importance of this tree species to red squirrels.
Implications: The continued spread of the grey squirrel may lead to red squirrels becoming restricted to areas of intense management such as parks and, accordingly, optimum tree planting is required from the onset for the long-term maintenance of red squirrels. With recent concern about the disease vulnerability of other coniferous species, this study emphasises the relative importance of yew and other tree species in the distribution of red squirrels.

• Holmes, M. (2015) The perfect pest: natural history and the red squirrel in nineteenth-century Scotland. Archives of Natural History 42: 113-125.

Abstract: Following the extirpation of the red squirrel from much of Scotland by the end of the eighteenth century, nineteenth-century naturalists strived to find evidence of its native Scottish status. As medieval accounts and Gaelic place names proved ambiguous, the true extent of the squirrel’s former habitat was a matter of some debate. While numerous reintroductions of the species were made from the late eighteenth century, general enthusiasm for the return of the squirrel quickly turned to dismay, ultimately followed by persecution. If the squirrel originally represented a symbolic mission to rediscover a lost species, the physical animal itself fell below expectations. It became publically perceived as both economically and ecologically destructive. The squirrel was despised by foresters and landowners for damaging trees, while naturalists condemned the species for the destruction of bird’s eggs and nests. This article will investigate naturalists’ quests to rediscover the red squirrel, before examining changing attitudes to the species upon its reintroduction and gradual proliferation. The narrative will emerge through the works and correspondence of Scottish naturalist John Alexander Harvie-Brown (1844–1916) and The new statistical account of Scotland. (1834–1845). The argument will be made that the red squirrel as an object of antiquarian curiosity initially made the species endearing to natural historians, as part of a wider fascination with extinct British fauna. However, the clash between naturalists’ established ornithological interests did little to endear the species to that community, leaving the red squirrel open to a policy of general persecution on economic grounds

• Lane J.E., McAdam A.G., Charmantier A., Humphries M.M., Coltman D.W., Fletcher Q., Gurrell, J.C. & Boutin, S. (2015) Post-weaning parental care increases fitness but is not heritable in North American red squirrels. Journal of Evolutionary Biology  28:1203-12.

Abstract: Most empirical attempts to explain the evolution of parental care have focused on its costs and benefits (i.e., fitness consequences). In contrast, few investigations have been made of the other necessary prerequisite for evolutionary change, inheritance. Here, we examine the fitness consequences and heritability (h2) of a post-weaning parental care behaviour (territory bequeathal) in a wild population of North American red squirrels. Each year, a subset (average across all years = 19%) of reproductive females bequeathed their territory to a dependent offspring. Bequeathing females experienced higher annual reproductive success and did not suffer a survival cost to themselves relative to those females retaining their territory. Bequeathing females thus realized higher relative annual fitness (ω = 1.18 ± 0.03 (SE)) than non-bequeathing females (ω = 0.96 ± 0.02 (SE)). Additive genetic influences on bequeathal behaviour, however, were not significantly different from 0 (h2 = 1.9 x 10-3; 95% Highest Posterior Density Interval = 3.04 x 10-8 to 0.37) and, in fact, bequeathal behaviour was not significantly repeatable (R = 2.0 x 10-3; 95% HPD interval = 0 to 0.27). In contrast, directional environmental influences were apparent. Females were more likely to bequeath in years following low food abundance and when food availability in the upcoming autumn was high. Despite an evident fitness benefit, a lack of heritable genetic variance will constrain evolution of this trait.

• Lucas, J. M., Prieto, P. & Galián, J. (2015) Red squirrels from south–east Iberia: low genetic diversity at the southernmost species distribution limit. Animal Biodiversity and Conservation 38: 129-138.

Abstract: Red squirrels from southeast Iberia: low genetic diversity at the southernmost species distribution limit.— South–east Iberia is the southernmost limit of this species in Europe. Squirrels in the region mainly inhabit coniferous forests of Pinus. In this study, we analyzed the pattern of mitochondrial genetic variation of southern Iberian red squirrels. Fragments of two mitochondrial genes, a 350–base pair of the displacement loop (D–loop) and a 359–bp of the cytochrome b (Cytb), were sequenced using samples collected from 88 road–kill squirrels. The genetic variation was low, possibly explained by a recent bottleneck due to historical over–exploitation of forest resources. Habitat loss and fragmentation caused by deforestation and geographic isolation may explain the strong genetic subdivision between the study regions. Six new haplotypes for the D–loop and two new haplotypes for the Cytb fragments are described. A Cytb haplotype of south–east Iberia was found to be present in Albania and Japan, suggesting local extinction of this haplotype in intermediate areas. No significant clustering was found for the south–east of Spain or for the other European populations (except Calabria) in the phylogenetic analysis.

• Macpherson, M. F., Davidson, R. S., Duncan, D. B., Lurz, P. W., Jarrott, A., & White, A. (2015) Incorporating habitat distribution in wildlife disease models: conservation implications for the threat of squirrelpox on the Isle of Arran. Animal Conservation n/a-n/a.

Abstract: Emerging infectious diseases are a substantial threat to native populations. The spread of disease through naive native populations will depend on both demographic and disease parameters, as well as on habitat suitability and connectivity. Using the potential spread of squirrelpox virus (SQPV) on the Isle of Arran as a case study, we develop mathematical models to examine the impact of an emerging disease on a population in a complex landscape of different habitat types. Furthermore, by considering a range of disease parameters, we infer more generally how complex landscapes interact with disease characteristics to determine the spread and persistence of disease. Specific findings indicate that a SQPV outbreak on Arran is likely to be short lived and localized to the point of introduction allowing recovery of red squirrels to pre-infection densities; this has important consequences for the conservation of red squirrels. More generally, we find that the extent of disease spread is dependent on the rare passage of infection through poor quality corridors connecting good quality habitats. Acute, highly transmissible infectious diseases are predicted to spread rapidly causing high mortality. Nonetheless, the disease typically fades out following local epidemics and is not supported in the long term. A chronic infectious disease is predicted to spread more slowly but can remain endemic in the population. This allows the disease to spread more extensively in the long term as it increases the chance of spread between poorly connected populations. Our results highlight how a detailed understanding of landscape connectivity is crucial when considering conservation strategies to protect native species from disease threats.

• Millins, C., Magierecka, A., Gilbert, L., Edoff, A., Brereton, A., Kilbride, E., et al. (2015) An invasive mammal (grey squirrel, Sciurus carolinensis) commonly hosts diverse and atypical genotypes of the zoonotic pathogen Borrelia burgdorferi sensu lato. Applied and Environmental Microbiology 

Abstract: Invasive vertebrate species can act as hosts for endemic pathogens and may alter pathogen community composition and dynamics. For the zoonotic pathogen Borrelia burgdorferi sensu lato, the agent of Lyme borreliosis, recent work shows invasive rodent species can be of high epidemiological importance and may support host specific strains. This study examined the role of grey squirrels (Sciurus carolinensis), (n=679), an invasive species in the United Kingdom (UK), as B. burgdorferi s.l. hosts. We found that grey squirrels were frequently infested with Ixodes ricinus, the main vector of B. burgdorferi s.l. in the UK, and 11.9% were infected with B. burgdorferi s.l. All four genospecies which occur in the UK were detected in grey squirrels, and unexpectedly, the bird associated genospecies B. garinii, was most common. The second most frequent infection was with B. afzelli. Genotyping of B. garinii and B. afzelli produced no evidence for strains associated with grey squirrels. Generalised linear mixed models (GLMM) identified tick infestation and date of capture as significant factors associated with B. burgdorferi s.l. infection in grey squirrels, with infection elevated in early summer in squirrels infested with ticks. Invasive grey squirrels appear to become infected with locally circulating strains of B. burgdorferi s.l., further studies are required to determine their role in community disease dynamics. Our findings highlight that the role of introduced host species in B. burgdorferi s.l epidemiology can be highly variable and thus difficult to predict.

• Mori, E., Mazzoglio, P., Rima, P., Aloise, G., & Bertolino, S. (2015) Bark-stripping damage by Callosciurus finlaysonii introduced into Italy. Mammalia 

Abstract: The Finlayson’s squirrel Callosciurus finlaysonii was introduced into Italy during the 1980s and has established two viable populations. The diet of this species includes a high proportion of tree barks, suggesting an intensive debarking behavior. We reported a severe bark-stripping impact in both colonized areas, and we tested whether a preference for some tree species existed. Results of this work showed the presence of a wide spectrum of damaged species, without any strong preference, mainly with large wounds. Old deciduous plants and conifers, which presented a hard bark, were usually avoided.

• Palmer, R.R. & Koprowski, J.L. (2015) How do Neotropical pygmy squirrels (Sciurillus pusillus) use seasonally flooded forests in the Peruvian Amazon? Journal of Mammalogy.

Abstract: Tree squirrels are important components of ecosystems but to understand their role, we must learn how squirrels select and use habitat. Tree squirrel species richness is highest in the tropics and, according to the International Union for Conservation of Nature, is also where the greatest number of data deficient species occurs. The Neotropical pygmy squirrel (Sciurillus pusillus) is one of these species. In 2009 and 2010, we conducted distance sampling to estimate population density, we measured vegetation variables to investigate forest characteristics that influence habitat selection at 3 different scales, and conducted observations to obtain knowledge about activity patterns and behavior of Neotropical pygmy squirrels in igapó forest in the Peruvian Amazon. Density of squirrels was 0.10 and 0.14 individuals/ha, respectively, for each year. Squirrel activity peaked at 0800h, individuals were found mainly in the canopy and never on the ground, and frequency of squirrel behaviors differed by time and story level. Squirrels mainly used high and low restinga and areas that had more trees that were ≥ 30cm diameter at breast height (DBH)/ha compared to random areas in our site in igapó forest. Squirrels used trees that were larger in DBH, taller, and had a larger live crown compared to random trees. Neotropical pygmy squirrels are associated with features related to mature forests.

• Pečnerová, P., Moravec, J.C., & Martínková, N. (2015)  A skull might lie: modelling ancestral ranges and diet from genes and shape of tree squirrels.. Systematic Biology 64: 1074-88.

Abstract: Tropical forests of Central and South America represent hotspots of biological diversity. Tree squirrels of the tribe Sciurini are an excellent model system for the study of tropical biodiversity as these squirrels disperse exceptional distances, and after colonizing the tropics of the Central and South America, they have diversified rapidly. Here, we compare signals from DNA sequences with morphological signals using pictures of skulls and computational simulations. Phylogenetic analyses reveal step-wise geographic divergence across the Northern Hemisphere. In Central and South America, tree squirrels form two separate clades, which split from a common ancestor. Simulations of ancestral distributions show western Amazonia as the epicenter of speciation in South America. This finding suggests that wet tropical forests on the foothills of Andes possibly served as refugia of squirrel diversification during Pleistocene climatic oscillations. Comparison of phylogeny and morphology reveals one major discrepancy: Microsciurus species are a single clade morphologically but are polyphyletic genetically. Modelling of morphology-diet relationships shows that the only group of species with a direct link between skull shape and diet are the bark-gleaning insectivorous species of Microsciurus. This finding suggests that the current designation of Microsciurus as a genus is based on convergent ecologically-driven changes in morphology.

• Posthumus E,Koprowski J, Steidl R (2015) Red squirrel middens influence abundance but not diversity of other vertebrates. PLoS One 10: e0123633.

Abstract: Some animals modify the environment in ways that can influence the resources available to other species. Because red squirrels (Tamiasciurus hudsonicus) create large piles of conifer-cone debris (middens) in which they store cones, squirrels concentrate resources that might affect biodiversity locally. To determine whether other animals are attracted to midden sites beyond their affinity for the same resources that attract red squirrels, we assessed associations between middens, mammals, and birds at population and community levels. We surveyed 75 middens where residency rates of red squirrels varied during the previous five years; sampling along this residency gradient permitted us to evaluate the influence of resources at middens beyond the influence of a resident squirrel. At each location, we quantified vegetation, landscape structure, abundance of conifer cones, and midden structure, and used capture-recapture, distance sampling, and remote cameras to quantify presence, abundance, and species richness of mammals and birds. Red squirrels and the resources they concentrated at middens influenced mammals and birds at the population scale and to a lesser extent at the community scale. At middens with higher residency rates of red squirrels, richness of medium and large mammals increased markedly and species richness of birds increased slightly. After accounting for local forest characteristics, however, only species richness of medium-to-large mammals was associated with a red squirrel being resident during surveys. In areas where red squirrels were resident during surveys or in areas with greater amounts of resources concentrated by red squirrels, abundances of two of four small mammal species and two of four bird species increased. We conclude that the presence of this ecosystem modifier and the resources it concentrates influence abundance of some mammals and birds, which may have implications for maintaining biodiversity across the wide geographic range inhabited by red squirrels and other larderhoarding animals.

• Ramos-Lara N, Koprowski J (2015) Spacing behavior of a non-larder-hoarding Tamiasciurus: a study of Mearns’s squirrels in xeric coniferous forests. Ethology 121: 196-205.

Abstract: In ecosystems with seasonal fluctuations in food supply many species use two strategies to store food: larder hoarding and scatter hoarding. However, because species at different geographic locations may experience distinct environmental conditions, differences in hoarding behavior may occur. Tree squirrels in the genus Tamiasciurus display variation in hoarding behavior. Whereas red (Tamiasciurus hudsonicus) and Douglas’s (Tamiasciurus douglasii) squirrels in mesic coniferous forests defend territories centered around larder hoards maintaining non-overlapping home ranges, red squirrels in deciduous forests defend small scatter-hoarded caches of cones maintaining overlapping home ranges. As in other rodent species, variation in hoarding behavior appears to influence the spacing behavior of red and Douglas’s squirrels. In contrast, Mearns’s squirrels (Tamiasciurus mearnsi) in xeric coniferous forests neither rely on larder hoards nor appear to display territorial behavior. Unfortunately, little is known about the ecology of this southernmost Tamiasciurus. Using radiotelemetry, we estimated home-range size, overlap, and maximum distance traveled from nest to examine the spacing behavior of Mearns’s squirrels. Similar to scatter-hoarding rodents, maximum distance traveled from nest was greater for males during mating season, whereas those of females were similar year round. Although no seasonal differences were detected, male home ranges were three times larger during mating season, whereas those of females were smaller and displayed a minor variation between seasons. Home ranges were overlapped year round but contrary to our expectations, overlap was greater during mating season for both sexes, with no detectable relationship between male home-range size and the number of females overlapped during mating season. Overall, the results appear to support our hypothesis that in the absence of larder hoards, the spacing behavior of Mearns’s squirrels should be different from larderhoarding congeners and more similar to scatter-hoarding rodents.

• Romeo, C., Ferrari, N., Lanfranchi, P., Saino, N., Santicchia, F., Martinoli, A., et al. (2015) Biodiversity threats from outside to inside: effects of alien grey squirrel (Sciurus carolinensis) on helminth community of native red squirrel (Sciurus vulgaris). Parasitology Research 114: 2621-2628. doi:10.1007/s00436-015-4466-3.

Abstract: Biological invasions are among the major causes of biodiversity loss worldwide, and parasites carried or acquired by invaders may represent an added threat to native species. We compared gastrointestinal helminth communities of native Eurasian red squirrels (Sciurus vulgaris) in the presence and absence of introduced Eastern grey squirrels (Sciurus carolinensis) to detect alterations induced by the alien species. In particular, we investigated whether spillover of a North American nematode Strongyloides robustus occurs and whether prevalence of a local parasite Trypanoxyuris sciuri in red squirrels is affected by grey squirrel presence. The probability of being infected by both parasites was significantly higher in areas co-inhabited by the alien species, where 61 % of examined red squirrels (n = 49) were infected by S. robustus and 90 % by T. sciuri. Conversely, in red-only areas, the two parasites infected only 5 and 70 % of individuals (n = 60). Overall, our findings support the hypothesis that red squirrels acquire S. robustus via spillover from the alien congener and suggest that invaders’ presence may also indirectly affect infection by local parasites through mechanisms diverse than spill-back and linked to the increased competitive pressure to which red squirrels are subjected. These results indicate that the impact of grey squirrel on red squirrels may have been underestimated and highlight the importance of investigating variation in macroparasite communities of native species threatened by alien competitors.

• Sanamxay, D., Douangboubpha, B., Bumrungsri, S., Satasook, C., & Bates Paul, J. J. (2015) A summary of the taxonomy and distribution of the red giant flying squirrel, Petaurista petaurista (Sciuridae, Sciurinae, Pteromyini), in mainland Southeast Asia with the first record from Lao PDR. Mammalia 79: 305-314.

Abstract: The occurrence of the red giant flying squirrel, Petaurista petaurista, in Lao PDR is confirmed on the basis of a single adult male specimen obtained from an informal food market in Thatlouang village, Xekong Province in the south of the country. This individual was reported to have been collected from close-by Thatlouang in the mixed deciduous forest or dry dipterocarp forest on the Bolaven Plateau. The record extends the known distribution of this species by approximately 700 km eastwards. Information is provided on the external, cranial, dental, and bacular characters of the new Lao specimen. It is compared with the holotypes and/or type descriptions of seven taxa, namely, barroni, candidula, cicur, melanotus, penangensis, taylori, and terutaus, described from mainland Southeast Asia, all of which are currently included in the synonymy of P. petaurista. On the basis of its external pelage colour, the Lao specimen is referred to P. p. barroni, which was described from southeast Thailand and is considered here to be a valid subspecies.

• Santicchia. F.,Romeo C., Grilli, G., Vezzoso, S., Wauters,  L.A., Mazzamuto, M., et al. (2015) The use of uterine scars to explore fecundity levels in invasive alien tree squirrels. Hystrix 26:

Abstract: In invasion ecology, reliable measures of female fecundity are necessary to infer population growth rate and develop control programs to determine the proportion of animals that should be culled to reduce population size. Here, we present a reliable staining technique of uterine scars to determine individual fecundity in terms of both seasonal and total (annual) number of young born per female. We applied this method to two alien squirrels species (grey squirrel, Sciurus carolinensis and Pallas’s squirrel, Callosciurus erythraeus) introduced in Northern Italy, obtaining carcasses from control campaigns from 2011 to 2013. We also investigated environmental and phenotypic variables that might affect individual variation in fecundity and compared annual reproductive output between the two species. For grey squirrels (n=44), 25% of examined females produced a single litter and 61% two litters. Females which reproduced in both seasons tended to have larger summer than spring litters (on average 2.61 and 1.94 offspring, respectively) and mean annual fecundity was 3.4 scars/female ranging from 1 to 8 births. There was no effect of year, eye lens weight, body size or body mass on total fecundity. For Pallas’s squirrel (n=31), 58% of females had a spring litter, some of these also produced a summer litter (35%) and a few even a third litter in autumn (10%). Heavier and older females (higher eye lens weight) had more uterine scars than younger animals with lower body mass. Finally, fecundity of the two IAS in Italy was similar or even higher than in the native range and/or in other countries of introduction, suggesting they are well adapted to their new environment and potentially have a high capacity to spread and recover after reduction of population size.

• Santicchia, F., Romeo, C., Martinoli, A., Lanfranchi, P., Wauters, L. A., & Ferrari, N. (2015) Effects of habitat quality on parasite abundance: do forest fragmentation and food availability affect helminth infection in the Eurasian red squirrel? Journal of Zoology 296:38-44.

Abstract: Habitat quality affects demography, population genetics, space use and phenotypic characteristics of mammals. However, little is known about the effects of habitat quality, fragmentation and/or food abundance, on host–parasite interactions. Here we present a first study on the relationships between the abundance of the dominant gastrointestinal helminth, Trypanoxyuris (Rodentoxyuris) sciuri, infecting the Eurasian red squirrel Sciurus vulgaris and three environmental factors: habitat type (mountain conifer forests vs. lowland mixed deciduous forests), fragmentation (fragmented woodlands vs. continuous forests) and food availability. Abundance of T. (R.) sciuri increased in fragmented woods. Furthermore, in mountain conifer forests, squirrels were more heavily infected after a poor Norway spruce seed crop than in years with medium or high seed production, indicating that squirrels are less capable of reducing parasite load when food availability is low. Hence, we suggest that T. (R.) sciuri abundance in red squirrels may be determined mainly by changes in host susceptibility induced by higher stress levels and/or poorer nutritional status, while in fragments, reduced genetic diversity may also increase host susceptibility to parasite infection. Although our data do not shed light on the mechanisms generating the observed patterns, results from other field studies highlighted the effect of stress and nutritional status on parasite infection, thus suggesting their implication in the changes in the abundance of T. (R.) sciuri.

• Selonen, V., Varjonen, R. & Korpimäki, E. (2015) Immediate or lagged responses of red squirrel population to pulsed resources. Oecologia 177: 401-411.

According to producer-consumer models, consumers should follow pulsed resources with a time lag. This view has been challenged by studies demonstrating that individuals may anticipate future resource pulses by increasing reproduction just before the pulse. We studied population fluctuations and reproduction in European red squirrels, Sciurus vulgaris, in relation to seed masting of the main food resource (the Norway spruce) in boreal coniferous forests between 1979 and 2013. Red squirrels are pre-dispersal seed predators, and previous studies have shown that they can anticipate the coming seed mast. We did not find any indication that anticipation of masting (year t ) increased red squirrel reproduction in the preceding spring to early summer. Instead, the reproductive output of the squirrels was highest in the spring following the mast, indicating that the population had to be at its largest size in the autumn after the mast (year(t+1)), when lots of subadults were around. However, we surmised, based on snow tracks and squirrel nest data, that the population crashed during the following winter (year(t+1)). These data reflected the adult population during winter, which peaked at the same time as the resource pulse. We can therefore conclude that the time lag between the resource pulse and the attainment of the peak number of squirrels was less than one year, and that the resource crash affected more juveniles and subadults than adults. The population increase overlapped with the occurrence of masting, but there was also a lagged response, supporting the classical view of producer-consumer models.

• Shuttleworth, C., Halliwell, E., & Lurz, P. W. W. (2015) Squirrel projects. European Squirrel Initiative, Woodbridge, Suffolk UK.

• Shuttleworth, C., Lurz, P., & Hayward, M. (2015) Red Squirrels: Ecology, Conservation & Management in Europe, 328 pp. European Squirrel Initiative, Woodbridge, Suffolk UK.
  • Bertolino, S., Martinoli, A., Paoloni, D., Marsan, A., & Wauters, L. (2015) The grey squirrel in Italy: impacts and management. In C. Shuttleworth, P. Lurz & M. Hayward (eds) Red Squirrels: Ecology, Conservation & Management in Europe, 163-173. European Squirrel Initiative, Woodbridge, Suffolk UK.
  • Everest, D., Shuttleworth, C., Grierson, S., Dastjerdi, A., Stidworthy, M., Duff, J., et al. (2015) The impact of adenoviruses on red squirrel (Sciurus vulgaris) conservation – with notes on other enteric viruses of potential significance. In C. Shuttleworth, P. Lurz & M. Hayward (eds) Red Squirrels: Ecology, Conservation & Management in Europe, 129-146. European Squirrel Initiative, Woodbridge, Suffolk UK.
  • Gurnell, J., Blackett, T., Butler, H., Lurz, P., Magris, L., & Shuttleworth, C. (2015) British red squirrel strongholds: challenges for conservation. In C. Shuttleworth, P. Lurz & M. Hayward (eds) Red Squirrels: Ecology, Conservation & Management in Europe, 213-232. European Squirrel Initiative, Woodbridge, Suffolk UK.
  • Gurnell, J., Lurz, P., & Wauters, L. (2015) Years of interactions and conflict in Europe: competition between Eurasian red squirrels and North American grey squirrels. In C. Shuttleworth, P. Lurz & M. Hayward (eds) Red Squirrels: Ecology, Conservation & Management in Europe. European Squirrel Initiative, Woodbridge, Suffolk UK.
  • Halliwell, E., Shuttleworth, C., Wilberforce, E., Denman, H., Lloyd, C., & Cartmel, S. (2015) Striving for success: an evaluation of local action to conserve red squirrels (Sciurus vulgaris) in Wales. In C. Shuttleworth, P. Lurz & M. Hayward (eds) Red Squirrels: Ecology, Conservation & Management in Europe, 176-192. European Squirrel Initiative, Woodbridge, Suffolk UK.
  • Jayne, K., & Leaver, L. (2015) Strategic decisions made by small mammals during scatter hoarding, cache recovery and cache pilferage. In C. Shuttleworth, P. Lurz & M. Hayward (eds) Red Squirrels: Ecology, Conservation & Management in Europe, 51-65. European Squirrel Initiative, Woodbridge, Suffolk UK.
  • Krauze-Gryz, D., & Gryz, J. (2015) A review of the diet of the red squirrel (Sciurus vulgaris) in different types of habitats. In C. Shuttleworth, P. Lurz & M. Hayward (eds) Red Squirrels: Ecology, Conservation & Management in Europe, 39-50. European Squirrel Initiative, Woodbridge, Suffolk UK.
  • Lawton, C., Waters, C., & Shuttleworth, C. (2015) Reintroductions and translocations of red squirrels within Europe. In C. Shuttleworth, P. Lurz & M. Hayward (eds) Red Squirrels: Ecology, Conservation & Management in Europe, 193-212. European Squirrel Initiative, Woodbridge, Suffolk UK.
  • Lurz, P., Bertolino, S., Koprowski, J., Willis, P., Tonkin, M., & Gurnell, J. (2015) Squirrel monitoring: snapshots of population presence and trends to inform management. In C. Shuttleworth, P. Lurz & M. Hayward (eds) Red Squirrels: Ecology, Conservation & Management in Europe, 279-297. European Squirrel Initiative, Woodbridge, Suffolk UK.
  • McInnes, C., Deane, D., & Fiegna, C. (2015) Squirrelpox virus: origins and the potential for its control. In C. Shuttleworth, P. Lurz & M. Hayward (eds) Red Squirrels: Ecology, Conservation & Management in Europe, 251-264. European Squirrel Initiative, Woodbridge, Suffolk UK.
  • Meredith, A., & Romeo, C. (2015) Disease and causes of mortality in red squirrel populations. In C. Shuttleworth, P. Lurz & M. Hayward (eds) Red Squirrels: Ecology, Conservation & Management in Europe, 116-127. European Squirrel Initiative, Woodbridge, Suffolk UK.
  • Sage, R., & Sotherton, N. (2015) Predation of woodland bird nests by tree squirrels in Britain Central Europe and North America. In C. Shuttleworth, P. Lurz & M. Hayward (eds) Red Squirrels: Ecology, Conservation & Management in Europe, 147-162. European Squirrel Initiative, Woodbridge, Suffolk UK.
  • Selonen, V., & Hanski, I. (2015) Occurrence and dispersal of the red squirrel and the Siberian flying squirrel in fragmented landscapes. In C. Shuttleworth, P. Lurz & M. Hayward (eds) Red Squirrels: Ecology, Conservation & Management in Europe, 67-82. European Squirrel Initiative, Woodbridge, Suffolk UK.
  • Seward, A., & O’Hare, S. (2015) The importance of community support and participation for regional red squirrel conservation: a case study from northern England. In C. Shuttleworth, P. Lurz & M. Hayward (eds) Red Squirrels: Ecology, Conservation & Management in Europe, 299-316. European Squirrel Initiative, Woodbridge, Suffolk UK.
  • Sheehy, E., & Lawton, C. (2015) Predators of red and grey squirrels in their natural and introduced ranges. In C. Shuttleworth, P. Lurz & M. Hayward (eds) Red Squirrels: Ecology, Conservation & Management in Europe, 83-96. European Squirrel Initiative, Woodbridge, Suffolk UK.
  • Shuttleworth, C., Lurz, P., Hayward, M., & Bertolino, S. (2015) Developing Integrated Red Squirrel Conservation in Europe. In C. Shuttleworth, P. Lurz & M. Hayward (eds) Red Squirrels: Ecology, Conservation & Management in Europe, 317-328. European Squirrel Initiative, Woodbridge, Suffolk UK.
  • Shuttleworth, C., Schuchert, P., Everest, D., McInnes, C., Rushton, S., Jackson, N., et al. (2015) Developing integrated and applied red squirrel conservation programmes: What lessons can Europe learn from a regional grey squirrel eradication programme in North Wales? In C. Shuttleworth, P. Lurz & M. Hayward (eds) Red Squirrels: Ecology, Conservation & Management in Europe, 233-250. Woodbridge, Suffolk UK, European Squirrel Initiative.
  • Signorile, L., Wang, J., Reuman, D., Bertolino, S., & Lurz, P. (2015) How population genetics can contribute to the management of grey squirrel invasions. In C. Shuttleworth, P. Lurz & M. Hayward (eds) Red Squirrels: Ecology, Conservation & Management in Europe, 97-113. European Squirrel Initiative, Woodbridge, Suffolk UK.
  • White, A., Lurz, P., Jones, H., Boots, M., Bryce, J., Tonkin, M., et al. (2015) The use of mathematical models in red squirrel conservation: Assessing the threat from grey invasion and disease to the Fleet basin stronghold. In C. Shuttleworth, P. Lurz & M. Hayward (eds) Red Squirrels: Ecology, Conservation & Management in Europe, 265-277. European Squirrel Initiative, Woodbridge, Suffolk UK.

• Simpson, V., Hargreaves, J., Butler, H., Blackett, T., Stevenson, K., & McLuckie, J. (2015) Leprosy in red squirrels on the Isle of Wight and Brownsea Island. Veterinary Record 177: 206-207. doi:10.1136/vr.h4491.

• Stevenson-Holt, C. D. & Sinclair, W. (2015) Assessing the geographic origin of the invasive grey squirrel using DNA sequencing: Implications for management strategies. Global Ecology and Conservation 3: 20-27.

Abstract: The invasive grey squirrel Sciurus carolinensis has become a major pest species causing negative effects to forestry and biodiversity. This study aims to assess the origin of grey squirrel within Cumbria using phylogeographic analysis to aid in management and control. The work reported analysed mitochondrial DNA sequences in the D-Loop gene of 73 grey squirrel individuals from multiple locations in the UK. The results indicate that individuals in north Cumbria are derived from individuals from Scotland and North East England. Other individuals in north Cumbria share a unique haplotype with south Cumbria and Lancashire suggesting a southerly origin and movement around or over the Cumbrian Mountain range which is thought of as a barrier to movements. The assessment of invasive species geographical origin and the identification of potential wildlife transit corridors through natural barriers are becoming more important as species shift range in response to environmental and ecological changes. With the grey squirrel population expansion also occurring in Italy, the European red squirrel may become threatened across its entire range. It is crucial to understand the population origins of the invasive grey squirrel and landscape usage to successfully manage the incursion routes and control the population.

• Studd, E. K., Boutin, S, McAdam, AG, Krebs, C.J. & Humphries, M.M. (2015). Predators, energetics and fitness drive neonatal reproductive failure in red squirrels. Journal of Animal Ecology 84: 249-259.

Abstract:

* Neonatal reproductive failure should occur when energetic costs of parental investment outweigh fitness benefits. However, little is known about the drivers of neonatal reproductive failure in free-ranging species experiencing continuous natural variation in predator abundance and in the energetic and fitness costs and benefits associated with parental investment.

* Long-term comprehensive studies are required to better understand how biotic, abiotic and life-history conditions interact to drive occurrences of reproductive failure in the wild.

* Using 24 years (1987–2011) of reproductive data from a northern boreal population of North American red squirrels in south-western Yukon, we examined the effects of predator abundance, energetics (resource availability, ambient temperature and litter size) and fitness benefits (probability of overwinter juvenile survival and maternal age) on occurrences of neonatal reproductive failure (494/2670 reproductive attempts; 18·5%).

* Neonatal reproductive failure was driven by a combination of predator abundance, and the energetic and fitness costs and benefits of parental investment. The abundance of mustelids and maternal age was positively related to the occurrence of neonatal reproductive failure. High energy costs associated with a combination of low resource availability and cold ambient temperatures or large litters, corresponded to increased occurrences of neonatal reproductive failure. However, the strength of these relationships was influenced by variation in juvenile overwinter survival (i.e. fitness benefits).

* We provide evidence that predation pressure is an important driver of neonatal reproductive failure. In addition, we found a trade-off occurs between resource-dependent energetic and fitness costs and benefits of raising the current litter to independence.

• Uchida, K., Suzuki, K., Shimamoto, T., Yanagawa, H., & Koizumi, I. (2015) Seasonal variation of flight initiation distance in Eurasian red squirrels in urban versus rural habitat. Journal of Zoology, n/a-n/a. doi:10.1111/jzo.12306.

Abstract: Urbanization has caused significant behavioural modifications in wild animals. Change in anti-predator behaviour is the most widespread example across different taxa in urban areas, which is probably due to a decrease in predation pressure and habituation towards humans. Seasonality or phenology has also been modified by urbanization since some resources in urban environments are highly controlled, for example, artificial feeding. Under natural conditions, anti-predator responses vary with seasonal variability in environmental and individual conditions. However, resource stability possibly reduces the seasonality of anti-predator behaviours in urban animals. Here, we compare the seasonal difference of flight initiation distance (FID), a measurement of anti-predator response, in Eurasian red squirrels Sciurus vulgaris between urban and rural areas in the Tokachi region, Hokkaido, Japan. Rural squirrels possessed FIDs two to three times longer than those of urban squirrels. We also found squirrels in rural areas lowered FID in autumn, but no seasonal difference was observed in urban squirrels. Our results suggest that continuous supplementary feeding may have buffered the seasonality in anti-predator response. In addition, strong habituation to humans may allow urban red squirrels to correctly assess human activity as benign rather than reacting unnecessarily.

• Vieira, B., Fonseca, C., & Rocha, R. (2015) Critical steps to ensure the successful reintroduction of the Eurasian red squirrel. Animal Biodiversity and Conservation 38: 49-58.

Abstract: Critical steps to ensure the successful reintroduction of the Eurasian red squirrel.— Wildlife reintroduction strategies aim to establish viable long–term populations, promote conservation awareness and provide economic benefits for local communities. In Portugal, the Eurasian red squirrel (Sciurus vulgaris) became extinct in the 16th century and was reintroduced in urban parks in the 1990s, mainly for aesthetic and leisure purposes. We evaluated the success of this reintroduction in two urban parks and here described the critical steps. We assessed habitat use, population density and abundance, and management steps carried out during reintroduction projects. Reintroductions have been successful to some extent given squirrels are present 20 years after release. However, populations in both parks are declining due to the lack of active management and poor quality habitat. Successful reintroduction of Eurasian red squirrel in areas without competition of alien tree squirrels involves three critical main stages. The pre–project stage includes studies on habitat quality, genetic proximity between donors and closest wild population, and health of donor stocks. In the release stage, the number of individuals released will depend on resource variability, and the hard release technique is an effective and economically viable method. Post–release activities should evaluate adaptation, mitigate mortality, monitor the need for supplementary feeding, provide veterinary support, and promote public awareness and education.

• Wilson, D. R., Goble, A. R., Boutin, S., Humphries, M. M., Coltman, D. W., Gorrell, J. C., et al. (2015) Red squirrels use territorial vocalizations for kin discrimination. Animal Behaviour 107: 79-85. doi:dx.doi.org/10.1016/j.anbehav.2015.06.011.

Abstract: The ability to discriminate among individuals, or among classes of individuals, can provide animals with important fitness benefits. Although several mechanisms for discrimination are possible, most require animals to show stable phenotypic variation that reflects their identity or their membership in a particular class (e.g. sex, mate, kin). For territorial animals that rarely interact physically, vocalizations could serve as long-distance signals that facilitate discrimination. In this study, we tested whether the territorial rattle vocalizations of North American red squirrels, Tamiasciurus hudsonicus, are repeatable, and whether they could hence provide the basis for multiple types of discrimination. We measured four structural features from two rattles from each of 76 marked squirrels. All four features were repeatable, which is consistent with territorial rattles being individually distinctive. We then conducted a playback experiment to determine whether squirrels use rattles for discrimination. Specifically, we tested whether squirrels discriminate between the rattles of neighbours and non-neighbours, and kin (coefficient of relatedness, r ≥ 0.25) and non-kin (r &lt; 0.125). Following a 2 × 2 factorial design, we broadcast a rattle from a non-neighbouring nonkin individual to 15 subjects, from a neighbouring nonkin individual to 14 subjects, from a non-neighbouring kin individual to 11 subjects, and from a neighbouring kin individual to 13 subjects. Subjects did not discriminate between the rattles of neighbours and non-neighbours, but did respond differently to the rattles of kin and nonkin. Specifically, squirrels were significantly more likely to produce a rattle of their own in response to the broadcasted rattles of nonkin versus the broadcasted rattles of kin. This result demonstrates that red squirrels can use territorial vocalizations for kin discrimination. It also suggests that they are more tolerant of territorial intrusions by kin.

• Yi, X. & Wang, Z. (2015) Dissecting the roles of seed size and mass in seed dispersal by rodents with different body sizes. Animal Behaviour 107: 263-267.

Abstract: The process of seed dispersal is influenced by animals’ responses to seeds with different traits. Seed size/mass is one of the key seed traits affecting animal-mediated seed dispersal. Although seed size is usually positively correlated with seed mass, it is not clear whether the two seed traits show similar effects on seed choice and seed dispersal by animals. In the present study, we artificially manipulated nuts of Juglans mandshurica to create two types of seeds: (1) seeds of the same size but different mass and (2) seeds of the same mass but different size. Our aim was to explore whether food-hoarding animals respond differently to seed size and mass. Our results showed that seed size was more important than seed mass in determining seed removal by wood mice, Apodemus peninsulae. However, the larger red squirrels, Sciurus vulgaris, evaluated seed profitability based on mass rather than size. Moreover, seed mass showed consistent positive effects on seed dispersal distances of J. mandshurica by both A. peninsulae and S. vulgaris. Our results indicate that the effects of seed size and mass on seed dispersal processes are not always consistently correlated. In addition, seed dispersers of different size show different responses to given seed traits such as size or mass. More effort should be made to investigate the role of seed traits and seed dispersers in seed dispersal systems.

• Zelditch, M.L.,Li, J.,Tran L.A.P. & Swiderski, D.L. (2015) Relationships of diversity, disparity and their evolutionary rates in squirrels (Sciuridae). Evolution 69:1284-1300.

Abstract: Several theories predict that rapidly diversifying clades will also rapidly diverge phenotypically; yet, there are also reasons for suspecting that diversification and divergence might not be correlated. In the widely distributed squirrel clade (Sciuridae), we test for correlations between per-lineage speciation rates, species richness, disparity and a time-invariant measure of disparity that allows for comparing rates when evolutionary modes differ, as they do in squirrels. We find that species richness and speciation rates are not correlated with clade age or with each other. Disparity appears to be positively correlated with clade age because young, rapidly diversifying Nearctic grassland clades are strongly pulled to a single stable optimum but older, slowly diversifying Paleotropical forest clades contain lineages that diverge along multiple ecological and morphological lines. That contrast is likely due to both the environments they inhabit and their phylogenetic community structure. Our results argue against a shared explanation for diversity and disparity in favor of geographically mediated modes of speciation and ecologically mediated modes of phenotypic evolution.

2014

• Ball, S.J., Daszak, P., Sainsbury, A.W. & Snow, K.R. (2014) Coccidian parasites of red squirrels (Sciurus vulgaris) and grey squirrels (Sciurus carolinensis) in England. Journal of Natural History 1-6.
• Chantrey, J., Dale, T. D., Read, J. M., White, S., Whitfield, F., Jones, D., et al. (2014) European red squirrel population dynamics driven by squirrelpox at a gray squirrel invasion interface. Ecology and Evolution 4: 3788-3799.
• Cudworth, N. & Koprowski, J. (2014) Survival and mortality of the Arizona gray squirrel (Sciurus arizonensis). The Southwestern Naturalist 59: 423-426.
  Bateman, P.W. & Fleming, P.A. (2014) Does human pedestrian behaviour influence risk assessment in a successful mammal urban adapter? Journal of Zoology 294:93-98.
• Bertolino, S., Montezemolo, N., Preatoni, D., Wauters, L. & Martinoli, A.  (2014) A grey future for Europe: Sciurus carolinensis is replacing native red squirrels in Italy. Biological Invasions 16: 53-62.
• Bonnington, C., Gaston, K. & Evans, K. (2014) Squirrels in suburbia: influence of urbanisation on the occurrence and distribution of a common exotic mammal. Urban Ecosystems 17: 533-546.
• Bonnington, C., Gaston, K., &  Evans,K.L. (2014) Relative roles of grey squirrels, supplementary feeding, and habitat in shaping urban bird assemblages. PLoS ONE 9: e109397.
• Boutin, S. & Lane, J.E. (2014) Climate change and mammals: evolutionary versus plastic responses. Evolutionary Applications 7: 29-41.
• Chavez, A. S., & Kenagy, G. J. (2014) Clinal colour variation within a panmictic population of tree squirrels, I (Rodentia: Sciuridae), across an ecological gradient. Biological Journal of the Linnean Society n/a-n/a.
• Collins, L.M., Warnock, N.D., Tosh, D.G., McInnes, C., Everest, D., Montgomery, W.I., Scantlebury, M., Marks, N., Dick, J.T.A. & Reid, N. (2014)  Squirrelpox virus: assessing prevalence, transmission and environmental degradation. PLoS ONE 9: e89521.
• Darby, A.C., McInnes, C.J., Kjaer, K.H., Wood, A.R., Hughes, M., Martensen, P.M., Radford, A.D., Hall, N. & Chantrey, J. (2014) Novel host-related virulence factors are encoded by squirrelpox virus, the main causative agent of epidemic disease in red squirrels in the UK. . PLoS One 9: e96439.
• Everest, D. J., Shuttleworth, C. M., Stidworthy, M. F., Grierson, S. S., Duff, J. P., & Kenward, R. E. (2014) Adenovirus: an emerging factor in red squirrel Sciurus vulgaris conservation. Mammal Review, n/a-n/a.
• Flaherty, S., Lurz, P.W.W. & Patenaude, G. (2014) Use of LiDAR in the conservation management of the endangered red squirrel (Sciurus vulgaris L.). Journal of Applied Remote Sensing 8: 083592-083592.
• Fletcher, Q.E., Speakman, J.R., Boutin, S., Lane, J.E., McAdam, A.G., Gorrell, J.C., Coltman, D.W. & Humphries, M.M. (2014) Daily energy expenditure during lactation is strongly selected in a free-living mammal. Functional Ecology 29: 1985-208.
• Gurnell, J., Lurz, P. W. W., & BertoldiI, W. (2014) The changing patterns in the distribution of red and grey squirrels in the North of England and Scotland between 1991 and 2010 based on volunteer surveys. Hystrix, the Italian Journal of Mammalogy 25:
• Hatten, J.R. (2014) Mapping and monitoring Mount Graham red squirrel habitat with Lidar and Landsat imagery. Ecological Modelling 289: 106-123.
• Liu, Z., Li, B., Ma, J., Zheng, D., & Xu, Y. (2014) Phylogeography and genetic diversity of the red squirrel (Sciurus vulgaris) in China: Implications for the species’ postglacial expansion history. Mammalian Biology – Zeitschrift für Säugetierkunde 79: 247-253.
• Luo, Y., Yang, Z., Steele, M. A., Zhang, Z., Stratford, J. A., & Zhang, H. (2014) Hoarding without reward: rodent responses to repeated episodes of complete cache loss. Behavioural Processes 106: 36-43.
• Lurz, P. (2014) Changing ‘Red to Grey’: Alien Species Introductions to Britain and the Displacement and Loss of Native Wildlife from our Landscapes. In Displaced Heritage. pp. 265-272. Boydell & Brewer.
• Martínez-Duque, P., Avila-Flores, R., Emerson, G. L., Carroll, D. S., Suzán, G., & Gallardo-Romero, N. F. (2014) Orthopoxvirus Antibodies in Grey Squirrels (Sciurus aureogaster) in Mexico City, Mexico. Journal of Wildlife Diseases 50, 696-698.
• McGowan, N.E., Marks, N.J., McInnes, C.J., Deane, D., Maule, A.G. & Scantlebury, M. (2014) Effects of parasitism and morphology on squirrelpox virus seroprevalence in grey squirrels (Sciurus carolinensis). PLoS ONE 9: e83106.
• Murrant, M. N., Bowman, J., & Wilson, P. J. (2014) A test of non-kin social foraging in the southern flying squirrel (Glaucomys volans). Biological Journal of the Linnean Society n/a-n/a.
• Palmer Rosa, R. & Koprowski John, L. (2014) Feeding behavior and activity patterns of Amazon red squirrels. Mammalia 78: 303.
• Palmer, G.H., Koprowski, J.L. & Pernas, AJ. (2014) Distribution and spread of an introduced insular population of red-bellied squirrels (Sciurus aureogaster) in Florida.  Mammalia 78: 67-73.
• Patterson, J. E. H., Neuhaus, P., Kutz, S. J., & Ruckstuhl, K. E. (2013) Parasite removal improves reproductive success of female North American red squirrels (Tamiasciurus hudsonicus). PLoS ONE 8: e55779.
• Pisanu, B., Chapuis, J.-L., Dozières, A., Basset, F., Poux, V. & Vourc’h, G. (2014) High prevalence of Borrelia burgdorferis. L in the European red squirrel Sciurus vulgaris in France. Ticks and Tick-borne Diseases 5: 1-6.
• Ramos-Lara, N., & Koprowski, J. L. (2014) Spacing behavior of a non-larder-hoarding Tamiasciurus: a study of Mearns’s squirrels in xeric coniferous forests. Ethology 121: 196-205.
• Ramos-Lara, N., & Koprowski, J. (2014) Deforestation and knowledge gaps threaten conservation of less charismatic species: status of the arboreal squirrels of Mexico. Mammalia 78: 417-427.
• Rézouki, C., Dozières, A., Le Cœur, C., Thibault, S., Pisanu, B., Chapuis, J.-L., et al. (2014) A viable population of the European red squirrel in an urban park. PLoS ONE 9: e105111.
• Rocha, R., Wauters, L., Mathias, M. L., & Fonseca, C. (2014) Will an ancient refuge become a modern one? A critical review on the conservation and research priorities for the red squirrel (Sciurus vulgaris) in the Iberian Peninsula.  Hystrix, Italian Journal of Mammalogy n/a-n/a..
• Romeo, C., Ferrari, N., Rossi, C., Everest, D.J., Grierson, S.S., Lanfranchi, P., Martinoli, A., Saino, N., Wauters, L.A. & Hauffe, H.C. (2014) Ljungan Virus and an Adenovirus in Italian Squirrel Populations. Journal of Wildlife Diseases 50: 409-411.
• Romeo, C., Wauters, L., Ferrari, N., Lanfranchi, P., Martinoli, A., Pisanu, B., Preatoni, D. & Saino, N. (2014)  Macroparasite Fauna of Alien Grey Squirrels (Sciurus carolinensis): Composition, Variability and Implications for Native Species. PLoS ONE 9, e88002.
• Romeo, C., Ferrari, N., Rossi, C., Everest, D., Grierson, S., Lanfranchi, P., Martinoli, A., Saino, N., Wauters, L. &  Hauffe, H. (2014) Ljungan Virus and an Adenovirus in Italian Squirrel Populations. Journal of Wildlife Diseases 50: 409-411.
• Schuchert, P., Shuttleworth, C., McInnes, C., Everest, D., & Rushton, S. (2014) 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? Biological Invasions n/a-n/a.
• Selonen, V., Hanski, I., & Wistbacka, R. (2014) Communal nesting is explained by subsequent mating rather than kinship or thermoregulation in the Siberian flying squirrel. Behavioral Ecology and Sociobiology 68: 971-980.
• Shearer, L., Bray, R., & Toner, C. (2014) An experimental study investigating the ability of volunteers to identify squirrel species from tail–hair samples. Animal Biodiversity and Conservation 37: 145-147.
• Shuttleworth, C., Everest, D., McInnes, C., Greenwood, A., Jackson, N., & Rushton, S., et al. (2014) Inter-specific viral infections: Can the management of captive red squirrel collections help inform scientific research? Hystrix, Italian Journal of Mammalogy 25: 18-24.
• Sheehy, E. & Lawton, C. (2014) Population crash in an invasive species following the recovery of a native predator: the case of the American grey squirrel and the European pine marten in Ireland. Biodiversity and Conservation n/a-n/a.
• Signorile, A., Paoloni, D. & Reuman, D. (2014)  Grey squirrels in central Italy: a new threat for endemic red squirrel subspecies. Biological Invasions n/a-n/a
• Signorile, A.L., Wang, J., Lurz, P.W.W., Bertolino, S., Carbone, C. & Reuman, D.C.  (2014) Do founder size, genetic diversity and structure influence rates of expansion of North American grey squirrels in Europe? Diversity and Distributions n/a-n/a.
• Stevenson-Holt, C. D., Watts, K., Bellamy, C. C., Nevin, O. T., & Ramsey, A. D. (2014) Defining Landscape Resistance Values in Least-Cost Connectivity Models for the Invasive Grey Squirrel: A Comparison of Approaches Using Expert-Opinion and Habitat Suitability Modelling. PLoS ONE 9: e112119.
• Studd, E. K., Boutin, S., McAdam, A. G., Krebs, C. J., & Humphries, M. M. (2014) Predators, energetics and fitness drive neonatal reproductive failure in red squirrels. Journal of Animal Ecology n/a-n/a.
• Vasilieva, N. A., & Tchabovsky, A. V. (2014) Timing is the only thing: reproduction in female yellow ground squirrels (Spermophilus fulvus). Canadian Journal of Zoology 92: 737-747.
• White, A., Bell, S.S., Lurz, P.W.W. & Boots, M. (2014) Conservation management within strongholds in the face of disease-mediated invasions: red and grey squirrels as a case study. Journal of Applied Ecology n/a-n/a.
• Williams, C. T., Barnes, B. M., Kenagy, G. J., & Buck, C. L. (2014) Phenology of hibernation and reproduction in ground squirrels: integration of environmental cues with endogenous programming. Journal of Zoology 292: 112-124.
• Zong, C., Mei, S., Santicchia, F., Wauters, L., Preatoni, D., & Martinoli, A. (2014) Habitat effects on hoarding plasticity in the Eurasian red squirrel (Sciurus vulgaris). Hystrix, Italian Journal of Mammalogy 25: 14-17.

2013

• Benitez, V.V., Almada Chavez, S., Gozzi, A.C., Messetta, M.L. &  Guichón, M.L (2013) Invasion status of Asiatic red-bellied squirrels in Argentina. Mammalian Biology – Zeitschrift für Säugetierkunde 78: 164-170.
• Bertolino, S., & Lurz, P. W. W. (2013) Callosciurus squirrels: worldwide introductions, ecological impacts and recommendations to prevent the establishment of new invasive populations. Mammal Review 43: 22-33.
• Bonnington, C., Gaston, K.J. &  Evans, K.L. (2013) Assessing the potential for Grey Squirrels Sciurus carolinensis to compete with birds at supplementary feeding stations. Ibis 156: 220-226.
• Bonnington, C., Gaston, K.J., & Evans, K.L. (2013) Fearing the feline: domestic cats reduce avian fecundity through trait-mediated indirect effects that increase nest predation by other species. Journal of Applied Ecology 50: 15-24.
• Bonnington, C., Gaston, K. & Evans, K., (2013) Squirrels in suburbia: influence of urbanisation on the occurrence and distribution of a common exotic mammal. Urban Ecosystems 1-14.
• Bosch, S. &  Lurz, PW..W. (2013) The process of drey construction in red squirrels – nestbox observations based on a hidden camera. Hystrix 24: 199-202.
• Bosson, C., Palme, R., & Boonstra, R. (2013) Assessing the impact of live-capture, confinement, and translocation on stress and fate in eastern gray squirrels. Journal of Mammalogy 94: 1401-1411.
• Brunner, J. L., Duerr, S., Keesing, F., Killilea, M., Vuong, H., & Ostfeld, R. S. (2013) An experimental test of competition among mice, chipmunks, and squirrels in deciduous forest fragments. PLoS ONE 8: e66798.
• Chavez, A.S., Maher, S.P., Arbogast, B.S. & Kenagy, G.J. (2013) Diversification and gene flow in nascent lineages of island and mainland north american tree squirrels (Tamiasciurus). Evolution n/a-n/a.
• Della-Flora, F., Melo, G., Sponchiado, J., & Cáceres, N. (2013) Association of the southern Amazon red squirrel Urosciurus spadiceus Olfers, 1818 with mixed species bird flocks. Mammalia 77 113-117.
• Desantis, L.M., Delehanty, B., Weir, J.T., & Boonstra, R. (2013) Mediating free glucocorticoid levels in the blood of vertebrates: are corticosteroid-binding proteins always necessary? Functional Ecology 27: 107-119.
• Di Febbraro, M., Lurz, P., Genovesi, P., Maiorano, L., Girardello, M., & Bertolino, S. (2013) The use of climatic niches in screening procedures for introduced species to evaluate risk of spread: a case with the american eastern grey squirrel. PLoS ONE 87.
• Everest, D. J., Butler, H., Blackett, T., Simpson, V. R., & Shuttleworth, C. M. (2013) Adenovirus infection in red squirrels in areas free from grey squirrels. Veterinary Record 173: 199-200.
• Fletcher, Q.E., Selman, C., Boutin, S., McAdam, A.G., Woods, S.B., Seo, A.Y., Leeuwenburgh, C., Speakman, J.R., & Humphries, M.M. (2013)  Oxidative damage increases with reproductive energy expenditure and is reduced by food-supplementation. Evolution 67: 1527-1536.
• Getschow, C.M., Rivers, P., Sterman, S., Lumpkin, D.C. & Tarvin, K.A. (2013) Does gray squirrel (Sciurus carolinensis) response to heterospecific alarm calls depend on familiarity or acoustic similarity? Ethology: n/a-n/a.
• Gür, H. (2013) The effects of the Late Quaternary glacial–interglacial cycles on Anatolian ground squirrels: range expansion during the glacial periods? Biological Journal of the Linnean Society 109: 19-32.
• Lin, Z. (2013) Health surveillance and MHC Class II DRB genetic diversity in the red squirrel (Sciurus vulgaris) population of the Isle of Arran. Master of Veterinary Science, University of Edinburgh.
• Mantooth, S. J., Hafner, D. J., Bryson, R. W., & Riddle, B. R. (2013) Phylogeographic diversification of antelope squirrels (Ammospermophilus) across North American deserts. Biological Journal of the Linnean Society n/a-n/a.
• Mayle, B.A. & Broome, A.C. (2013) Changes in the impact and control of an invasive alien: the grey squirrel (Sciurus carolinensis) in Great Britain, as determined from regional surveys. Pest Management Science 69: 323-333.
• McInnes, C. J., Coulter, L., Dagleish, M. P., Deane, D., Gilray, J., Percival, A., et al. (2013) The emergence of squirrelpox in Ireland. Animal Conservation 16: 51-59.
• Monclús, R., von Holst, D., Blumstein, D. T., & Rödel, H. G. (2013) Long-term effects of litter sex ratio on female reproduction in two iteroparous mammals. Functional Ecology n/a-n/a.
• Palmer, G., Koprowski, J., & Pernas, A. (2013) Nest tree and site selection of an introduced population of red-bellied squirrels (Sciurus aureogaster). Journal of Mammalogy 94: 1274-1281.
• Ramos-Lara, N., Koprowski, J., & Swann, D. (2013) Nest-site characteristics of the montane endemic Mearns’s squirrel (Tamiasciurus mearnsi): an obligate cavity-nester? Journal of Mammalogy 94: 50-58.
• Říčanová, Š., Koshev, Y., Říčan, O., Ćosić, N., Ćirović, D., Sedláček, F., et al. (2013) Multilocus phylogeography of the European ground squirrel: cryptic interglacial refugia of continental climate in Europe. Molecular Ecology 22: 4256-4269.
• Romeo, C., Pisanu, B., Ferrari, N., Basset, F., Tillon, L., Wauters, L.A., Martinoli, A., Saino, N. & Chapuis, J.L. (2013) Macroparasite community of the Eurasian red squirrel (Sciurus vulgaris): poor species richness and diversity. Parasitol Res 112, 3527-3536.
• Rong, K., Yang, H., Ma, J., Zong, C., & Cai, T. (2013) Food availability and animal space use both determine cache density of Eurasian red squirrels. PLoS ONE 8: e80632.
• Selonen, V., Painter, J., Rantala, S. & Hanski, I. (2013) Mating system and reproductive success in the Siberian flying squirrel. Journal of Mammalogy 94: 1266-1273.
• Signorile, A. (2013) Genetic determinants of the expansion of eastern grey squirrel populations across Europe. PhD, Imperial College London.
• Simpson, S., Bleamoied, N., Peniche, G., Dozières, A., Blackett, T., Coleman, S., Cornish, N., & Groombridge, J. (2013) Genetic structure of introduced populations: 120-year-old DNA footprint of historic introduction in an insular small mammal population. Ecology and Evolution 3: 614-628.
• Simpson, V., Hargreaves, J., Butler, H., Davison, N. & Everest, D. (2013) Causes of mortality and pathological lesions observed post-mortem in red squirrels (Sciurus vulgaris) in Great Britain. BMC Veterinary Research 9: 229.
• Stevenson, C.D., Ferryman, M., Nevin, O.T., Ramsey, A.D., Bailey, S. & Watts, K. (2013) Using GPS telemetry to validate least-cost modeling of gray squirrel (Sciurus carolinensis) movement within a fragmented landscape. Ecology and Evolution n/a-n/a.
• Xiao, Z., Gao, X. & Zhang, Z. (2013) Sensitivity to seed germination schedule by scatter-hoarding Pére David’s rock squirrels during mast and non-mast years. Ethology 119: 472-479.
• Yi, X., Liu, G., Steele, M. A., Shen, Z., & Liu, C. (2013) Directed seed dispersal by a scatter-hoarding rodent: the effects of soil water content. Animal Behaviour 86: 851-857.
• Zapponi, L., Del Bianco, M., Luiselli, L., Catorci, A. & Bologna, M.A. (2013) Assessing environmental requirements effects on forest fragmentation sensitivity in two arboreal rodents. Mammalian Biology – Zeitschrift für Säugetierkunde 78: 157-163.

2012

• Bosch, S. &  Lurz, P.W.W.  (2012) The Eurasian Red SquirrelSciurus vulgaris. Die Neue Brehm-Bücherei, English Edition Vol. 183, Hohenlohe, Germany. ISBN 978-3-89432-258-8.
• Bosson, C.O., Islam, Z. & Boonstra, R. (2012) The impact of live trapping and trap model on the stress profiles of North American red squirrels. Journal of Zoology 288: 159-169.
• Di Cerbo, A. and C. Biancardi (2012) Monitoring small and arboreal mammals by camera traps: effectiveness and applications. Acta theriologica: 1-5.
• Digweed, S., Rendall, D., & Imbeau, T. (2012) Who’s your neighbor? Acoustic cues to individual identity in red squirrel Tamiasciurus hudsonicus rattle calls. Current Zoology 58: 758-764.
• Dozières, A., Chapuis, J.-L., Thibault, S., & Baudry, E. (2012). Genetic structure of the French red squirrel populations: implication for conservation. PLoS ONE 7:  e47607.
• Everest, D. J., Shuttleworth, C. M., Grierson, S. S., Duff, J. P., Jackson, N., Litherland, P., Kenward, R.E. & Stidworthy, M.F. (2012). Systematic assessment of the impact of adenovirus infection on a captive reintroduction project for red squirrels (Sciurus vulgaris). Veterinary Record 171:  176.
• Flaherty, S., Patenaude, G., Close, A., & Lurz, P.W.W. (2012) The impact of forest stand structure on red squirrel habitat use.Forestry 85: 437-444.
• Fletcher, Q.E., Speakman, J.R., Boutin, S., McAdam, A.G., Woods, S.B., & Humphries, M.M. (2012) Seasonal stage differences overwhelm environmental and individual factors as determinants of energy expenditure in free-ranging red squirrels. Functional Ecology 26: 677-687.
• Gurnell, J. & Lurz, P.W.W. (2012) Red Squirrel, In:  Cresswell, W., Birks, J., Dean, M.D., Pacheco, M., Trewhella, W., Wells, D., Wray, S. (Eds.) UK BAP Mammals; Interim Guidance for Survey Methodologies, Impact Assessment and Mitigation. The Mammal Society, Southampton, pp. 9-21.
• Gurnell, J., Lurz, P.W.W. & Wauters, L.A.  (2012) Squirrels. TheMammal Society,  Southampton, UK.  Pp. 35.
• Mayle B, Ferryman M, Peace A, Yoder C, Miller L, & Cowan D. (2012) The use of DiazaCon™ to limit fertility by reducing serum cholesterol in female grey squirrel, Sciurus carolinensis. Pest management Science (2012).
• McInnes, C.J., Coulter, L., Dagleish, M.P., Deane, D., Gilray, J., Percival, A., Willoughby, K., Scantlebury, M., Marks, N., Graham, D., Everest, D.J., McGoldrick, M., Rochford, J., McKay, F. & Sainsbury, A.W. (2012) The emergence of squirrelpox in Ireland.Animal Conservation, n/a-n/a.
• Pečnerová, P. & Martínková, N. (2012) Evolutionary history of tree squirrels (Rodentia, Sciurini) based on multilocus phylogeny reconstruction. Zoologica Scripta 41: 211-219.
• Previtali, M.A., Ostfeld, R.S., Keesing, F., Jolles, A.E., Hanselmann, R., & Martin, L.B., (2012) Relationship between pace of life and immune responses in wild rodents. Oikos 121: 1483-1492
• Selonen, V. & Hanski, I. (2012) Dispersing Siberian flying squirrels (Pteromys volans) locate preferred habitats in fragmented landscapes. Canadian Journal of Zoology 90: 885-892.
• Selonen, V., Hanski, I. & Mäkeläinen, S. (2012) Predictors of long-distance dispersal in the Siberian flying squirrel. Evolutionary Ecology 6: 1361-1369.
• Shuttleworth, C., Lurz, P., Geddes, N. & Browne, J. (2012) Integrating red squirrel (Sciurus vulgaris) habitat requirements with the management of pathogenic tree disease in commercial forests in the UK. Forest Ecology and Management 279: 167-175.
• Strauss, A., White, A., & Boots, M. (2012) Invading with biological weapons: the importance of disease-mediated invasions. Functional Ecology 26(6): 1249-1261.
• Taylor, R.W., Boon, A.K., Dantzer, B., Réale, D., Humphries, M.M., Boutin, S., Gorrell, J.C., Coltman, D.W. & McAdam, A.G. (2012) Low heritabilities, but genetic and maternal correlations between red squirrel behaviours. Journal of Evolutionary Biology 25: 514-624.
• Xiao, Z. & Zhang, Z. (2012) Behavioural responses to acorn germination by tree squirrels in an old forest where white oaks have long been extirpated. Animal Behaviour 83,:945-951.

2011

• Bertolino, S., Lurz, P.W.W. (2011) Callosciurus squirrels: worldwide introductions, ecological impacts and recommendations to prevent the establishment of new invasive populations. Mammal Review: no-no.
• Chapuis, J.-L., Obolenskaya, E., Pisanu, B. & Lissovsky, A. (2011) Datasheet on Tamias sibiricius. CABI, Wallingford, UK.www.cabi.org/isc
• Chavez, A. S., Saltzberg, C. J. & Kenagy, G. J. (2011) Genetic and phenotypic variation across a hybrid zone between ecologically divergent tree squirrels (Tamiasciurus). Molecular Ecology 20: 3350-3366.
• Dantzer, B., McAdam, A. G., Palme, R., Humphries, M. M., Boutin, S. & Boonstra, R. (2011) Maternal androgens and behaviour in free-ranging North American red squirrels. Animal Behaviour 81: 469-479.
• Dinets, V. (2011) Observations of the woolly flying squirrel Eupetaurus cinereus in Pakistan. Mammalia 75: 277-280.
• Garroway, C. J., Bowman, J., Holloway, G. L., Malcolm, J. R. & Wilson, P. J. (2011) The genetic signature of rapid range expansion by flying squirrels in response to contemporary climate warming. Global Change Biology 17: 1760-1769.
• Gooderham, K.  & Schulte-Hostedde, A. (2011) Macroparasitism influences reproductive success in red squirrels (Tamiasciurus hudsonicus). Behavioral Ecology 22: 1195-1200.
• Gurnell, J., McDonald, R. & Lurz, P. W. W. (2011) Making red squirrels more visible: the use of baited visual counts to monitor populations. Mammal Review 41:244-250.
• Huang, Z., Y. Wang, H. Zhang, F. Wu & Z. Zhang (2011) Behavioural responses of sympatric rodents to complete pilferage. Animal Behaviour: 81, 831.
• Lowney, A. (2011) Impact of mountain bike trails on red squirrel population (Sciurus vulgaris) in Whinlatter Forest, Cumbria.Bioscience Horizons 4, 99-107.
• Marmet, J., Pisanu, B. & Chapuis, J.-L. (2011) Natal dispersal of introduced Siberian chipmunks, Tamias sibiricus, in a suburban forest. Journal of Ethology, 29: 23-29.
• Marsot, M., Sigaud, M., Chapuis, J.-L., Ferquel, E., Cornet, M. & Vourc’h, G. (2011). Introduced Siberian chipmunks (Tamias sibiricus barberi) harbour more diverse Borrelia burgdorferi sensu lato genospecies than native bank voles (Myodes glareolus). Applied and Environmental Microbiology, 77: 5716-5721.
• McCleery, R. A. & Parker, I. D. (2011) Influence of the urban environment on fox squirrel range overlap. Journal of Zoology:no-no.
• Obon, E., Juan-Sallés, C., McInnes, C. & Everest, D. (2011) Poxvirus identified in a red squirrel (Sciurus vulgaris) from Spain. Veterinary Record January 22, 86.
• Oshida, T., Lin, L.-K., Chang, S.-W., Chen, Y.-J. & Lin, J.-K.  (2011) Phylogeography of two sympatric giant flying squirrel subspecies, Petaurista alborufus lena and P. philippensis grandis (Rodentia: Sciuridae), in Taiwan. Biological Journal of the Linnean Society 102: 404-419.
• Pelech, S. A., Smith, J. N. M. & Boutin, S. (2009) A predator’s perspective of nest predation: predation by red squirrels is learned, not incidental. Oikos, no-no.
• Schmidt, A. (2011) Functional differentiation of trailing and leading forelimbs during locomotion on the ground and on a horizontal branch in the European red squirrel (Sciurus vulgaris, Rodentia). Zoology 114: 155-164.
• Summers, R. W. (2011) Patterns of exploitation of annually varying Pinus sylvestris cone crops by seed-eaters of differing dispersal ability. Ecography: 723-728.
• Waters, C. and Lawton, C. (2011) Red Squirrel Translocation in Ireland. Irish Wildlife Manuals, No. 51. National Parks and Wildlife Service, Department of the Environment, Heritage and Local Government, Dublin, Ireland.
• Yoder, C., Mayle, B., Furcolow, C., Cowan, D. & Fagerstone, K. (2011) Feeding of grey squirrels (Sciurus carolinensis) with the contraceptive agent DiazaCon™: effect on cholesterol, haematology, and blood chemistry. Integrative Zoology 6: 409-419.

2010

• Atkin, J., Radford, A., Coune, K., Stavisky, J. & Chantrey, J. (2010) Detection of squirrel poxvirus by nested and real-time PCR from red (Sciurus vulgaris) and grey (Sciurus carolinensis) squirrels. BMC Veterinary Research 6: 33-42.
• Boyer, N., Réale, D., Marmet, J., Pisanu, B. & Chapuis, J.-L. (2010) Personality, space use, and tick load in an introduced population of Siberian chipmunks, Tamias sibiricus. Journal of Animal Ecology, 79 : 538-547.
• Chapuis, J.-L., Ferquel, E., Patey, O., Vourc’h, G. & Cornet, M. (2010) Borréliose de Lyme : situation générale et conséquences de l’introduction en Ile-de-France d’un nouvel hôte, le tamia de Sibérie. Bulletin épidémiologique hebdomadaire, Hors-série, 14 sept. 2010, 6-8.
• Cheng Z., Wauters L.A., Van Dongen S., Mari V., Romeo C., Martinoli A., Preatoni D. & Tosi G. (2010) Annual variation in predation and dispersal of Arolla pine (Pinus cembra L.) seeds by Eurasian red squirrels and other seed-eaters. Forest Ecology and Management
• Di Pierro E., Bertolino S., Martinoli A., Preatoni D., Tosi g. & Wauters, L.A. (2010) Estimating offspring production using capture-mark-recapture and genetic methods in red squirrels.Ecological Research 25: 395-402.
• Dozieres, A., Pisanu, B., Gerriet, O., Lapeyre, C., Stuyck, J. & Chapuis, J.L. (2010) Macroparasites of Pallas’s squirrels (Callosciurus erythraeus) introduced into Europe. Vet Parasitol 172: 172-176.
• Eason, P. (2010) Alarm signaling in a facultatively social mammal, the southern Amazon red squirrel Sciurus spadiceus.Mammalia 74: 343-345.
• Everest, D. J., Grierson, S. S., Meredith, A. L. & Milne, E. M. (2010) Adenovirus in a red squirrel (Sciurus vulgaris) from Scotland. Vet Rec. 167: 184.
• Everest, D., Stidworthy, M., Milne, E., Meredith, A., Chantrey, J., Shuttleworth, C., Blackett, T., Butler, H., Wilkinson, M. & Sainsbury, A. (2010) Retrospective detection by negative contrast electron microscopy of faecal viral particles in free-living wild red squirrels (Sciurus vulgaris) with suspected enteropathy in Great Britain. Veterinary Record 167:  1007-1010.
• Garroway C., Bowman J., Casceden T., Holloway G., Maham C., Malcolm J., Steele, M.A., Turner, G. Wilson, P.J. (2010) Climate change induced hybridization in flying squirrels. Global Change Biology 16:113-21.
• Garroway, C. J., Bowman, J., Holloway, G. L., Malcolm, J. R. & Wilson, P. J. (2010) The genetic signature of rapid range expansion by flying squirrels in response to contemporary climate warming. Global Change Biology
• Gorrell, J. C., McAdam, A. G., Coltman, D. W., Humphries, M. M. & Boutin, S. (2010) Adopting kin enhances inclusive fitness in asocial red squirrels. Nature Communications 1: Article number 22: 4 pages.
• Hopewell, L., Leaver, L., Lea, S. & Wills, A. (2010) Grey squirrels (Sciurus carolinensis) show a feature-negative effect specific to social learning. Animal Cognition 13: 219-227.
• Johnson, K. M., Boonstra, R. & Wojtowicz, J. M. (2010) Hippocampal neurogenesis in food-storing red squirrels: the impact of age and spatial behavior. Genes, Brain and Behavior 9:  583-591.
• Lane, J. E., Boutin, S., Speakman, J. R. & Humphries, M. M. (2010) Energetic costs of male reproduction in a scramble competition mating system. Journal of Animal Ecology 79: 27-34.
• Larivée, M. L., Boutin, S., Speakman, J. R., McAdam, A. G. & Humphries, M. M. (2010) Associations between over-winter survival and resting metabolic rate in juvenile North American red squirrels. Functional Ecology 24: 597-607.
• LaRose, J. P., Meredith, A., Everest, D., Fiegna, C., McInnes, C., Shaw, D. & Milne, E. (2010) Epidemiological and postmortem findings in 262 red squirrels (Sciurus vulgaris) in Scotland, 2005 to 2009.  Veterinary Record 167: 297-302.
• Lawton, C., Cowan, P., Bertolino, S., Lurz, P. & Peters, A. 2010: The consequences of introducing non-indigenous species: two case studies, the grey squirrel in Europe and the brushtail possum in New Zealand. Rev. sci. tech. Off. int. Epiz. 29, 287-298.
• Lurz, P. (2010) Red Squirrels: Naturally Scottish. Edinburgh: Scottish Natural Heritage, 50 pp.
• Madsen, O., Kortum, T., Hupkes, M., Kohlen, W., van Rheede, T. & de Jong, W. 2010. Loss of octarepeats in two processed prion pseudogenes in the red squirrel, Sciurus vulgaris. Journal of Molecular Evolution 71: 356-363.
• McKinney, S. & Fiedler, C. (2010) Tree squirrel habitat selection and predispersal seed predation in a declining subalpine conifer. Oecologia 162:  697-707.
• Mezquida, E. T. & Benkman, C. W. (2010) Habitat area and structure affect the impact of seed predators and the potential for coevolutionary arms races. Ecology 91: 802-814.
• Newson, S., Leech, D., Hewson, C., Crick, H. & Grice, P. (2010) Potential impact of grey squirrels Sciurus carolinensis on woodland bird populations in England. Journal of Ornithology151: 211-218.
• Newson, S. E., Rexstad, E. A., Baillie, S. R., Buckland, S. T. & Aebischer, N. J. (2010) Population change of avian predators and grey squirrels in England: is there evidence for an impact on avian prey populations? Journal of Applied Ecology 47: 244-252.
• Pelech, S., J.N.M., S. & Boutin, S. (2010) A predator’s perspective of nest predation: predation by red squirrels is learned, not incidental. Oikos 119: 841-851.
• Pisanu, B., Marsot, M., Marmet, J., Chapuis, J-L., Réale, D. Vourc’h, G.(2010). Introduced Siberian chipmunks are more heavily infested by ixodid ticks than are native bank voles in a suburban forest in France. International Journal for Parasitology, 40: 1277-1283.
• Rodrigues D., Wauters L.A., Romeo C., Mari V., Preatoni D., da L. Mathias M., Tosi G. & Martinoli A. (2010) Living on the edge: can Eurasian red squirrels (Sciurus vulgaris) persist in extreme high-elevation habitats? Arctic, Antarctic and Alpine Research 42: 106-112.
• Samaras, A. & Youlatos, D. (2010) Use of forest canopy by European red squirrels Sciurus vulgaris in Northern Greece: claws and small branch niche. Acta theriologica 55: 351-360.
• Scantlebury, M., Maher McWilliams, M., Marks, N. J., Dick, J. T. A., Edgar, H. & Lutermann, H.  (2010) Effects of life-history traits on parasite load in grey squirrels. Journal of Zoology.
• Selonen, V. & Hanski, I. K. (2010) Decision making in dispersing Siberian flying squirrels. Behavioral Ecology 21: 219-225.
• Selonen, V. & Hanski, I. K. (2010) Condition-dependent, phenotype-dependent and genetic-dependent factors in the natal dispersal of a solitary rodent. Journal of Animal Ecology79:1093-1100.
• Selonen, V. & Hanski, I. (2010) Movements of dispersing flying squirrels in relation to siblings and parents. Behavioral Ecology and Sociobiology 64: 1019-1027 .
• Selonen, V., Hanski, I. & Painter, J. (2010) Gene flow in Siberian flying squirrels based on direct and indirect data. Conservation Genetics 11: 1257-1264.
• Selonen, V., Sulkava, P., Sulkava, R., Sulkava, S. & Korpimäki, E. (2010) Decline of flying and red squirrels in boreal forests revealed by long-term diet analyses of avian predators. Animal Conservation 13: 579-585..
• Simpson, V., Davison, N., Hudson, L. & Whatmore, A. M. (2010) Staphylococcus aureus ST49 infection in red squirrels. Vet Rec.167: 69.
• Simpson, V. R., Hargreaves, J., Everest, D. J., Baker, A. S., Booth, P. A., Butler, H. M. & Blackett, T. (2010) Mortality in red squirrels (Sciurus vulgaris) associated with exudative dermatitis. Vet Rec.167: 59-63.
• Wauters L.A., Verbeylen G., Preatoni D., Martinoli A. & Matthysen E. (2010) Dispersal and habitat cuing of Eurasian red squirrels in fragmented habitats. Population Ecology
• Wauters L.A., Preatoni D., Martinoli A., Verbeylen G. & Matthysen E. (2010) No sex bias in natal dispersal of Eurasian red squirrels.Mammalian Biology
• Xiao, Z., Gao, X., Steele, M. A. & Zhang, Z. (2010) Frequency-dependent selection by tree squirrels: adaptive escape of nondormant white oaks. Behavioral Ecology 21: 169-175.

2009

• Amori, G., Gippoliti, S., Luiselli, L. & Battisti, C. (2009). Sciuridae, Rapoport’s effect and the mismatch between range size, conservation needs, and scientific productivity: an approach at the genus level. Web Ecology 9: 1-7.
• Amori, G., Gippoliti, S., Luiselli, L. & Battisti, C. 2009. Do interlinks between geography and ecology explain the latitudinal diversity patterns in Sciuridae? An approach at the genus level. Canadian Journal of Zoology 87: 246-253.
• Andrea, G., Giovanni, A., Gaetano, A., Irene, L., Guido, T., Lucas A, W. & Ettore, R. (2009) Molecular phylogeography of EuropeanSciurus vulgaris: refuge within refugia? Molecular Ecology 18: 2687-2699.
• Bangari, D., Miller, M., Stevenson, G., Thacker, H., Sharma, A. & Mittal, S. (2009) Cutaneous and Systemic Poxviral Disease in Red (Tamiasciurus hudsonicus) and Gray (Sciurus carolinensis) Squirrels. Veterinary Pathology 46: 667-672.
• Bertolino, S. (2009) Animal trade and non-indigenous species introduction: the world-wide spread of squirrels. Diversity and Distributions 15: 701-708.
• Bertolino S., Wauters L.A., Pizzul A., Molinari A., Lurz P.W.W. & Tosi G. (2009) A general approach of using hair-tubes to monitor the European red squirrel: a method applicable at regional and national scales. Mammalian Biology 74: 210-219.
• Carroll, B., Russell, P., Gurnell, J., Nettleton, P. & Sainsbury, A. (2009). Epidemics of squirrelpox virus disease in red squirrels (Sciurus vulgaris): temporal and serological findings.Epidemiology and Infection 137: 247-265.
• Dipineto, L., Gargiulo, A., Cuomo, A., Santaniello, A., Sensale, M., Borrelli, L., D’Angelo, L., Menna, L. & Fioretti, A. (2009) Campylobacter jejuni in the red squirrel (Sciurus vulgaris) population of Southern Italy. Veterinary Journal 179: 149-150.
• Digweed, S. M. & D. Rendall (2009) Predator-associated vocalizations in North American red squirrels, Tamiasciurus hudsonicus: are alarm calls predator specific? Animal Behaviour78: 1135-1144.
• Digweed, S. and D. Rendall (2009) Predator-associated vocalizations in North American red squirrels (Tamiasciurus hudsonicus): to whom are alarm calls addressed and how do they function? Ethology 115: 1190-1199.
• Edelman, A. J., Koprowski, J. L. & Bertelsen, S. R. (2009) Potential for nest site competition between native and exotic tree squirrels. Journal of Mammalogy 90: 167-174.
• Finnegan, L., Poole, A., Lawton, C. & Rochford, J. (2009) Morphological diversity of the red squirrel, Sciurus vulgaris , in Ireland. European Journal of Wildlife Research 55: 145-151.
• Grill A., Amori G., Aloise G., Lisi I., Tosi G., Wauters L.A. & Randi E. (2009) Molecular phylogeography of European Sciurus vulgaris: refuge within refugia? Molecular Ecology 18: 2687-2699.
• Hanski, I. K. & Selonen, V. (2009). Female-biased natal dispersal in the Siberian flying squirrel. Behavioural Ecology 20: 60-67.
• Himsworth, C., Musil, K., Bryan, L. & Hill, J. (2009) Poxvirus infection in an American red squirrel (Tamiasciurus hudsonicus) from northwestern Canada. Journal of Wildlife Diseases 45: 1143-1149.
• Koyabu, D.B., Oshida, T., Dang, N.X., Can D.N., Kimura, J., Sasaki, M.,  Motokawa, M., Son, N.T., Hayashida, A., Shintaku, Y. & Endo, H. (2009) Craniodental mechanics and the feeding ecology of two sympatric callosciurine squirrels in Vietnam.Journal of Zoology 279: 372-80.
• Lane, J. E., Boutin, S., Gunn, M. R. & Coltman, D. W. (2009). Sexually selected behaviour: red squirrel males search for reproductive success. Journal of Animal Ecology 78: 296-304.
• Marmet, J., Pisanu, B. & J Chapuis, J.-L. (2009) Home range, range overlap and site fidelity of introduced Siberian chipmunks in a suburban French forest.European Journal of Wildlife Research, 55: 497-504.
• Mayle, B. A., Proudfoot, J. & Poole, J. (2009) Influence of tree size and dominance on incidence of bark stripping by grey squirrels to oak and impact on tree growth. Forestry 82: 431-444.
• McInnes, C., Coulter, L., Dagleish, M., Fiegna, C., Gilray, J., Willoughby, K., Cole, M., Milne, E., Meredith, A., Everest, D. & Macmaster, A. (2009) First cases of squirrelpox in red squirrels (Sciurus vulgaris) in Scotland. Veterinary Record: 164: 528-531
• Lane, J. E., Boutin, S., Speakman, J. R. & Humphries, M. M. (2010) Energetic costs of male reproduction in a scramble competition mating system. Journal of Animal Ecology 79: 27-34.
• Ozmen, O., Yukari, B. & Haligur, M. (2009) First report of Eimeria lancasterensis in a Red Squirrel (Sciurus vulgaris L.) in Turkey. Turkiye Parazitol Derg 33: 245-247.
• Partan, S. R., Larco, C. P. & Owens, M. J. (2009) Wild tree squirrels respond with multisensory enhancement to conspecific robot alarm behaviour. Animal Behaviour 77: 1127-1135.
• Pisanu, B., Lebailleux, L. & Chapuis, J.-L. (2009).Why Siberian chipmunksTamias sibiricus (Sciuridae) introduced in French forests acquired so few intestinal helminth species from native sympatric Murids?Parasitology Research, 104: 709-714.
• Poole, A. & Lawton, C. (2009) The translocation and post release settlement of red squirrels Sciurus vulgaris to a previously uninhabited woodland. Biodiversity and Conservation 18: 3205-3218.
• Preston, S. & Jacobs, L.F. (2009) Mechanisms of cache  decision making in fox squirrels (Sciurus niger). Journal of Mammalogy90: 787-795.
• Rima P.C., Cagnin M., Aloise G., Preatoni D & Wauters L.A. (2009) Scale-dependent environmental variables affecting red squirrel (Sciurus vulgaris meridionalis) distribution. Italian Journal of Zoology 77: 92-101.
• Salmaso F., Molinari A., Di Pierro E., Ghisla A., Martinoli A., Preatoni D., Cerabolini B., Tosi G., Bertolino S. & Wauters, L.A. (2009) Estimating and comparing food availability for tree-seed predators in typical pulsed-resource systems: alpine conifer forests. Plant Biosystems 143: 258-267.
• Simpson, V., Davison, N., Borman, A., Linton, C. & Everest, D. (2009) Fatal candidiasis in a wild red squirrel (Sciurus vulgaris).Veterinary Record: 164, 342-344.
• Tatsuo, O., Ryuichi, M. & Koichi, I. (2009) Phylogeography of the Japanese giant flying squirrel, Petaurista leucogenys (Rodentia: Sciuridae): implication of glacial refugia in an arboreal small mammal in the Japanese Islands. Biological Journal of the Linnean Society 98: 47-60.
• Vander Wall, S. B., Enders, M. S. & Waitman, B. A. (2009) Asymmetrical cache pilfering between yellow pine chipmunks and golden-mantled ground squirrels. Animal Behaviour 78:  555-561.
• Verbeylen, G., Wauters, L.A., De Bruyn, L. & Matthysen, E. (2009) Woodland fragmentation affects space use of Eurasian red squirrels. Acta Oecologica 35: 94-103.
• Xiao, Z., Gao, X., Jiang, M. & Zhang, Z. (2009) Behavioral adaptation of Pallas’s squirrels to germination schedule and tannins in acorns. Behavioural Ecology 20: 1050-1055.

2008

• Beaucournu, J.-C., Pisanu, B. & Chapuis, J.-L. (2008) Enderleinellus tamiasis Fahrenholz 1916 (Anoplura- Enderleinellidae), espèce importée, nouvelle pour la faune de France. Parasite, 15 : 175-178.
• Bertolino, S. (2008) The introduction of the American grey squirrel (Sciurus carolinensis) in Europe: a case study in biological invasion. Current Science 95: 903-906.
• Bertolino, S., Lurz, P. W. W., Sanderson, R. & Rushton, S. P. (2008) Predicting the spread of the American grey squirrel (Sciurus carolinensis) in Europe: A call for a co-ordinated European approach. Biological Conservation 141: 2564-2575.
• Blois, J. L., Feranec, R. S. & Hadly, E. A. (2008) Environmental influences on spatial and temporal patterns of body-size variation in California ground squirrels (Spermophilus beecheyi).Journal of Biogeography 35: 602-613.
• Boon, A., Réale, D. & Boutin, S. (2008) Personality, habitat use, and their consequences for survival in North American red squirrels Tamiasciurus hudsonicus. Oikos 117: 1321-1328.
• Boonstra, R., Lane, J., Boutin, S., Bradley, A., Desantis, L., Newman, A. & Soma, K. (2008) Plasma DHEA levels in wild, territorial red squirrels: seasonal variation and effect of ACTH.Gen Comp Endocrinol: 158, 61-67.
• Court, I. & Fawcett, H. (2008) Distribution of red squirrels Sciurus vulgaris in the Yorkshire Dales National Park, 1990-2006. Naturalist 133: 55-62.
• Descamps, S., Boutin, S., Berteaux, D., McAdam, A.G., & Gaillard, J.-M. (2008) Cohort effects in red squirrels: the influence of density, food abundance and temperature on future survival and reproductive success. Journal of Animal Ecology 77: 305-314.
• Di Pierro, E., Molinari, A., Tosi, G. & Wauters, L. (2008) Exclusive core areas and intrasexual territoriality in Eurasian red squirrels (Sciurus vulgaris) revealed by incremental cluster polygon analysis. Ecological Research 23: 529-542.
• Finnegan, L., Edwards, C. & Rochford, J. (2008) Origin of, and conservation units in, the Irish red squirrel ( Sciurus vulgaris ) population. Conservation Genetics 9: 1099-1109.
• Freeman, P. & Lemen, C. (2008) A simple morphological predictor of bite force in rodents. Journal of Zoology 275: 418-422.
• Hanski, I. K. & V. Selonen (2009) Female-biased natal dispersal in the Siberian flying squirrel. Behavioral Ecology 20: 60-67.
• Hayssen, V. (2008) Patterns of Body and Tail Length and Body Mass in Sciuridae. Journal of Mammalogy 89: 852-873.
• Hayssen, V. (2008) Reproductive effort in squirrels: ecological, phylogenetic, allometric, and latitudinal patterns. Journal of Mammalogy 89: 582-606.
• Hillegass, M., Waterman, J. & Roth, J. (2008) The influence of sex and sociality on parasite loads in an African ground squirrel.Behavioural Ecology 19: 1006-1011.
• Hopewell, L.J. & Leaver, L.A. (2008). Evidence of social influences on cache-making by grey squirrels (Sciurus carolinensis). Ethology 114(11): 1061-1068.
• Hopewell, L. J., Leaver, L. A. & Lea, S. E. G. (2008) Effects of competition and food availability on travel time in scatter-hoarding gray squirrels (Sciurus carolinensis). Behavioral Ecology 19, 1143-1149.
• Kvac, M., Hofmannova, L., Bertolino, S., Wauters, L., Modr}, D. (2008) Natural infection with two genotypes of Cryptosporidium in red squirrels (Sciurus vulgaris) in Italy. Folia Parasitologica 55: 95-99.
• Lane, J.E., Boutin, S., Gunn, M.R., Slate, J., & Coltman, D.W. (2008) Female multiple mating and paternity in free-ranging North Aerican red squirrels. Animal Behaviour 75: 1927-1937.
• Li, S., F. Yu, et al. (2008). Molecular phylogeny of five species ofDremomys (Rodentia: Sciuridae), inferred from cytochrome b gene sequences. Zoologica Scripta 37: 349-354.
• Lurz, P., Shirley, M. & Geddes, N. (2008) Monitoring low density populations: a perspective on what level of population decline we can truly detect. Animal Biodiversity and Conservation, 31, 29-39.
• Guichon, M.L. & Doncaster, C.P. (2008) Invasion dynamics of an introduced squirrel in Argentina. Ecography 31: 211-220.
• Manjerovic, M., Kinahan, A., Waterman, J., Bennett, N. & Bateman, P. (2008) Structure and allometry of genitalia in males and females of a social African ground squirrel with high polygynandry. Journal of Zoology 275: 375-380.
• Michaux, J., Hautier, L., Simonin, T. & Vianey-Liaud, M. (2008) Phylogeny, adaptation and mandible shape in Sciuridae (Rodentia, Mammalia). Mammalia 72: 286-296.
• Parchman, T. L. and C. W. Benkman (2008). The geographic selection mosaic for ponderosa pine and crossbills: a tale of two squirrels. Evolution 62: 348-360.
• Pisanu, B., Marmet, J., Beaucournu, J.C. & Chapuis, J.-L. (2008). Diversité du cortège en Siphonaptères chez le tamia de Sibérie (Tamias sibiricusLaxmann) introduit en Forêt de Sénart (Ile-de-France). Parasite, 15 : 35-43.
• Perry, D.A.N. & Perry, G.A.D. (2008) Improving interactions between animal rights groups and conservation biologists.Conservation Biology 22: 27-35.
• Preston, S. and L.F. Jacobs, Mechanisms of cache decision making in fox squirrels (Sciurus niger). Journal of Mammalogy, 2009. 90(4): p. 787-795.
• Sainsbury, A., R. Deaville, B. Lawson, W. Cooley, S. Farelly, M. Stack, P. Paul Duff, C. McInnes, J. Gurnell, P. Russell, S. Rushton, D. Pfeiffer, P. Nettleton & P. Lurz (2008) Poxviral disease in red squirrels Sciurus vulgaris in the UK: spatial and temporal trends of an emerging threat. Ecohealth Online 5: 305-16.
• Steele, M.A., Halkin, S.L., Smallwood, P.D., McKenna, T.J., Mitsopoulos, K., & Beam, M. (2008) Cache protection strategies of a scatter-hoarding rodent: do tree squirrels engage in behavioural deception. Animal Behaviour 75, 705-714.
• Wauters, L.A, Githiru, M., Bertolino, S., Molinari, A., Tosi, G., & Lens, L. Demography of alpine red squirrel populations in relation to fluctuations in seed crop size. Ecography 31, 104-114.

2007

• Boon, A.K., Reale, D., & Boutin, S. (2007) The interaction between personality, offspring fitness and food abundance in North American red squirrels. Ecology Letters 10: 1094-1104.
• Descamps, S., Boutin, S., Berteaux, D., & Gaillard, J.-M. (2007) Female red squirrels fit Williams’ hypothesis of increasing reproductive effort with increasing age. Journal of Animal Ecology, 76: 1192-1201.
• Duff, J., Higgins, R., & Farrelly, S. (2007) Enteric adenovirus infection in a red squirrel (Sciurus vulgaris). The Veterinary Record March 17, 2007: 384.
• Edelman, A.J. & Koprowski, J.L. (2007) Communal nesting in asocial Abert’s squirrels: the role of social thermoregulation and breeding strategy. Ethology 113: 147-154.
• Finnegan, L., Hamilton, G., Perol, J. & J Rochford, J. (2007) The use of hair tubes as an indirect method for monitoring red and grey squirrel populations. Biology and the Environment 107B: 55-60.
• Hurme, E., Reunanen, P., Monkkonen, M., Nikula, A., Nivala, V., & Oksanen, J. (2007) Local habitat patch pattern of the Siberian flying squirrel in a managed boreal forest landscape. Ecography30: 277-287.
• Lane, J., Boutin, S., Slate, J., & Coltman, D. (2007) Genetic relatedness of mates does not predict patterns of parentage in North American red squirrels. Animal Behaviour74: 611-619.
• Lawton, C. & Rochford, J. (2007) The recovery of grey squirrel (Sciurus carolinensis) populations after intensive control programmes. Biology and Environment: Proceedings of the Royal Irish Academy 107B: 19-29.
• Leaver, L., Hopewell, L., Caldwell, C. & Mallarky, L. (2007) Audience effects on food caching in grey squirrels (Sciurus carolinensis): evidence for pilferage avoidance strategies.Animal Cognition 10: 23-27.
• Marmet, J. & Chapuis, J.-L. (2007) Répartition de l’Ecureuil de Corée (Tamias sibiricus), animal de compagnie exotique introduit en France : résultats de l’enquête nationale de 2005. Pp : 27-31, in : L. Tillon (ed), Les mammifères forestiers, Actes du 28^ème Colloque de la SFEPM, 21-23 octobre 2005, Rambouillet (78), Collection dossiers forestiers, n° 18.
• Pisanu, B., Jérusalem, C., Huchery, C., Marmet, J. & J.-L. Chapuis (2007).Helminth fauna of the Siberian chipmunk, Tamias sibiricus Laxmann (Rodentia, Sciuridae) introduced in suburban French forests. Parasitology Research, 100: 1375-1379.
• Selonen, V., Hanski, & Desrochers, A. (2007) Natal habitat-biased dispersal in the Siberian flying squirrel. Proceedings of the Royal Society London B 274: 2063-2068.
• Siepielski, A.M. & Benkman, C.W. (2007) Selection by a predispersal seed predator constrains the evolution of avian seed dispersal in pines. Functional Ecology 21: 611-618.
• Signorile, A. & Evans, J. (2007) Damage caused by the American grey squirrel (Sciurus carolinensis) to agricultural crops, poplar plantations and semi-natural woodland in Piedmont, Italy. Forestry, 80: 89-98. Forestry 80: 89-98.
• Tamura, N. & Hayashi, F. (2007) Five-year study of the genetic structure and demography of two subpopulations of the Japanese squirrel (Sciurus lis) in a continuous forest and an isolated woodlot. Ecological Research 22: 261-267.
• Vourc’h, G., Marmet, J., Chassagne, M., Bord, S. & , J.-L. Chapuis (2007)Borrelia burgdorferi sensu lato in Siberian chipmunks (Tamias sibiricus) introduced in suburban forests in France. Vector-Borne and Zoonotic Diseases, 7: 637-642.
• Wauters, L., Preatoni, D., Molinari, A., & Tosi, G. (2007) Radio-tracking squirrels: performance of home range density and linkage estimators with small range and sample size. Ecological Modelling 202: 333-344.
• Wauters, L.A., Vermeulen, M., Van Dongen, S., Bertolino, S., Molinari, A., Tosi, G., & Matthysen, E. (2007) Effects of spatio-temporal variation in food supply on red squirrel Sciurus vulgaris body size and body mass and its consequences for some fitness components. Ecography 30: 51-65.

2006

• Boutin, S., Wauters, L., McAdam, A., Humphries, M., Tosi, G., & Dhondt, A. (2006) Anticipatory reproduction and population growth in seed predators. Science 314, 1928 – 1930.
• Carvalho, L.d.S., Cowing, J., Wilkie, S., Bowmaker, J., & Hunt, D. (2006) Shortwave visual sensitivity in tree and flying squirrels reflects changes in lifestyle. Current Biology 16: R81-R83.
• Edelman, A.J. & Koprowski, J.L. (2006) Diet and tree use of Abert’s squirrels (Sciurus aberti) in a mixed-conifer forest.Southwestern Naturalist 50: 461-465.
• Edelman, A.J. & Koprowski, J.L. (2007) Communal Nesting in Asocial Abert’s Squirrels: the Role of Social Thermoregulation and Breeding Strategy. Ethology 113, 147-154.
• Ferryman, M., Mayle, B., & Morgan, G. (2006) Visual methods for evaluating the state of sexual development in male grey squirrels (Sciurus carolinensis). Reproduction, Fertility and Development 18, 283-293.
• Gurnell, J., Rushton S.P., Lurz, P.W.W., Sainsbury, A.W., Nettleton, P, Shirley, M.D.F., Bruemmer, C. & Geddes, N. (2006) Squirrel poxvirus; landscape scale strategies for managing disease threat. Biological Conservation 131: 287-295.
• Jansen, P.A., Bongers, F., & H. T. Prins, H. (2006) Tropical rodents change rapidly germinating seeds into long-term food supplies. Oikos 113, 449-458.
• Meijaard, E., Kitchener, A.C., & Smeenk, C. (2006) ‘New Bornean carnivore’ is most likely a little known flying squirrel. Mammal Review 36, 318-324.
• McInnes, C.J., Wood, A.R., Thomas, K., Sainsbury, A.W., Gurnell, J., Dein, F.J. and Nettleton. P.F. (2006) Genomic characterisation of a novel poxvirus contributing to the decline of the red squirrel (Sciurus vulgaris) in the UK. Journal General Virology 87: 2115-2125.
• Molinari, A., Wauters L.A., Airoldi, G., Cerinotti, F., Martinoli, A. & Tosi, G.(2006) Cone selection by Eurasian red squirrels in mixed conifer forests in the Italian Alps.Acta Oecologica 30: 1-10. (doi:10.1016/j.actao.2005.11.004).
• Moore, J. & Swihart, R. (2006) Modeling patch occupancy by forest rodents: incorporating detectability and spatial autocorrelation with hierarchically structured data. Journal of Wildlife Management 69, 933-949.
• Ozgul, A., Armitage, K.B., Blumstein, D.T., Vanvuren, D.H., & Oli, M.K. (2006) Effects of patch quality and network structure on patch occupancy dynamics of a yellow-bellied marmot metapopulation. Journal of Animal Ecology 75, 191-202.
• Robitaille, J.-F. & Linley, R.D. (2006) Structure of forests used by small mammals in the industrially damaged landscape of Sudbury, Ontario, Canada. Forest Ecology and Management 225: 160-167.
• Rushton S.P., Lurz, P.W.W., Gurnell, J., Nettleton, P. , Bruemmer, C.,Shirley, M.D.F. and Sainsbury, A.W. (2006) Disease threats posed by alien species: the role of a poxvirus in the decline of the native red squirrel in Britain. Epidemiology and Infection134: 521-533.
• Selonen,V. & Hanski,I.K. (2006) Habitat exploration and use in dispersing juvenile flying squirrels. Journal of Animal Ecology75:1440-1449.
• Simpson, V., Birtles, R., Bown, K., Panciera, R., Butler, H. & Davison, N. (2006) Hepatozoon species infection in wild red squirrels (Sciurus vulgaris) on the Isle of Wight. Veterinary Record 159: 202-205.
• Steele, M.A., Manierre, S., Genna, T., Contreras, T.A., Smallwood, P.D., & Pereira, M.E. (2006) The innate basis of food-hoarding decisions in grey squirrels: evidence for behavioural adaptations to the oaks. Animal Behaviour 71: 155-160.
• Sushma, H.S. & Singh, M. (2006) Resource partitioning and interspecific interactions among sympatric rain forest arboreal mammals of the Western Ghats, India. Behavioral Ecology 17: 479-490.
• Tattoni, C., Preatoni D.G., Lurz, P.W.W., Rushton, S.P., Tosi, G., Bertolino, S., Martinoli, A., & Wauters, L.A. (2006) Modelling the expansion of a grey squirrel population: implications for squirrel control. Biological Invasions8: 1605-1619.
• Thorington Jr, R. & Ferrell, K. (2006) Squirrels: The Animal Answer Guide. 208 pp. John Hopkins University Press, Baltimore, USA. Link to JHUP.
• Wagner, D.M., Drickamer, L.C., Krpata, D.M., Allender, C.J., Van Pelt, W.E., & Keim, P. (2006) Persistence of Gunnison’s prairie dog colonies in Arizona, USA. Biological Conservation 130, 331-339.
• Winterrowd, M.F. & Weigl, P.D. (2006) Mechanisms of Cache Retrieval in the Group Nesting Southern Flying Squirrel (Glaucomys volans). Ethology 112, 1136-1144.
• Xiao, Z., Jansen, P.A., & Zhang, Z. (2006) Using seed-tagging methods for assessing post-dispersal seed fate in rodent-dispersed trees. Forest Ecology and Management 223: 18-23.

2005

• Bertolino, S. & Genovesi, P. (2005) The application of the European strategy on invasive alien species: an example with introduced squirrels. Hystrix Italian Journal of Mammalogy 16: 59-69.
• Bryce, J., Cartmel, S. & Quinn, C.P. (2005) Habitat use by red and grey squirrels: results of two recent studies and implications for management. Information Note. Forestry Commission, Edinburgh. 12pp.
• Chapuis, J.-L. (2005) Répartition en France d’un animal de compagnie naturalisé, le Tamia de Sibérie (Tamias sibiricus). Revue d’Ecologie (Terre Vie), 60 : 239-253.
• Edelman, A.J. & Koprowski, J.L. (2005) Selection of drey sites by Abert’s squirrels in an introduced population. Journal of Mammalogy, 86(6):1220-1226.
• Edelman, A.J., Koprowski, J.L. & Edelman, J.R. (2005) Kleptoparasitic behavior and species richness at Mt. Graham red squirrel middens. In Gottfried, Gerald J.; Gebow, Brooke S.; Eskew, Lane G.; and Edminster, Carleton B., compilers.Connecting mountain islands and desert seas: biodiversity and management of the Madrean Archipelago II. 2004 May 11-15; Tucson, AZ. Proceedings RMRS-P-36. pp 395-398. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station. 631 p.
• Gerhardt, F. (2005) Food pilfering in larder-hoarding red squirrels (Tamiascsiurus hudsonicus). Journal of Mammalogy86: 108-114.
• Goheen, J. & Swihart, R. (2005) Resource selection and predation of North American red squirrels in deciduous forest fragments.Journal of Mammalogy 86: 22-28.
• Hurme, E., Monkkonen, M., Nikula, A.,Nivala, V., Reunanen, P., Heikkinen, T., & Ukkola, M. (2005) Building and evaluating predictive occupancy models for the Siberian flying squirrel using forest planning data. Forest Ecology and Management216:241-256.
• Koprowski, J.L. (2005) Management and conservation of tree squirrels: the importance of endemism, species richness, and forest condition. In Gottfried, Gerald J.; Gebow, Brooke S.; Eskew, Lane G.; and Edminster, Carleton B., compilers.Connecting mountain islands and desert seas: biodiversity and management of the Madrean Archipelago II. 2004 May 11-15; Tucson, AZ. Proceedings RMRS-P-36. pp. 245-250. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station. 631 p.
• Koprowski, J.L. (2005) Annual cycles in body mass and reproduction of endangered Mt. Graham red squirrels. Journal of Mammalogy 86: 309-313.
• Koprowski, J.L. (2005) The response of tree squirrels to fragmentation: a review and synthesis. Animal Conservation 8: 369-376.
• Koprowski, J.L. (2005, July 20). Pine Squirrel (Tamiasciurus hudsonicus): a technical conservation assessment. [Online]. USDA Forest Service, Rocky Mountain Region. Available: www.fs.fed.us/r2/projects/scp/assessments/pinesquirrel.pdf [date of access].
• Koprowski, J.L., Alanen, M.I. & Lynch, A.M.(2005) Nowhere to run and nowhere to hide: Response of endemic Mt. Graham red squirrels to catastrophic forest damage. Conservation Biology126: 491-498.
• Koprowski, J.L. & Corse, M.C. (2005) Time budgets, activity periods, and behavior of Mexican fox squirrels. Journal of Mammalogy 86: 947-952.
• Koprowski, J.L. Edelman, A.J., Pasch, B.S. and Buecher, D.C. (2005) A dearth of data on the mammals of the Madrean Archipelago: What we think we know and what we actually do know. In Gottfried, Gerald J.; Gebow, Brooke S.; Eskew, Lane G.; and Edminster, Carleton B., compilers. Connecting mountain islands and desert seas: biodiversity and management of the Madrean Archipelago II. 2004 May 11-15; Tucson, AZ. Proceedings RMRS-P-36. pp 412-415. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station. 631 p.
• Lawton, C. (Ed.) (2005). Proceedings of the Irish Red Squirrel Conservation Symposium , 22nd April, 2005. 20pp. National University of Ireland, Galway. (PDF)
• Lu, J. & Zhang, Z. (2005) Food hoarding behaviour of David’s rock squirrel Sciurotamias davidianus. Acta Zoologica Sinica 51: 376-382.
• Mezquida, E.T. & Benkman, C.W. (2005) The geographic selection mosaic for squirrels, crossbills and Aleppo pine.Journal of Evolutionary Biology 18: 348-357.
• Ogden, R., Shuttleworth, C., McEwing, R. & Cesarini, S. (2005) Genetic management of the red squirrel, Sciurus vulgaris: a practical approach to regional conservation. Conservation Genetics 6: 511-525.
• Oshida, T., AramovV, A., Yanagawa, H., & Masuda, R. (2005) Phylogeography of the Russian flying squirrel (Pteromys volans): implication of refugia theory in arboreal small mammal of Eurasia. Molecular Ecology, 14, 1191-1196.
• Pasch, B.S. & Koprowski, J.L. (2005) Correlates of vulnerability in Chiricahua fox squirrels. In Gottfried, Gerald J.; Gebow, Brooke S.; Eskew, Lane G.; and Edminster, Carleton B., compilers.Connecting mountain islands and desert seas: biodiversity and management of the Madrean Archipelago II. 2004 May 11-15; Tucson, AZ. Proceedings RMRS-P-36. pp 426-428. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station. 631 p.
• Selonen, V., Painter, J. N. & Hanski, I. K. (2005) Microsatellite variation in the Siberian flying squirrel in Finland. Annales Zoologici Fennici 42: 505:511.
• Steele, M., Wauters, L.A. & Larsen, K.A. (2005) Selection, Predation and Dispersal of Seeds by Tree Squirrels in Temperate and Boreal Forests: are Tree Squirrels Keystone Granivores? InSeed Fate (Eds. P.-M. Forget, J.E. Lambert, P.E. Hulme and S.B. Vander Wall). pp 205-221. CAB International.
• Tattoni, C., Preatoni, D., Martinoli, A., Bertolino, S. & Wauters, L. (2005) Application of modelling tecniques to manage a population of grey squirrels (Sciurus carolinensis) in Lombardy, northern Italy, and analysis of parameters estimates used in simulations. Hystrix It. J. Mamm. 16: 99-112.
• Trizio, I., B. Crestanello,B., Galbusera, P., Wauters, L.A., Tosi, G. Matthysen, E. & Hauffe, H. (2005) Geographical distance and physical barriers shape the genetic structure of Eurasian red squirrels (Sciurus vulgaris) in the Italian Alps. Molecular Ecology14: 469:481.
• Wauters,L.A., Bertolino, S., Adamo, M., Van Dongen, S., Tosi, G. (2005) Food shortage disrupts social organization: the case of red squirrels in conifer forests. Evolutionary Ecology 19:375-404
• Wauters, L., Tosi, G., & Gurnell, J. (2005) A review of the competitive effects of alien grey squirrels on behaviour, activity and habitat use of red squirrels in mixed, deciduous woodland in Italy. Hystrix It. J. Mamm. 16: 27-40.

2004

• Bakker, V.J.& Van Vuren, D.H. (2004) Gap-crossing decisions by the Red Squirrel, a forest-dependent small mammal.Conservation Biology 18: 689-697.
• Gurnell, J. (2004) Grey Squirrels Invade Europe. Mammals UK. Mammals Trust UK, London
• Gurnell,J., Lurz, P.W.W., Shirley, M.D.F., Cartmel, S., Garson, P.J., Magris, L., & Steele, J. (2004) Monitoring red (Sciurus vulgaris) and grey (Sciurus carolinensis) squirrels in Britain. Mammal Review 34: 51-74.
• Gurnell, J., Wauters, L.A., Lurz, P.W.W. & Tosi, G. (2004) Alien species and interspecific competition: effects of introduced eastern grey squirrels on red squirrel population dynamics.Journal of Animal Ecology 73: 26-35.
• Hale, M. L., Lurz, P. W. W. & Wolff, K. (2004). Patterns of genetic diversity in the red squirrel: footprints of biogeographic history and artificial introductions. Conservation Genetics 5:167-179.
• Haughland, D. & Larsen, K. (2004) Ecology of North American red squirrels across contrasting habitats: relating natal dispersal to habitat.Journal of Mammalogy 85: 225-236.
• Haughland, D.L. & Larsen, K.W. (2004) Exploration correlates with settlement: red squirrel dispersal in contrasting habitats.Journal of Animal Ecology 73: 1024-1034.
• Hewson, C., Fuller, R., Mayle, B. & Smith, K. (2004) Possible impacts of grey squirrels on birds and other wildlife. British Wildlife 15: 183-191.
• Lehmkuhl, J.F. Gould, L.E.,Cázaresc, E & Hosford, D.R. (2004) Truffle abundance and mycophagy by northern flying squirrels in eastern Washington forests. Forest Ecology and Management200: 49-65.
• Linders, M., West, S.& Vander Haegen, W.M. (2004) Seasonal variability in the use of space by western gray squirrels in Southcentral Washington. Journal of Mammalogy 85: 511-516.
• Mayle, B. & Gurnell, J. (2004) Increasing efficiency of grey squirrel control in conifer habitats. Forestry and British TimberApril 2004.
• Mayle, B. & Gurnell, J. (2004) Comparison of trap location and bait type for grey squirrel control in conifers. 20 pp. Report to the Sustainable Forestry Group, Forestry Commission, Edinburgh.
• Miyamoto, A., Tamura, N., Sugimura, K. & Yamada, F.(2004) Predicting habitat distribution of the alien Formosan squirrels using logistic regression model. Global Environmental Research8: 13-21.
• Painter, J., Selonen, V. & Hanski, I. K. (2004) Microsatellite loci for the Siberian flying squirrel, Pteromys volans. Molecular Ecology Notes 4:119-121.
• Sainsbury, A., Kountouri, A., DuBoulay, G., & Kertesz, P. (2004) Oral disease in free-living red squirrels (Sciurus vulgaris) in the United Kingdom.Journal of Wildlife Diseases, 40, 185-196.
• Selonen, V. & Hanski, I. K. (2004) Young flying squirrels (Pteromys volans)dispersing in fragmented forests. Behavioral Ecology 15: 564-571.
• Tamura, N. (2004) Effects of habitat mosaic on home range size of the Japanese squirrel Sciurus lis.” Mammal Study 29: 9-14.
• Wauters L.A., Zaninetti M., Tosi G., Bertolino S.(2004) Is coat-colour polymorphism in Eurasian red squirrels (Sciurus vulgarisL.) adaptive? Mammalia 68:37-48.
• Wauters, L.A., Matthysen, E. Adriaensen, F. & Tosi, G. (2004) Within-sex density-dependence and population dynamics of red squirrels Sciurus vulgaris Journal of Animal Ecology 73:11-25.
• Wells, K., Pfeiffer, M., Lakim, M.B. & Linsenmair, K.E. (2004) Use of arboreal and terrestrial space by a small mammal community in a tropical rain forest in Borneo, Malaysia. Journal of Biogeography 31: 641-652.

2003

• Anon (2003)Urban grey squirrels. DEFRA Rural Develeopment Setrvices Technical Advisory Note WM09 [PDF]
• Bertolino, S. Wauters, L.A., De Bruyn L. & Canestri-Trotti, G. (2003) Prevalence of coccidia parasites (Protozoa) in red squirrels ( Sciurus vulgaris): effects of host phenotype and environmental factors. Oecologia 137, 286-295.
• Desrochers, A., Hanski, I. K. & Selonen, V. (2003) Siberian flying squirrel responses to high- and low- contrast forest edges.Landscape Ecology 18:543-552.
• Goheen, J. & Swihart, R. (2003) Food-hoarding behavior of gray squirrels and North American red squirrels in the central hardwoods region: implications for forest regeneration.Canadian Journal of Zoology, 81, 1636-1639.
• Goheen, J., Swihart, R., Gehring, T., & Miller, M. (2003) Forces structuring tree squirrel communities in landscapes fragmented by agriculture: species differences in perceptions of forest connectivity and carrying capacity. Oikos, 102, 95-103.
• Goheen, J., Swihart, R., & Robins, J. (2003) The anatomy of a range expansion: changes in cranial morphology and rates of energy extraction for North American red squirrels from different latitudes. Oikos, 102, 33-44.
• Hale, M. L. & Lurz, P. W. W. (2003). Morphological changes in a British mammal as a result of introductions and changes in landscape management: the red squirrel (Sciurus vulgaris).Journal of Zoology, London: 260: 159-167.
• Hutton KA, Koprowski JL, Greer VL, Alanen MI, Schauffert CA & Young PJ. (2003) Use of mixed-conifer and spruce-fir forests by an introduced population of Abert’s squirrels (Sciurus aberti).Southwestern Naturalist 48:257-260.
• Lurz, P. W. W., Geddes, N., Lloyd, A. J., Shirley, M. D. F., Rushton, S. P. & Burlton, B. (2003) Planning a red squirrel conservation area: using a spatially explicit population dynamics model to predict the impact of felling and forest design plans.Forestry, 76, 95-108.
• Lurz, P. W. W., Rushton, S. P. & Gurnell. J. (2003) Kielder Red Squirrel Project. Report for FE Kielder Forest and Mammals Trust UK. 22 pp.
• Petty, S. J., Lurz, P. W. W. & Rushton, S. P. (2003) Predation of red squirrels by northern goshawks in a conifer forest in northern England: can this limit squirrel numbers and create a conservation dilemma? Biological Conservation, 111, 105-114.
• Réale, D., McAdam, A., Boutin, S. & Berteaux, D. (2003) Genetic and plastic response of a northern mammal to climate change. Proceedings of the Royal Society of London B 270: 591-596.
• Réale, D., Berteaux, D., McAdam, A. & Boutin, S. (2003) Lifetime selection on heritable life-history traits in a natural population of red squirrels. Evolution 57: 2416-2423.
• Selonen, V. & Hanski, I. K. (2003) Movements of the flying squirrel Pteromys volans in corridors and in matrix habitat.Ecography 26:641-651.
• Stapleford, D. (2003) An Affair with Red Squirrels The Larks Press, Ordnance Farmhouse, Guist Bottom, Dereham, Norfolk NR20 SPF UK. Price £5. 58 pp.
• Swihart, R., Atwood, T., Goheen, J., Scheiman, D., Munroe, K., & Gehring, T. (2003) Patch occupancy of North American mammals: is patchiness in the eye of the beholder? Journal of Biogeography, 30, 1259-1279.
• Tompkins, D., Sainsbury, A.W., Netteleton, P., Buxton, D. & Gurnell. J. (2002) Parapoxvirus causes a deleterious disease of red squirrels associated with UK population declines.Proceedings of the Royal Society of London 269: 529-533.
• Verbeylen, G., De Bruyn, L. & Matthysen, E. (2003) Patch occupancy, population density and dynamics in a fragmented red squirrel Sciurus vulgaris population. Ecography 26, 118-128.

2002

• Barr, J. J., Lurz, P. W. W., Shirley, M. D. F. & Rushton, S. P. (2002) Evaluation of immunocontraception as a publicly acceptable form of vertebrate pest species control: the introduced grey squirrel in Britain as an example. Environmental Management, 30, 342-351.
• Bertolini,S; Genovesi,P (2002) Spread and attempted eradication of the grey squirrel (Sciurus carolinensis) in Italy, and consequences for the red squirrel (Sciurus vulgaris) in Eurasia.Biological Conservation 109, 351-358.
• Bryce,J; Johnson,PJ; Macdonald,DM (2002) Can niche use in red and grey squirrels offer clues for their apparent coexistence?Journal of Applied Ecology 39, 875-887.
• Gurnell, J., Cartmel, S., Lurz, P.W.W. & Rushton, S.P. (2002)Modelling the effects of forest composition and management on conserving red squirrels in Clocaenog Forest, North Wales.Unpublished Report for Forestry Commission and Countryside Council for Wales.
• Gurnell, J. & Steele, J. (2002) Grey Squirrel Control for Red Squirrel Conservation – Thetford Forest. Published Report for English Nature and the Forestry Commission.
• Gurnell, J., Clark, M.J., Lurz, P.W.W., Shirley, M.D.F. & Rushton, S.P. (2002) Conserving red squirrels (Sciurus vulgaris): mapping and forecasting habitat suitability using a Geographic Information Systems Approach. Biological Conservation 105: 53-64.
• Koprowski, J.L. (2002) Handling tree squirrels with an efficient and safe restraint. Wildlife Society Bulletin 30:101-103.
• Kurtila,M; Pukkala,T; Loikkanen,J (2002) The performance of alternative spatial objective types in forest planning calculations: a case for flying squirrel and moose. Forest Ecology and Management 166, 245-260.
• Lee,TH (2002) Feeding and hoarding behaviour of the Eurasian red squirrel Sciurus vulgaris during autumn in Hokkaido, Japan.Acta theriologica 147, 459-470.
• Magris, L. & Gurnell, J. (2002) Population ecology of the red squirrel (Sciurus vulgaris) in a fragmented woodland ecosystem on the Island of Jersey, Channel islands. Journal of Zoology, London 256: 99-112.
• McAdam,AG; Boutin,S; R‚ale,D; Berteaux,D (2002) Maternal effects and the potential for evolution in a natural population of animals. Evolution 56, 846-851.
• Schauffert, C., J. Koprowski, V. Greer, M. Alanen & K. Hutton. (2002) Potential predators of Mt. Graham red squirrels.Southwestern Naturalist 47:498-501.
• Wauters, L.A., Tosi, G. & Gurnell, J. (2002) Interspecific competition of grey on reds: do grey squirrels deplete tree seeds cached by red squirrels. Behavioral Ecology and Sociobiology 51:360-367
• Tompkins, D., Sainsbury, A.W., Netteleton, P., Buxton, D. & Gurnell. J. (2002) Parapoxvirus causes a deleterious disease of red squirrels associated with UK population declines.Proceedings of the Royal Society of London 269: 529-533.

2001

• • Gurnell, J. (2001) Red squirrels: a species in decline. Mammals UK Summer 2001 Pp7-8, Mammals Trust UK, London.
  • Gurnell, J., Lurz, P.P.W. & Pepper, H. (2001) Practical techniques for surveying and monitoring squirrels. Forestry Commission Practice Note 11. Forestry Commission, Edinburgh. 12pp.
  • Gurnell, J. , Wauters, L.A., Preatoni, D. & Tosi, G. (2001) Spacing behaviour, kinship and dynamics of the grey squirrel in a newly colonised deciduous woodland in north Italy. Canadian Journal of Zoology 79: 1533-1543.
  • Hale, M. L., Peter W.W. Lurz, Mark D.F. Shirley, Steven Rushton, Robin M. Fuller & Kirsten Wolff (2001). The Impact of Landscape Management on the Genetic Structure of Red Squirrel Populations. Science 293: 2246-2248.
  • Koprowski, J.L. & M.C. Corse (2001) Food habits of Chiricahua Fox Squirrels (Sciurus nayaritensis chiricahuae). Southwestern Naturalist, 46:62-65.
  • Lurz, P.W.W., Rushton, S.P., Wauters, L.A., Bertolino, S., Corrado, I., Mazzoglio, P. & Shirley, M.D.F. (2001). Predicting gray squirrel expansion in North Italy: a spatially explicit modelling approach. Landscape Ecology , 16, 407-420
  • Sainsbury, A.W., Adair, B., Graham, D., Gurnell, J., Cunningham, A.A., Benko, M. & Papp, T. (2001) Isolation of a novel adenovirus associated with splenitis, diarrhoea, and mortality in translocated red squirrels, Sciurus vulgaris. Verh. ber. Ekrg. Zootiere 40: 265-270.

Selonen, V., Hanski, I. K & Stevens, P. (2001) Space use of Siberian flying squirrel Pteromys volans in fragmented forest landscapes. Ecography 24:588-600.

• Shuttleworth, C.M. & Gurnell, J. (2001) The management of coastal sand dune woodland for red squirrels (Sciurus vulgaris L.). In Coastal Dune Management: Shared Experience of European Conservation Practice (eds. J.A. Houston, S.E. Edmondson & P.J. Rooney) pp. 117 – 127. Liverpool University Press, Liverpool, England.
• Steele, M.A. & Koprowski, J.L. (2001) North American Tree Squirrels Smithsonian Institution Press, 201 pp.
• Wauters, L.A., Gurnell, J., Martinoli, A. & Tosi, G. (2001) Does interspecific competition with grey squirrels affect the foraging behaviour and food choice of red squirrels. Animal Behaviour 61: 1079-1091.
• Wauters, L.A., Gurnell, J. ,Preatoni, D. & Tosi, G (2001) Effects of spatial variation in food availability on spacing behaviour and demography of Eurasian red squirrels. Ecography 24: 525-538.

2000

• Bruemmer, C., Lurz, P., Larsen, K. & Gurnell, J. (2000) Impacts and management of the alien Eastern Gray Squirrel in Great Britain and Italy: lessons for British Columbia. Pp. 341 – 349. In L. M. Darling (editor). Proceedings of a Conference on the Biology and Management of Species and Habitats at Risk, Kamloops, B.C., 15 – 19 Feb., 1999. Volume One. B.C. Ministry of Environment, Lands and Parks, Victoria, B.C. and University College of the Cariboo, Kamloops, B.C. 490 pp.
• Cagnin, M., Aloise, G., Fiore, F., Oriolo, V. & Wauters, L.A. (2000). Habitat use and population densityof the red squirrel, Sciurus vulgaris meridionalis, in the Sila Grande mountain range (Calabria, South Italy). Ital. J. Zool. 67: 81-87.
• Lurz, P.W.W., Garson, P.J. & Wauters, L.A.. (2000). Effects of temporal and spatial variations in food supply on the space and habitat use of red squirrels, Sciurus vulgaris L. J. Zool., Lond. 251: 167-178.
• Rushton, S.P., Lurz, P.W.W. & Gurnell, J. (2000) Modelling the spatial dynamics of parapoxvirus disease in red and grey squirrels: a possible cause of the decline in the red squirrel in the United Kingdom? Journal of Applied Ecology 37:1-18.
• Sainsbury, A.W., Nettleton, P., Gilray, J. & Gurnell, J. (2000) Grey squirrels have high seroprevalence to a parapoxvirus associated with deaths in red squirrels. Animal Conservation 3: 229-233.
• Verbeylen, G. & De Bruyn, L. (2000). Inventarisatie van de Aziatische grondeekhoorn in De Panne. Een project van het IBW en AMINAL afdeling Natuur. Rapport 3 – September 2000.
• Wauters, L.A. (2000). Squirrels – Medium-sized Granivores in Woodland Habitats. – In: Halle, S. and Stenseth N. C. (eds), Activity Patterns in Small Mammals: a Comparative Ecological Approach. Ecological Studies 141, Springer-Verlag, Heidelberg, Germany, pp. 131-143.
• Wauters, L.A., Lurz, P.W.W. & Gurnell, J. (2000). The effects of interspecific competition by grey squirrels (Sciurus carolinensis) on the space use and population dynamics of red squirrels (S. vulgaris) in conifer plantations. Ecological Research 15: 271-284.
• Teangana, D.Ó., Russ, J.M., Mathers, R.G., & Montgomery, W.I. (2000) Habitat associations of the red squirrel Sciurus vulgaris and grey squirrel S. carolinensis in Northern Ireland. Biology and Environment: Proceedings of the Royal Irish Academy, Vol. 100B, No. 1, 27-33.

1999

• Barratt, E.M., Gurnell, J., Malarky, G. Deaville, R. & Bruford, M.W. (1999) Genetic structure of fragmented populations of red squirrel (Sciurus vulgaris) in Britain. Molecular Ecology 8: S55-S63.
• David-Gray, Z.K., Gurnell, J. & Hunt, D.M. (1999) Estimating the relatedness in a population of grey squirrels, Sciurus carolinensis, using DNA fingerprinting. Acta theriologica 44: 243-251.
• David-Gray, Z.K., Gurnell, J. & Hunt, D.M. (1999) DNA fingerprinting reveals high levels of genetic diversity in British populations of the introduced non-native grey squirrel, Sciurus carolinensis. Journal of Zoology, London 246: 443-486.
• Gurnell, J. (1999) Grey squirrels in woodlands: managing grey squirrels to prevent woodland damage. Enact 7: 10-14.
• Gurnell, J. and Wauters, L. (1999) Sciurus vulgaris. In The Atlas of European Mammals. Mitchell-Jones, A.J., Amori, G., Bogdanowicz, W., Krystufek, B., Reijnders, P.J.H., Spitzenberger. F., Stubbe, M., Thissen, J.B.M., Vohohralik, V. & Zima, J. (Eds.) pp. 180-181. Academic press, London.
• Gurnell, J. and Wauters, L. (1999) Sciurus carolinensis. In The Atlas of European Mammals. Mitchell-Jones, A.J., Amori, G., Bogdanowicz, W., Krystufek, B., Reijnders, P.J.H., Spitzenberger. F., Stubbe, M., Thissen, J.B.M., Vohohralik, V. & Zima, J. (Eds.) pp. 178-179. Academic press, London.
• Gurnell, J. and Wauters, L. (1999) Callosciurus erythraeus. In The Atlas of European Mammals. Mitchell-Jones, A.J., Amori, G., Bogdanowicz, W., Krystufek, B., Reijnders, P.J.H., Spitzenberger. F., Stubbe, M., Thissen, J.B.M., Vohohralik, V. & Zima, J. (Eds.) pp. 182-183. Academic press, London.
• Garvish, L. and Gurnell, J. (1999) Sciurus anomalus. In The Atlas of European Mammals. Mitchell-Jones, A.J., Amori, G., Bogdanowicz, W., Krystufek, B., Reijnders, P.J.H., Spitzenberger. F., Stubbe, M., Thissen, J.B.M., Vohohralik, V. & Zima, J. (Eds.) pp. 176-177. Academic press, London.
• Massingham Hart, R. (1999) Squirrel Proofing your Home and Garden. Workman Publishing Company. ISBN: 1580171915
• Mathias, M.L. & Gurnell, J. (1999) Status and conservation of the red squirrel (Sciurus vulgaris L) in Portugal. Hystrix 10: 13-19.
• Osborn, D.A. (1999) Squirrel dog basics: a guide to hunting squirrels with dogs. Tree Top Publications, Watkinsville, Georgia, 151 pp.
• Rushton, S.P., Lurz, P.W.W., South, A.B. & Mitchell-Jones, A. (1999). Modelling the distribution of red squirrels on the Isle of Wight. Animal Conservation, 2, 111-121
• Verbeylen, G., De Bruyn, L. & Matthysen, E. (1999). Inventarisatie van de Aziatische grondeekhoorn in De Panne. Een project van de UA groep Dierenecologie in opdracht van AMINAL afdeling Natuur. Rapport 2 – Augustus-Oktober 1999.
• Wauters, L.A. & Gurnell, J. (1999) The mechanism of replacement of red squirrels by grey squirrels: a test of the interference competition hypothesis. Ethology 105: 1053-1071.

Selected Pre-1999

• Lurz, P.W.W., Garson, P.J. & Rushton, S.P., (1995). The ecology of squirrels in spruce dominated plantations: implications for management. Forest Ecology and Management, 79, 79-90.
• Lurz, P.W.W., Garson, P.J., Wauters, L. (1997). Effect of temporal and spatial variation in habitat quality on red squirrel, Sciurus vulgaris, dispersal behaviour. Animal Behaviour 54, 427-435.
• Lurz, P.W.W., Garson P.J. & Ogilvie J.F. (1998) Conifer species mixtures, cone crops and red squirrel conservation Forestry 71, 67-71.
• Rushton, S.P., Lurz, P.W.W., Fuller, R. & Garson, P.J. (1997). Modelling the distribution of the red and grey squirrel at the landscape scale: a combined GIS and population dynamics approach. J. Appl. Ecol., 34, 1137-1154.
• Rushton, S.P. , Lurz, P.W., Fuller, R. and Garson, P.J. (1997) Modelling the distribution of the red and grey squirrel at the landscape scale: a combined GIS and population dynamics approach. J. Appl. Ecol, 34, 1137-1154.
• Verbeylen, G. & Matthysen, E. (1998). Inventarisatie van de Aziatische grondeekhoorn in De Panne. Een project van de UIA groep Dierenecologie in opdracht van AMINAL afdeling Natuur. Rapport September-November 1998.
• Verbeylen, G. & Wauters, L. (1997). Habitatversnippering en eekhoorns : een vergelijking met vogelstudies. Limosa, 70, 38.
• Villalba, S., Gulinck, H., Verbeylen, G. & Matthysen, E. (1998). Relationship between patch connectivity and the occurrence of the European red squirrel, Sciurus vulgaris, in forest fragments withi n heterogeneous landscapes. In : Dover, J.W. & Bunce, R.G.H. (eds.). Key Concepts in Landscape Ecology. IALE(UK), Preston, 205-220.

Bedtime listening

On Monday 4th August 2003, a radio programme was broadcast of BBC Radio 4 at 21.00 (British Summer Time) called, Can the red squirrel survive in Europe if the grey squirrel spreads over the Italian Alps?. To quote from the BBC website:

“Lionel Kelleway travels to Italy, to discover how the grey squirrel arrived there after World War Two and has now travelled north to the Italian Alps. In its wake it has devastated the native red squirrel population and is poised to disperse across Europe. With the help of continuous forest and few physical barriers, their spread is likely to be rapid. Lionel asks if there is hope for the red squirrel if the grey squirrel crosses Italy’s border and colonises the rest of mainland Europe.”

The programme features Piero Genovesi, Italo Currado, Sandro Bertolino and Luc Wauters. To listen to the programme again, go to the Radio 4 Wild Europe website:www.bbc.co.uk/radio4/science/wildeurope.shtml. You will need a copy of “Real One Player” to listen to the broadcast, downloadable free from the link on the BBC page.

Forestry Commission Publications

Management of grey squirrels web pages:www.forestresearch.gov.uk/greysquirrels

Information about FC publications in general can be found here:www.forestry.gov.uk/website/publications.nsf

Finnegan, L., Edwards, C. & Rochford, J. (2008) Origin of, and conservation units in, the Irish red squirrel ( Sciurus vulgaris ) population. Conservation Genetics 9: 1099-1109.