A battle of wits? Problem-solving abilities in invasive Eastern grey squirrels and native Eurasian red squirrels

This is the author accepted manuscript. The final version is available from Elsevier via the DOI in this recordBehavioural 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.This project is supported by the Postgraduate Research Enhancement Fund from University of Exeter and Gilbert’s private fund

Mixture or mosaic? Genetic patterns in UK grey squirrels support a human-mediated ‘long-jump’ invasion mechanism

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

Squirrelpox in a red squirrel in Fife

SQUIRRELPOX has been identified as a key factor in red squirrel (Sciurus vulgaris) decline in the UK.1 Grey squirrels (Sciurus carolinensis) are thought to act as reservoir hosts for squirrelpox virus (SQPV), the causative agent of squirrelpox, with a reported asymptomatic seroprevalence of 61 per cent.2SQPV is implicated in the complete replacement of red squirrels by grey squirrels throughout mainland England and Wales,1 and poses a major threat to Scottish red squirrel populations since being first detected in 2007.3 To combat this threat, an mortality surveillance programme has been established at the Royal (Dick) School of Veterinary Studies, University of Edinburgh, using opportunistic sampling of red squirrel carcases. This has been ongoing for several years, including a summary publication covering 262 cases submitted from 2005 to 2009.4As part of this monitoring programme, an adult female red squirrel carcase was submitted by a member of the public, after having been found in Townhall Wood (NT110894), on the outskirts of Dunfermline, Fife, in March 2024. Postmortem examination identified multiple lesions typically associated with squirrelpox, including ulcerative and exudative dermatitis of the periocular and perioral skin (Fig 1). Histopathology of the affected skin identified extensive ulceration alongside remnant areas of epithelium with marked ballooning degeneration and eosinophilic intracytoplasmic inclusion bodies. Transmission electron microscopy of the affected tissue also identified numerous pox virions within the affected tissue, which through their size, shape and available surface morphology (Fig 2), were consistent with those of SQPV. In Britain there have been no additional identified diseases in red squirrels that present with periocular or perioral, ulcerative to exudative dermatitis due to a poxvirus,5indicating this case is highly likely due to SQPV.This finding represents the first identification of squirrelpox north of the central belt and is consistent with the predictions of previous modelling, which identified a high risk of northern SQPV spread from 2023 onwards.6 This modelling also suggests a rapid increase and spread of squirrelpox into more northerly and naive red squirrel populations is likely following establishment north of the central belt in central Scotland.6This case and modelling supports an increased requirement for targeted investigations, ongoing monitoring and grey squirrel interventions both around Dunfermline itself and within adjacent areas to establish the disease burden in this locality and limit further northerly squirrelpox spread.LA Wilson, veterinary pathology lecturer, M Marr, postdoctoral research fellow, C Logie, postmortem room technician, K Beckmann, conservation medicine lecturer, PWW Lurz, squirrel ecologist, R Ogden, director of conservation science, E Milne, professor emerita of veterinary clinical pathology Royal (Dick) School of Veterinary Studies and Roslin Institute, University of Edinburgh, Easter Bush Campus, Roslin, Midlothian EH25 9RG email: [email protected] DJ Everest, pathology scientist APHA Weybridge, Addlestone, Surrey KT15 3NBReferences1 Tompkins DM, White AR, Boots M. Ecological replacement of native red squirrels by invasive greys driven by disease. Ecol Lett2003;6:189–962 Sainsbury AW, Nettleton P, Gilray J, et al. Grey squirrels have high seroprevalence to a parapoxvirus associated with deaths in red squirrels. Anim Conserv2000;3:229–33 3 Mclnnes CJ, Coulter L, Dagleish MP, et al. First cases of squirrelpox in red squirrels (Sciurus vulgaris) in Scotland. Vet Rec2009;164:528–314 LaRose JP, Meredith AL, Everest DJ, et al. Epidemiological and postmortem findings in 262 red squirrels (Sciurus vulgaris) in Scotland, 2005 to 2009. Vet Rec2010;167:297–3025 Everest DJ, Tolhurst-Cherriman DAR, Davies H, et al. Assessing a potential non-invasive method for viral diagnostic purposes in European squirrels. Hystrix 2019;30:44–506 White A, Lurz PWW. A modelling assessment of control strategies to prevent/reduce squirrelpox spread. 2014. Scottish Natural Heritage Commissioned Report No 627. https://bit.ly/441TOgM (accessed 8 April 2024)PROFESSIONHistory of the veterinary professionI WRITE in response to the debate article by Bruce Vivash Jones (VR, 16/23 March 2024, vol 194, p 236). As a PhD researcher on the history of the RCVS and veterinary regulation, and a veterinary nurse, I would like to raise some issues with Vivash Jones’ historical evidence. He states that changes to the council structure would create an oligarchy. From his interpretation of an oligarchy I can assure him and our profession that the first RCVS council did work as This finding represents the first identification of squirrelpox north of the central belt20/27 April 2024 | VET RECORD312Fig 2: Squirrelpox virus virions detected in the affected tissue. Bar = 200 nmFig 1: Macroscopic lesions of ulcerative and exudative dermatitis surrounding the eye in a red squirrel (Sciurus vulgaris)Letters 20 April.indd 312Letters 20 April.indd 31216/04/2024 12:4816/04/2024 12:4

Do founder size, genetic diversity and structure influence rates of expansion of North American grey squirrels in Europe?

Aim This study investigates how founder size may affect local genetic diversity and spatial genetic structure of the invasive American eastern grey squirrel (Sciurus carolinensis) in European areas. It also examines whether dispersal propensity and invasion rate may be related to founder size, genetic diversity and structure. Location Piedmont, Italy; Northern Ireland, Northumberland and East Anglia, UK. Methods Across the invaded range in Europe, 315 squirrels from 14 locations, grouped in four areas, were sampled and examined at 12 highly polymorphic microsatellite loci. We estimated both genetic variation and population structure using AMOVA, Mantel tests and Bayesian analysis. We also estimated migration rates and range expansion rates. Results Genetic diversity varied in accordance with numbers of founders across populations. For instance, the Italian population had the smallest founder size and lowest genetic variability, whereas Northumberland had high values for both. Significant levels of genetic differentiation were observed in all the examined regions. Gene flow, migration and population range expansion rate were also higher in England and Ireland than in Italy. Main conclusions Populations descending from human-mediated releases of few individuals were more genetically depauperate and more differentiated than populations established from a greater number of founders. Propagule pressure is therefore a significant factor in squirrel invasions. There is a trend whereby larger founder sizes were associated with greater genetic diversity, more dispersal, less local genetic differentiation and faster range expansion rate in squirrels. These findings have important management implications for controlling spread rate of squirrels and other invasive species: good practice should prioritize preventing further releases and the merging of genetically distinct populations as these events can augment genetic diversity

Implications of squirrelpox virus for successful red squirrel translocations within mainland UK

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

Impact of piglet oral vaccination against tuberculosis in endemic free-ranging wild boar populations

This is the author accepted manuscript. The final version is available from Elsevier via the DOI in this recordThe Eurasian wild boar (Sus scrofa) is the main wild reservoir of the Mycobacterium tuberculosis complex in Mediterranean woodlands and a key risk factor for cattle tuberculosis (TB) breakdowns. Wild boar vaccination therefore has the potential to be a valuable tool for TB control. We tested two orally delivered vaccines, heat-inactivated Mycobacterium bovis (IV) and BCG, in four sites (two per vaccine type: one Managed and one Natural or unmanaged) during four years. TB was also monitored in 15 unvaccinated sites (spatial control), as well as in all sites from one year prior to intervention (temporal control). The rationale is that by vaccinating 2-6 month old wild boar piglets we can reduce disease at the population level during the study period. This is achievable due to the fast turnover of wild boar populations. Vaccine baits were deployed using selective piglet feeders and this method proved highly successful with uptake rates of 50 to 74% in Natural sites and 89 to 92% in Managed sites. This is relevant for the potential delivery of vaccines to control other diseases, too. Local wild boar TB prevalence at the beginning of the study was already high ranging from 50 to 100%. TB prevalence increased in unvaccinated sites (6%), while a significant decline occurred in the Managed IV site (34%). Changes recorded in the remaining sites were not significant. The short-term impact of vaccination observed in the field was complemented by mathematical modelling, representative of the field system, which examined the long-term impact and showed that vaccination of piglets reduced prevalence and increased abundance at the population level. We conclude that IV could become part of integrated TB control schemes, although its application must be tailored for each specific site.Engineering and Physical Sciences Research Council (EPSRC)Scottish Funding CouncilHeriot-Watt UniversityUniversity of EdinburghMINEC

Grey squirrels in central Italy: a new threat for endemic red squirrel subspecies

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