Summary1. Throughout much of Britain, Ireland and north Italy, red squirrels ( Sciurus vulgaris L.) have been replaced by alien grey squirrels ( S. carolinensis Gmelin) introduced from North America. We have studied squirrels in two mixed woodlands in north Italy and two conifer forests in north England. In each country, one site was occupied by red squirrels and one site by both species. 2. We have previously considered interference competition and exploitation competition for food and space between red and grey squirrels and have showed that grey squirrels caused reduced body growth in juvenile and subadult red squirrels, and compete for tree seeds cached by adult red squirrels in spring. Here we report on the effects of grey squirrels on three fitness components in red squirrels that have consequence at the population level: fecundity, residency and recruitment. 3. Litter production peaked in the spring and summer, but fewer females bred in the summer with grey squirrels present. In addition, fewer individual red squirrel females produced two litters per year in the sites with grey squirrels. Moreover, red squirrel recruitment rate and, in the mixed broadleaf sites, red squirrel juvenile residency, decreased with increasing grey squirrel density. 4. Fecundity of individual female red squirrels was lower in red-grey than in red-only sites because they had a lower body mass in sites with grey squirrels. 5. Overall, there was no significant effect of grey squirrels on residency of adult red squirrels or on population turnover rate. However, the presence of grey squirrels resulted in a reduction in red squirrel fitness which was evident by lower population summer breeding and a lower recruitment. Over time, this will result in a decline in population size and eventually population extinction.

The disease implications of novel pathogens need to be considered when investigating the ecological impact of species translocations on native fauna. Traditional explanations based on competition or predation may often not be the whole story. Evidence suggests that an emerging infectious disease, caused by a parapoxvirus, may be a significant component of the impact that the introduced grey squirrel has had on UK red squirrel populations. Here we validate the potential role of parapoxvirus by proving that the virus is highly pathogenic in the red squirrel while having no detectable effect on grey squirrel health.

Red squirrels are declining in the United Kingdom. Competition from, and squirrel poxvirus (SQPV) disease carried by, grey squirrels are assumed to be determining the decline. We analyse the incidence of disease and changes in distribution of the two species in Cumbria, from 1993 to 2003 and compare these to the predictions of an individual-based (IB) spatially explicit disease model simulating the dynamics of both squirrel species and SQPV in the landscape. Grey squirrels increased whilst red squirrels declined over 10 years. The incidence of disease in red squirrels was related to the time since grey squirrels arrived in the landscape. Analysis of rates of decline in red squirrel populations in other areas showed that declines are 17-25 times higher in regions where SQPV is present in grey squirrel populations than in those where it is not. The IB model predicted spatial overlap of 3-4 years between the species that was also observed in the field. The model predictions matched the observed data best when contact rates and rates of infection between the two species were low. The model predicted that a grey squirrel population control of>60% effective kill was needed to stop the decline in red squirrel populations in Cumbria.

Summary1. A stochastic individual-based model for simulating the dynamics of an infectious disease in sympatric red and grey squirrel populations is described. The model simulates the spread of parapoxvirus between squirrels in fragmented populations based on the dispersal of infected animals, the probability of encounters between individuals, exposure to the virus and subsequent mortality. 2. The disease model was integrated with a spatially explicit population dynamics model that simulated red and grey squirrel populations in real landscapes, using habitat information held in a geographical information system. Latin hypercube sampling was used to create a range of realistic life-history and infection scenarios and the model was used to investigate the dynamics of red and grey squirrels in Norfolk between 1966 and 1980. 3. The model predicted that parapoxvirus, like interspeci®c competition, could have led to the extinction of the red squirrel in Norfolk. The results suggest that the red squirrel±grey squirrel±parapoxvirus interaction represents a system of apparent competition mediated by an infectious agent, as seen in other interactions between resident and exotic species. 4. The need for further epidemiological research on the virus is emphasized. We believe that the combined eects on disease transmission of habitat, behaviour and grey squirrels acting as reservoir hosts will lead to a patchy prevalence and sporadic incidence of parapoxvirus disease in red squirrels and a more rapid local replacement by grey squirrels. 5. These results have implications for conservation management of the red squirrel in the UK. Schemes in which animals are translocated or given supplementary feeding may enhance disease spread by bringing infected animals into contact with others.

No abstract

In heterogeneous environments, differential niche selection by two competing species will result in niche partitioning so that individuals of each species can maximise their fitness under different sets of environmental variables. Thus, niche partitioning is considered essential to allow co-existence of ecologically related species. To assess whether niche partitioning was occurring between native red squirrels and alien grey squirrels living together in a 13-ha high-quality mixed deciduous woodland in north Italy, we investigated temporal and spatial patterns in their activity and foraging behaviour between 1996 and 1998. We used live trapping and radio-tracking to study numbers, distribution and behaviour of squirrels. Daily and seasonal temporal activity patterns, and activity on the ground and in the trees, were similar in the two species. However, grey squirrels were more tree specialists and had a narrower tree-species niche width than red squirrels, in particular making greater use of oak. Other studies of red and grey squirrels in allopatry show that the two species differ in the extent they utilise oak. Overall, tree-species niche overlap was about 70%. Grey squirrels had larger home ranges than red squirrels. Home ranges and core areas of both species were larger in males than females. Also, intraspecific home range and core-area overlap patterns were similar to those found in allopatric populations of these species. Overall, there was no evidence that the use of space of one species was affected by the other. Our results show that there was no niche partitioning of activity or foraging behaviour in time or space during the study. This suggests that, at moderate grey-squirrel densities, red squirrels are unable to avoid competition with grey squirrels, and that competition for food and/or space will occur when these resources become limiting

This is a literature review of the relationship between the abundance of tree seeds and the productivity, survival and dispersal of Holarctic tree squirrels. Causal links between changes in tree seed supplies and changes in squirrel numbers are often masked by various factors including intraspecifk and interspecific competition for seed, weather, predation and seed hoarding; density‐dependent relationships have not been clearly established. Tamiasciurus and Sciurus species have different seed‐hoarding strategies which are associated with differences in their social and spatial organization. Average squirrel densities in the long‐term are discussed in relation to forest habitat type, and directions for future research indicated.

The squirrel poxvirus (SQPV) is the probable mediator of apparent competition between the introduced invading gray squirrel (Sciurus carolinensis) and the red squirrel (Sciurus vulgaris) in the UK, and modeling studies have shown that this viral disease has had a significant impact on the decline of the red squirrel in the UK. However, given our limited understanding of the epidemiology of the disease, and more generally the effects of invasive species on parasite ecology, there is a need to investigate the transmission dynamics and the relative pathogenicity of the virus between species. We aimed to increase our knowledge of these processes through an empirical study in which we: (i) used pathological signs and transmission electron microscopy (TEM) to diagnose SQPV disease in red squirrels found dead during scanning surveillance between 1993 and 2005; (ii) detected antibody to SQPV using an enzyme-linked immunosorbent assay (ELISA) in the same animals; and (iii) mapped cases of the disease, and the gray squirrel distribution, using a geographical information system. We analyzed the distribution of cases of SQPV disease according to woodland type, a measure of squirrel density. SQPV disease occurred only in areas of England also inhabited by seropositive gray squirrels, and as the geographical range of gray squirrels expanded, SQPV disease occurred in these new gray squirrel habitats, supporting a role for the gray squirrel as a reservoir host of the virus. There was a delay between the establishment of invading gray squirrels and cases of the disease in red squirrels which implies gray squirrels must reach a threshold number or density before the virus is transmitted to red squirrels. The spatial and temporal trend in SQPV disease outbreaks suggested that SQPV disease will have a significant effect on Scottish populations of red squirrels within 25 years. The even spread of cases of disease across months suggested a direct rather than vector-borne transmission route is more likely. Eight juvenile and sub-adult free-living red squirrels apparently survived exposure to SQPV by mounting an immune response, the first evidence of immunity to SQPV in free-living red squirrels, which possibly suggests a changing host-parasite relationship and that the use of a vaccine may be an effective management tool to protect remnant red squirrel populations.