Morphological diversity of the red squirrel, <Emphasis Type="Italic">Sciurus vulgaris</Emphasis>, in Ireland

The red squirrel in Britain and Ireland has been described as a separate subspecies, Sciurus vulgaris leucourus, based on bleaching of the tail and ear tufts. However, recent investigations in northern England found this light colour confined to one area, probably due to the rapid spread of introduced continental European red squirrels. This study reports the first detailed survey of tail colour and cranial measurements in the Irish red squirrel population to (1) investigate the distribution of the light colour morph in Ireland and (2) determine whether the Irish red squirrel population is morphologically divergent from populations elsewhere in the species range. The light tail colour was found in 57% of individuals and in all regions, although it was most common in the northwest. The mixture of different colour morphs indicates the Irish population is a mixture of different subspecies, including S. vulgaris leucourus, while the cranial measurements suggest the Irish squirrel may be morphologically divergent from populations elsewhere. Combined, these results support previous suggestions that conservation measures seek to maintain the diversity within the Irish red squirrel population.     

A grey future for Europe: Sciurus carolinensis is replacing native red squirrels in Italy

Introduced mammals can cause extinction of native species due to replacement competition, disease, predation or hybridization. We studied the colonization of Piedmont (NW-Italy) by American grey squirrel (Sciurus carolinensis) and its effect on the native red squirrel (Sciurus vulgaris). Presence/absence data (2 × 2 km²), of both species were (re)constructed using questionnaires, literature, existing databases, unpublished information, and direct monitoring with hair-tubes. In 1970 red squirrels were still widespread and greys were restricted to forests near the introduction site. By 1990, grey squirrels had increased their range to 220 km², which coincided with the disappearance of native squirrels from 33 squares inside this range. The invasive species continued its spread occupying an area of 2,016 km² in 2010; within this area red squirrels went extinct in 88 squares. Overall, from 1970 to 2010 red squirrel went extinct in 62 % of 2 × 2 km² (ca. 1,689 km²), and were replaced by grey squirrels. The spread of the alien species was slow in the first 20 years, but doubled in the successive two decades. Nevertheless spread was slower than in Ireland and England. Grey squirrel adapt to climate and habitats in both North and South Europe, causing extinction of the native red squirrel. A EU LIFE co-funded project with the aim to control the grey squirrel in North Italy and recent trade-restrictions and trade-ban are a first step in reducing the risk of grey squirrels invading other countries, but their effectiveness will have to be evaluated.     

Limited diversity associated with duplicated class II MHC-<Emphasis Type="Italic">DRB</Emphasis> genes in the red squirrel population in the United Kingdom compared with continental Europe

The red squirrel (Sciurus vulgaris) population in the United Kingdom has declined over the last century and is now on the UK endangered species list. This is the result of competition from the eastern grey squirrel (S. carolinensis) which was introduced in the 19th century. However, recent evidence suggests that the rate of population decline is enhanced by squirrelpox disease, caused by a viral infection carried asymptomatically by grey squirrels but to which red squirrels are highly susceptible. Population genetic diversity provides some resilience to rapidly evolving or exotic pathogens. There is currently no data on genetic diversity of extant UK squirrel populations with respect to genes involved in disease resistance. Diversity is highest at loci involved in the immune response including genes clustered within the major histocompatibility complex (MHC). Using the class II DRB locus as a marker for diversity across the MHC region we genotyped 110 red squirrels from locations in the UK and continental Europe. Twenty-four Scvu-DRB alleles at two functional loci; Scvu-DRB1 and Scvu-DRB2, were identified. High levels of diversity were identified at both loci in the continental populations. In contrast, no diversity was observed at the Scvu-DRB2 locus in the mainland UK population while a high level of homozygosity was observed at the Scvu-DRB1 locus. The red squirrel population in the UK appears to lack the extensive MHC diversity associated with continental populations, a feature which may have contributed to their rapid decline.     

Conservation of genetic uniqueness in remaining populations of red squirrels (Sciurus vulgaris L.) in the South of England

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.     

Modelling the Expansion of a Grey Squirrel population: Implications for Squirrel Control

A recently discovered population of the North American grey squirrel (Sciurus carolinensis), introduced to Ticino Park, Lombardy (N Italy), is likely to spread into continuous prealpine broadleaf forests of Lombardy and the south of Switzerland. We used Spatially Explicit Population Dynamics Models (SEPMs), successfully used to predict the spread of grey squirrels in England and Piedmont, Italy, to examine the effects of different control scenarios on grey squirrel expansion in a 20000 km² area around Ticino Park. Without control, grey squirrels will invade Switzerland within the next two decades, and, concomitantly, the size and distribution of local populations of native red squirrels will be reduced. Simulating different grey squirrel control or removal scenarios suggests that: (i) efficient control is possible and mainly determined by the spatial distribution and woodland size of the ‘target’ control areas; and (ii) immediate actions must be taken, since delay in grey squirrel control will result in the population increasing and spreading, which makes the problems of successful containment more difficult. Model scenarios were based on surveys that may underestimate the real distribution range and current population size of grey squirrels. Therefore, a combination of hair–tube monitoring and a public participation survey to detect grey squirrel presence, which may also help to increase public awareness, is recommended. Successful containment of further grey squirrel spread will require local co-operation between Italian and Swiss authorities involved in wildlife management.     

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?

The control of grey squirrels (Sciurus carolinensis) is widely undertaken as a conservation measure to protect red squirrel (Sciurus vulgaris) populations in the UK. However, inconsistencies and omissions in data collection, as well as fluctuating financial resourcing of control efforts, have meant that it has to date proved difficult to quantify the impact of any regional control initiative upon populations. Here we have scrutinized a 13 years period (1998–2010) within an ongoing grey squirrel control project that reflects the resource challenges typically faced by red squirrel conservation programmes. We present evidence that despite variation in grey squirrel control intensity, the abundance of grey squirrels ultimately decreased significantly. Trapping success was significantly higher in spring and summer months and a greater abundance of grey squirrels was found in deciduous woodland and hazel dominated scrub relative to other habitats; two findings that reinforce existing guidance within national control best practice. Grey squirrels carry an infection that causes epidemic pathogenic disease if spread to the native red squirrel. We observed that the proportion of seropositive grey squirrels decreased constantly from 2003 to 2010 when only 4 % of sampled animals were seropositive. This discovery indicates that culling can in parallel remove both the competitive and disease threat posed to red by grey squirrels. The historical paucity of scientific data on the effectiveness of grey squirrel control as a tactic in UK red squirrel conservation means that the findings of this study will significantly advance conservation best practice and inform the development of future national strategy.     

The Mechanism of Replacement of Red Squirrels by Grey Squirrels: A Test of the Interference Competition Hypothesis

Introduced American grey squirrels have replaced native red squirrels in most of the range currently occupied in Britain and northern Italy. The mechanisms of the replacement are not yet fully understood. We restated the commonly cited Interference Competition Hypothesis (ICH) that grey squirrels interfere with the behaviour of red squirrels in three possible ways: 1. by direct aggressive interactions; 2. by interrupting red squirrel mating-chases; or 3. by forcing red squirrels to actively avoid areas intensively used by grey squirrels. We compared the activity pattern, behaviour and reproductive performance of red squirrels in two study areas in northern Italy, one with only red squirrels (control area C1), the other with both species (experimental area E1). The following predictions were tested: 1. the total time spent in both intraspecific and interspecific interactions by red squirrels increases in the experimental area; 2. most interspecific interactions are aggressive, with grey squirrels being the dominant species; 3. the proportion of breeding female red squirrels that are unsuccessful at weaning offspring increases in area E1; 4. grey squirrels take part and interfere with red squirrel mating-chases, and thereby decrease the reproductive output of red squirrel females; 5. the activity pattern of red squirrels in the mixed-species area is shifted with respect to that in the control area to the hours of the day during which grey squirrels show little activity; and 6. red squirrels will shift their home range (or at least their core-area) when grey squirrel densities increase to avoid interspecific core-area overlap. Our results supported only the first prediction of the ICH: they failed to support all the other predictions. Moreover, the increase in the percentage of active time red squirrels spent interacting with other squirrels in the experimental study area was very small (only 1–2 min/day). Red squirrels did not avoid the woodland patches most intensively used by grey squirrels and the interspecific core-area overlap was similar to red squirrel intraspecific core-area overlap. This suggested that red squirrels avoided spatial overlap with grey squirrels in a similar manner as with conspecifics and that an increase in grey squirrel numbers will augment the intensity of resource competition. We therefore conclude that our results do not lend support to the Interference Competition Hypothesis and that interference competition by grey squirrels cannot explain the large-scale replacement of red by grey squirrels that has occurred in Britain and in Piedmont.     

Genetic Structure of the French Red Squirrel Populations: Implication for Conservation

The decline of the red squirrel (Sciurus vulgaris) in several European countries due to the introduction of the American grey squirrel (S. carolinensis) and the predicted arrival of the grey squirrel in France in the near future has lead to the development of a preventative conservation project in this country. In this study, we conducted an extensive survey of mitochondrial DNA variation in French red squirrels using a fragment of the mitochondrial D-loop and we compared the results with previously published data from other European populations. Our main aims were: (1) to determine whether genetically differentiated populations, which could represent prioritized units for conservation purposes, were present in France and (2) to determine whether the French population, which is currently largely undisturbed, could provide information on the postglacial recolonization history of the species. We found that French D-loop haplotypes show almost no tendency to cluster by geographic origin, be it region or country, suggesting that French red squirrels have not been isolated from other populations during an evolutionarily significant period and that they do not constitute an Evolutionary Significant Unit. The French red squirrels showed strong signals of population expansion, the opposite to what is observed in most other European populations, making them of particular interest to study the postglacial expansion history of the species.     

Grauhörnchen vs. Eichhörnchen

Competition for resources in Europe's forests – grey versus red squirrels As a result of the introduction of the north American grey squirrel in Britain, Ireland and Italy in the 19th and 20th century, the sole native tree squirrel species in Europe – the red squirrel – faced a completely novel competitor. In order to understand the interactions between the two species and to develop a conservation strategy, the two species' biology and ecology was the focus of intense research over the last 25 years. We provide an overview of research findings and the complex competitive interactions for resources and the curcial role of disease in the replacement of native red by introduced grey squirrels. Key factors in the competitive replacement process are habitat composition and patterns of seed food availability, which directly influence individual body condition and reproductive success. In addition, in the British Isles a squirrelpox virus for which grey squirrels act as a reservoir and vector has a drastic impact on the competitive interactions between the two species.     

Genetic diversity and population divergence in fragmented habitats: Conservation of Idaho ground squirrels

The Idaho ground squirrel, which consists of a northern (Spermophilus brunneus brunneus) and a southern subspecies (S. b. endemicus), has suffered from habitat loss and fragmentation, resulting in a reduction in both numbers and geographic range of the species. The northern Idaho ground squirrel (NIDGS) is listed as a threatened subspecies under the Endangered Species Act, and the southern Idaho ground squirrel (SIDGS) is a candidate. Because Idaho ground squirrel populations are small and often isolated, they are susceptible to inbreeding and loss of genetic diversity through drift. This research evaluates levels of genetic diversity and patterns of population divergence in both subspecies of Idaho ground squirrels. We hypothesized that NIDGS would exhibit lower genetic diversity and greater population divergence due to a longer period of population isolation relative to most SIDGS populations. Genetic diversity and divergence were quantified using 8 microsatellite loci. Contrary to expectations, SIDGS populations exhibited consistently lower levels of microsatellite diversity. Additionally, NIDGS exhibited only modest divergence among populations, while divergence levels among SIDGS populations were highly varied. Preliminary evaluations of mitochondrial DNA diversity and structure revealed lower diversity in NIDGS and some differences in gene flow that warrant further study. Based on our results, we suggest different management strategies for the two subspecies. Habitat restoration appears to be the most desirable conservation strategy for NIDGS populations. In contrast, low genetic diversity observed in SIDGS may warrant supplementation of isolated populations through translocations or captive breeding to mitigate further loss of genetic variability.     

Making red squirrels more visible: the use of baited visual counts to monitor populations

Understanding changes in the distribution and abundance of the red squirrel Sciurus vulgaris in Europe, as a result of the spread of the introduced North American grey squirrel Sciurus carolinensis, is vital to planning and implementing red squirrel conservation measures. We studied the effectiveness of a variation on a common method of monitoring squirrels, standardised visual counts that made use of a diffuse baiting approach to increase their detectability. Significantly more sightings of squirrels occurred on baited than on unbaited visual transect lines in trials with the assistance of volunteers, and we recommend the use of baits in future studies.     

Effects of Parasitism and Morphology on Squirrelpox Virus Seroprevalence in Grey Squirrels (Sciurus carolinensis)

Invasive species have been cited as major causes of population extinctions in several animal and plant classes worldwide. The North American grey squirrel (Sciurus carolinensis) has a major detrimental effect on native red squirrel (Sciurus vulgaris) populations across Britain and Ireland, in part because it can be a reservoir host for the deadly squirrelpox virus (SQPV). Whilst various researchers have investigated the epizootiology of SQPV disease in grey squirrels and have modelled the consequent effects on red squirrel populations, less work has examined morphological and physiological characteristics that might make individual grey squirrels more susceptible to contracting SQPV. The current study investigated the putative relationships between morphology, parasitism, and SQPV exposure in grey squirrels. We found geographical, sex, and morphological differences in SQPV seroprevalence. In particular, larger animals, those with wide zygomatic arch widths (ZAW), males with large testes, and individuals with concurrent nematode and/or coccidial infections had an increased seroprevalence of SQPV. In addition, males with larger spleens, particularly those with narrow ZAW, were more likely to be exposed to SQPV. Overall these results show that there is variation in SQPV seroprevalence in grey squirrels and that, consequently, certain individual, or populations of, grey squirrels might be more responsible for transmitting SQPV to native red squirrel populations.     

European red squirrel population dynamics driven by squirrelpox at a gray squirrel invasion interface

Infectious disease introduced by non‐native species is increasingly cited as a facilitator of native population declines, but direct evidence may be lacking due to inadequate population and disease prevalence data surrounding an outbreak. Previous indirect evidence and theoretical models support squirrelpox virus (SQPV) as being potentially involved in the decline of red squirrels (Sciurus vulgaris) following the introduction of the non‐native gray squirrel (Sciurus carolinensis) to the United Kingdom. The red squirrel is a major UK conservation concern and understanding its continuing decline is important for any attempt to mitigate the decline. The red squirrel–gray squirrel system is also exemplary of the interplay between infectious disease (apparent competition) and direct competition in driving the replacement of a native by an invasive species. Time series data from Merseyside are presented on squirrel abundance and squirrelpox disease (SQPx) incidence, to determine the effect of the pathogen and the non‐native species on the native red squirrel populations. Analysis indicates that SQPx in red squirrels has a significant negative impact on squirrel densities and their population growth rate (PGR). There is little evidence for a direct gray squirrel impact; only gray squirrel presence (but not density) proved to influence red squirrel density, but not red squirrel PGR. The dynamics of red SQPx cases are largely determined by previous red SQPx cases, although previous infection of local gray squirrels also feature, and thus, SQPV‐infected gray squirrels are identified as potentially initiating outbreaks of SQPx in red squirrels. Retrospective serology indicates that approximately 8% of red squirrels exposed to SQPV may survive infection during an epidemic. This study further highlights the UK red squirrel – gray squirrel system as a classic example of a native species population decline strongly facilitated by infectious disease introduced by a non‐native species. It is therefore paramount that disease prevention and control measures are integral in attempts to conserve red squirrels in the United Kingdom.     

Patterns of genetic diversity in the red squirrel (Sciurus vulgaris L.): Footprints of biogeographic history and artificial introductions

British S. vulgaris are classified as aseparate subspecies, S. v. leucourus, tomainland Europe. While S. vulgaris is notunder threat across most of its Eurasian range,in Britain, Ireland and Italy populations aredeclining, mainly due to the introduction ofthe American grey squirrel (S.carolinensis). In this study, we conducted anextensive survey of mitochondrial DNA variationin British S. vulgaris populations and apreliminary survey of continental Europeanpopulations. Our main aims were to determinethe extent to which any populations of S.vulgaris in Britain are partially or whollythe product of artificial translocation of redsquirrels from continental Europe, and whethercontinental population variation will provideinformation on post-glacial reafforestationpatterns in Europe. We found that the majorityof extant populations of British S.vulgaris are of continental ancestry, manywith a very recent (last 40 years) Scandinavianancestry. The Scandinavian haplotype hasrapidly become the most dominant innortheastern Britain, despite not appearing innorthern English populations until 1966. Thissuggests that these squirrels may have anadaptive advantage in the non-native sprucedominated conifer plantations of northernEngland. Our preliminary examination ofcontinental populations demonstrated that theyare sufficiently differentiated to allow aphylogeographic study of this species.     

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

In Ireland, the UK and Italy, the invasive North American grey squirrel, Sciurus carolinensis, threatens the survival of the Eurasian red squirrel, Sciurus vulgaris, as the effects of competition and disease almost inevitably lead to total replacement of red squirrel populations. However the results of a recent national squirrel survey suggested that the normally invasive grey squirrel had gone into decline in the Irish midlands, which was anecdotally attributed to an increase in European pine marten, Martes martes, range and numbers. This study aimed to quantify changes in squirrel distribution in Ireland and to investigate the role, if any, of the pine marten in red and grey squirrel population dynamics. A distribution survey of the midlands was carried out which confirmed the grey squirrel population has crashed in approximately 9,000 km² of its former range and the red squirrel is common after an absence of up to 30 years. At landscape level, pine marten and red squirrel abundance were positively correlated, whereas a strong negative correlation between pine marten and grey squirrel presence at woodland level was found to exist. Squirrel demographics were determined by means of live trapping programs which confirmed that the red squirrel in the midlands is now in competitive release and the grey squirrel is present at unusually low density. This study provides the first evidence of a regional grey squirrel population crash and suggests that European pine marten abundance may be a critical factor in the American grey squirrel’s success or failure as an invasive species.     

Current and future impacts of nest predation and nest-site competition by invasive eastern grey squirrels Sciurus carolinensis on European birds

1. 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.
2. 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.
3. 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.
4. 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.
5. 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.

     

What future for British red squirrels?

The geographic range of red squirrels contracted sharply in Britain during the 1940s arid 1950s, as increasingly large areas were colonized by the congeneric North American grey squirrel. Red squirrels remain common only on offshore islands, and in the large conifer forests of northern England and Scotland. The initial replacement of red squirrels was in arras dominated by oak woodland, probably because acorn crops are exploited less efficiently by red squirrels than by grey squirrels. Dirt studies have shown that acorns are digested less efficiently by the red squirrel, which occurs in conifers through most of its Eurasian range, than by the introduced grey squirrel, which is primarily a native of deciduous woodland. The red squirrel will probably be replaced in deciduous and mixed woodland throughout mainland Britain, and may eventually persist only in large areas of conifers which arc far from oak trees. The conservation of red squirrels on islands is therefore particularly important for their survival, perhaps making it worthwhile to create new island populations where they do not at present exist.     

Subspecies of the Central American Squirrel Monkey (Saimiri oerstedii) as Units for Conservation

The accurate diagnosis of conservation units now typically includes recognition of genetic diversity and unique evolutionary lineages and is necessary to inform the conservation management of endangered species. We evaluated whether the two currently recognized subspecies of the endangered Central American squirrel monkey (Saimiri oerstedii) in Costa Rica are evolutionarily significant units (ESUs) that should be managed separately in conservation efforts. We used previously published sequences of 50 individuals of Saimiri oerstedii for 880 bp of the mtDNA d-loop and genotypes of 244 individuals for 16 microsatellites and conducted novel analyses to characterize genetic differentiation between subspecies of Saimiri oerstedii. We measured sequence differentiation and inferred an intraspecific molecular phylogeny and a haplotype network, and found consistent results supporting statistically significant divergence and reciprocal monophyly between subspecies. A population aggregation analysis also supported Saimiri oerstedii citrinellus and S. o. oerstedii as diagnosably distinct units. These results confirm previous genetic studies with smaller sample sizes and are consistent with other factors including differences in pelage and morphology and divergence at nuclear markers. Conservation managers should manage these subspecies separately to prevent the loss of genetic diversity via artificially induced outbreeding. High levels of genetic diversity may buffer populations against outside extinction pressures, to which Saimiri oerstedii are vulnerable because of their dwindling habitat and small population size.     

Genetic variation in natural and translocated populations of the endangered Delmarva fox squirrel (Sciurus niger cinereus)

The Delmarva fox squirrel, Sciurus niger cinereus, is a federally listed endangered subspecies whose range has been reduced by 90%. In an attempt to increase both population size and range, translocation sites were established beginning in the 1960's by moving squirrels from the natural range to sites outside the current range. Although translocations have served as the primary component of the DFS recovery program, there has been very little post-release examination of the genetics of the translocation sites. In this study, we developed ten microsatellite loci, screened the three polymorphic loci, and sequenced a 330 bp fragment of the mitochondrial control region in order to assess levels of genetic variation in natural and translocated regions of Delmarva fox squirrels and to compare them to Southeastern fox squirrels (S. n. niger). Although we found low levels of microsatellite polymorphism, there were no differences in heterozygosity between natural and translocated regions, or between Delmarva and Southeastern fox squirrels. We found high levels of polymorphism in the mitochondrial control region. Our patterns of haplotype diversity suggest incomplete lineage sorting of the two subspecies. In general, our data suggest that the current levels of genetic variation in the translocated sites are representative of those found in the natural population, and we encourage the continued use of translocations as a major component of Delmarva fox squirrel recovery. This revised version was published online in July 2006 with corrections to the Cover Date.     

Adenovirus: an emerging factor in red squirrel Sciurus vulgaris conservation

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