Scottish Wildcat Action (SWA) - Final Summary Report 2023

Published: March 2023

This report should be cited as: Campbell R. D., Gaywood M.J., & Kitchener A.C. (Eds.) 2023. Scottish Wildcat Action: Final Summary Report. NatureScot, Inverness.

For more information, and to download this and associated technical reports, go to: Scottish Wildcat Action (SWA) - Specialist Reports 2023.

Key partners and funders

Other supporters and funders are listed in the acknowledgements at the end. 

LIST OF CONTRIBUTORS

The text of this document, in particular the summaries of the individual projects set out in chapter 2, were provided by the following contributors (note that the contact details describe the affiliations of the personnel at the time of SWA):

• Beatriz S. G. Alves. The Royal (Dick) School of Veterinary Studies (R(D)SVS), University of Edinburgh, Easter Bush Campus, Midlothian, EH25 9RG
• Neil E. Anderson. The Royal (Dick) School of Veterinary Studies (R(D)SVS) and the Roslin Institute, University of Edinburgh, Easter Bush Campus, Midlothian, EH25 9RG
• Alice Bacon. The Royal (Dick) School of Veterinary Studies (R(D)SVS) and the Roslin Institute, University of Edinburgh, Easter Bush Campus, Midlothian, EH25 9RG
• David Barclay. Royal Zoological Society of Scotland, Highland Wildlife Park, Kincraig, Kingussie PH21 1NL
• Katie M. Beckmann. The Royal (Dick) School of Veterinary Studies (R(D)SVS) and the Roslin Institute, University of Edinburgh, Easter Bush Campus, Midlothian, EH25 9R
• Vicky Burns. Scottish Wildcat Action, Great Glen House, Leachkin Road, Inverness, IV3 8NW
• Roo Campbell. Scottish Wildcat Action, Great Glen House, Leachkin Road, Inverness, IV3 8NW
• Hebe Carus. Scottish Wildcat Action, Great Glen House, Leachkin Road, Inverness, IV3 8NW
• Martin Gaywood. NatureScot, Fodderty Way, Dingwall Business Park, Dingwall, IV15 9XB
• Calum Hislop. Scottish Wildcat Action, Great Glen House, Leachkin Road, Inverness, IV3 8NW
• Kerry Kilshaw. Wildlife Conservation Research Unit, University of Oxford, Recanati-Kaplan Centre, Tubney House, Abingdon Road, Tubney, Oxfordshire, OX13 5QL
• Andrew C Kitchener. Department of Natural Sciences, National Museums Scotland, Chambers Street, Edinburgh, EH1 1JF
• Kenny Kortland. Forestry and Land Scotland, Tower Road, Smithton, Inverness, IV2 7NL
• Keri Langridge. Scottish Wildcat Action, Great Glen House, Leachkin Road, Inverness, IV3 8NW
• David W. Macdonald. Wildlife Conservation Research Unit, University of Oxford, Recanati-Kaplan Centre, Tubney House, Abingdon Road, Tubney, Oxfordshire, OX13 5QL
• Duncan McKenzie. Scottish Wildcat Action, Great Glen House, Leachkin Road, Inverness, IV3 8NW
• Anna L. Meredith. The Royal (Dick) School of Veterinary Studies (R(D)SVS), University of Edinburgh, Easter Bush Campus, Midlothian, EH25 9RG
• Rob Ogden. The Royal (Dick) School of Veterinary Studies (R(D)SVS) and the Roslin Institute, University of Edinburgh, Easter Bush Campus, Midlothian, EH25 9RG
• Emma Rawling. Scottish Wildcat Action, Great Glen House, Leachkin Road, Inverness, IV3 8NW
• Helen Senn. Royal Zoological Society of Scotland, Edinburgh Zoo, 134 Corstorphine Road, Edinburgh, EH12 6TS
• Nicola Tallach. Scottish Wildcat Action, Great Glen House, Leachkin Road, Inverness, IV3 8NW
• Jenny Wemyss. Scottish Wildcat Action & NatureScot, Great Glen House, Leachkin Road, Inverness, IV3 8NW

Note that the views and opinions expressed within this and associated specialist reports may not reflect the views and opinions of all contributors.

SUMMARY

• The wildcat in Scotland is threatened by a combination of low population size, hybridisation with domestic cats, accidental persecution and potentially, road traffic accidents, disease and habitat loss and / or change. 
• The Scottish Wildcat Conservation Action Plan (SWCAP) was produced in 2013 with the input of over 20 partner organisations and was based on earlier work to conserve the wildcat in the Cairngorms. The Scottish Wildcat Action (SWA) project was established to take forward many of the activities identified within the SWCAP.
• The SWA work was centred on three pillars: to work within wildcat geographical ‘Priority Areas’ to reduce the risks of hybridisation, disease and accidental persecution; to establish a conservation breeding programme; and to improve our knowledge of the status and ecology of the wildcat.
• Within the Priority Areas, we worked with over 140 volunteers to detect and trap, for neutering and vaccinating, 205 feral domestic and obvious hybrid cats. We also encouraged the neutering and vaccination of pet cats. We engaged with estates to reduce the risks to wildcats during predator control procedures. We also worked closely with forestry staff to reduce risks to wildcats during forestry operations and to develop ways of improving the forest habitat for wildcats, including testing the use of artificial dens.
• To establish a conservation breeding programme, we genetically screened all fertile captive wildcats in the Scottish Wildcat Studbook. We also screened 13 potential wildcats from the wild outside of the Priority Areas, of which two individuals were taken into the conservation breeding programme. Between August 2015 and October 2019 we increased the captive wildcat breeding population by 67% from 64 to 107 animals, with an average annual offspring survival rate of 85%.
• To improve our knowledge of the wildcat in Scotland, we collected over one million images in camera-trap surveys of wild-living cats, identifying 356 cats within Priority Areas, including 31 wildcats as identified by coat pattern. We also collected 769 cat records from the public, 45 of these we verified as wildcat based on appearance. We fitted GPS collars to 14 wildcat-hybrids, using the data to inform forestry management, identify rest sites and examine habitat use and movement patterns of the cats. We investigated disease and toxins in wild-living cats, identifying a broad range of infectious agents and showing that cats accumulate rodenticides (rodent poisons) in their tissues. We ran genetic tests on 529 cat samples, finding that no samples taken from wild-living cats alive during the project scored as wildcat. We examined the morphology of 118 dead cats, over half of which were killed on our roads, with none proving to be wildcats. Morphology and genetic results indicate high levels of hybridisation in the current wild-living cat population and that the level of hybridisation has increased markedly since about 1995.
• Overall we detected too few cats that looked like wildcats in our Priority Areas for their populations to be sustainable in the short term, and too few nationally outside Priority Areas to be viable in the long term. In 2019 we shared survey and genetic results with members of the IUCN SSC (International Union for the Conservation of Nature, Species Survival Commission) Cat Specialist Group, who concluded in their own independent review that the wild population was no longer viable without reinforcement or reintroduction.

Moving Forward

• Following the recommendations set out in the IUCN Review and drawing on the expertise built during Scottish Wildcat Action, project partners secured funding from EU LIFE and other sources for a major new project called Saving Wildcats (#SWAforLIFE). This new project will establish a dedicated wildcat ‘breeding for release’ centre and aims to release wildcats bred there into the wild. This six-year programme represents the next, but not final, step in restoring the wildcat in Scotland.
• The ultimate goal for wildcat conservation in Scotland must be to establish a long-term, viable population of wildcats that does not require further intervention to secure its viability. Wildcats hybridised because the population was small and fragmented. Conservation interventions are extremely challenging when hybridisation has occurred. We need to remove the threat from hybridisation. This includes continuing efforts to neuter pets, feral domestic cats and obvious hybrids. Changes to the legislation to ensure pets are microchipped and neutered would make a significant advance towards this goal. However, our long-term vision must also include removing the root causes of hybridisation, including improvements to habitats used by wildcats and their prey, and a reduction in mortality from persecution and other causes such as road traffic. Wildcat conservation should be a countrywide undertaking that will require long-term commitments from all stakeholders to succeed.

CHAPTER 1: THE BACKGROUND TO SCOTTISH WILDCAT ACTION

Introduction

The wildcat, Felis silvestris, is one of our most iconic species. Elusive and rarely seen, it is threatened by a combination of extremely challenging factors, in particular hybridisation with domestic cats. Its secretive nature has made it a difficult animal to study in the wild, making it harder to design the most efficient and effective conservation measures.

However, despite these challenges, a unique partnership came together to find out more about wildcats in Scotland, and to see if it was still possible to save the species in the wild. This resulted in the multi-partner Scottish Wildcat Action (SWA) project that ran from 2015-2020. New technologies, such as camera trapping, GPS tracking and genetic tools, provided new insights into the ecology and occurrence of wildcats, feral domestic cats, Felis catus, and their hybrids. This helped to give us a far better idea of the perilous status of the species and the approaches that are now necessary if we are to restore the wildcat in Scotland.

This report summarises the work carried out by SWA. The project recognised the importance of publishing and sharing information, and using scientific, evidence-based, approaches to inform decision-making. Those readers who wish to learn more about the SWA work can download the detailed technical reports produced on the various topics covered, for which links are provided below, as well as various scientific papers and other published outputs.  All of this work has helped to inform the design of the ‘Saving Wildcats’ project, which represents a new phase of action that aims to release wildcats into a prepared Highland site over the coming years until 2025.

Background to SWA

A number of studies took place from the 1980s through to the late 2000s that showed that the wildcat had become progressively restricted to northern Scotland and that hybridisation with domestic cats, and the associated issues surrounding how a wildcat could be defined and identified, was a significant problem. A first major attempt in developing a more strategic and planned approach to wildcat conservation, and carrying out action on the ground, was the Cairngorm Wildcat Project which ran from 2007-2012 as part of the Species Action Framework.

Informed by this work, over 20 partner organisations contributed to the production of the Scottish Wildcat Conservation Action Plan (SWCAP) in 2013. The action plan was ambitious, setting out the need to identify at least five ‘Priority Areas’ for conserving wildcats, to take forward conservation work in the Priority Areas, and to underpin a wider conservation programme (such as the development of conservation breeding).

The SWA project was established to take forward this action plan. It ran from 2015-2020, funded by the Scottish Government, National Lottery Heritage Fund and others. SWA operated with a team of project staff managed by NatureScot, with associated work carried out by partner organisations and a wide volunteer network. Much of the work focussed initially on six Priority Areas, later reduced to five when the first surveys failed to detect any wildcats in the Strathavon Priority Area (see map below). The overall work programme was steered by a group of ten partners, which are listed at the end.

The SWA work was based around these key principles:

• To work with local people in wildcat ‘Priority Areas’ to reduce the risks of hybridisation, disease and accidental persecution;
• To breed healthy wildcats for later release to bolster the population through a conservation breeding programme;
• To gather extensive data and share our findings to improve understanding of this elusive predator.

This summary report, and the accompanying technical reports, describe the substantial amount of information and experience collated during the short life of SWA. Through these efforts we have learned that the wildcat is imperilled more seriously than initially feared. Therefore, we asked the IUCN SSC (International Union for the Conservation of Nature, Species Survival Commission) Cat Specialist Group to carry out an independent review of the work of the SWA, and other continuing wildcat conservation work, to help assess the situation and provide recommendations on the future actions that should follow.

What is a wildcat?

The wildcat is Britain’s only remaining native wild felid. Not much larger than a domestic cat, the wildcat is recognised by its tabby coloured markings and thick blunt ringed tail. Wildcats were once found across Britain but are now only found in Scotland, where they are widely but unevenly distributed throughout the north and central Scottish mainland.  Here, they mainly hunt rabbit and small mammals like voles and mice and require a mixture of cover habitat such as scrub or woodland and more open habitat such as grassland or riparian areas that support prey.

The wildcat is currently classified as Critically Endangered in the Red List for Scotland. Current threats include a combination of hybridisation with domestic cats, accidental persecution and potentially, road traffic accidents, disease, secondary rodenticide poisoning (an issue newly identified by this project) and habitat loss and / or change.  

Hybridisation is the interbreeding of individuals from genetically distinct populations (in this case, the wildcat and the domestic cat) to produce hybrid offspring. Hybridisation can also lead to genetic introgression, which occurs after many generations of hybrids (in this case wildcat × domestic cat hybrids) cross breeding with each other or with one of the parent species. This eventually produces individuals with a complex mixture of genes and morphological characteristics from the parent species, compared to a first generation hybrid, which would have typically half of its genes from each parent. A “hybrid swarm” occurs where most of the individuals in the wild-living population have a mixture of genes and physical characteristics from both original parent species. Hybridisation presents an identification challenge because wildcats are protected legally but hybrids and feral domestic cats are not.

Wildcats were first legally protected under a 1988 amendment to the Wildlife and Countryside Act, 1981. They are listed on Annex IV of the European Directive 92/43/EEC on the conservation of natural habitats of wild fauna and flora as a species in need of strict protection. This is transposed into Scottish law through The Conservation (Natural Habitat, &c.) Regulations, 1994 (amended in Scotland in 2004, 2007 and 2008), where it is listed on Schedule 2 as a ‘European protected species of animal’.  As a result, wildcats were removed from the Wildlife and Countryside Act, 1981 by 2007 amendment to the Conservation (Natural Habitat, &c.) Regulations, 1994.

 

Identifying a wildcat

Recently, wildcats have been identified from their external appearance by a ‘pelage score’ developed by Kitchener et al., (2005). The method gives a score of between 1 (domestic cat trait) and 3 (wildcat trait) for seven key pelage or coat patterns. Individuals that score 19 or more for the resulting pelage score (called the ‘7PS’ score) are classified as a wildcat.

During Scottish Wildcat Action, project partners developed a genetic test (see below) that estimates the proportion of wildcat ancestry (Q) in an individual, from 0 (domestic cat) to 1 (wildcat). The certainty of Q ranges from a lower (LBQ) to upper boundary (UBQ) based on a statistical calculation. The project considers a cat to be a wildcat if LBQ is greater or equal to 0.75, i.e. we have a high certainty that is at least 75% wildcat and therefore it “passes the genetic threshold”. Further details of the test can be found in Senn and Odgen (2015).

These two methods can be used to identify whether an individual is a wildcat or not. However, during the SWA project, the method(s) applied are generally determined by the practicalities of what is possible in the environment in which the activity takes place. For example, it is much easier to have high confidence in the pelage scores from a sedated cat on a well-lit veterinary table, than from a camera-trap image taken at night. Furthermore, genetic and pelage results can only be generated jointly in certain scenarios where a genetic sample can be collected and analysed. Selecting cats to breed from within the conservation breeding programme provides this opportunity, whereas decisions made during the Trap, Neuter, Vaccinate and Return programme do not because genetic results from samples are not returned rapidly enough..  Therefore, each method has a different level of confidence associated with it. For the purposes of the SWA project therefore, we developed a hierarchy of confidence in identification of a wildcat based on the identification method(s) used at the time. In addition, because it was not always possible to score all the pelage characters correctly in certain situations, such as through camera-trap photographs and live-trapped cats in cage traps, we set a conservative pelage score of 17 or more to distinguish wildcats in these field scenarios. The hierarchy of confidence is given below and is the approach adopted for the SWA project.  This approach may be reviewed in the future as new projects will have additional, practical conservation challenges.

An ‘obvious hybrid’ is a cat that fails to reach levels 4 or 5 in the figure above, depending on the experience of the observer. Others terms used to describe cats in this report refer to their behaviour and are not taxonomic definitions. These include ‘pet cat’, which is a domestic cat with an owner; ‘stray cat’, which is a domestic cat without an owner, but reliant on humans as a food source; ‘feral cat’, which is a domestic cat living without reliance on humans; ‘farm cat’, which is a domestic cat living on a farm; and ‘wild-living cat’ which is any cat (wildcat, hybrid or domestic cat) that is living without reliance on humans. Note that hybrid cats can occasionally be pets, strays or farm cats, but are usually wild-living.

CHAPTER 2: THE WORK OF SCOTTISH WILDCAT ACTION

In this section, we summarise the main areas of work that SWA and its partners focussed on between 2015 and 2020. We have divided this into nine main topics:

1. Survey and monitoring – Assessing the numbers of wildcats and other wild-living cats in the Priority Areas and elsewhere in Scotland.
2. Genetics and morphology – The work done to investigate the genetics and morphology (in particular their coat or pelage pattern) of wild-living cats, and how this has been complicated by hybridisation.
    
3. Ecology – In particular the results of studies tracking wildcat hybrids fitted with GPS radio-collars between 2018-2020.
    
4. Disease and health – Investigating infectious and non-infectious diseases in wildcats, hybrids free-living domestic cats and the threats these may pose to wildcat population health.
5. Trap, Neuter, Vaccinate, Return (TNVR) – The main method used to manage the feral domestic cat population in the Priority Areas, and the challenges the team faced.
6. Land management – The issues that relate to wildcats and game management, forestry, farming and other land uses, and ways to reduce risks and increase benefits to wildcats.
7. Conservation breeding – The development of a captive ‘insurance population’ of wildcats with the help of a coordinated network of conservation breeding facilities.
8. Responsible cat ownership – The risks posed by domestic pet cats to the wildcat population, and how simple measures can be used by pet owners and others to make a significant difference to wildcat conservation.
9. Communications – The approaches and methods we have taken to try and highlight the work we and our partners do, and to promote actions that benefit wildcats.

The infographic below provides an overview of the scale of work done over the last five years. Below we have provided brief summaries for all nine of these topics, but for those who would like to know more, we have also produced detailed technical reports that can be downloaded from the SWA Specialist Reports 2023 page. The technical reports provide the details of additional scientific papers and other publications produced so far. However, it is also important to highlight that a lot of this work carries on, and more knowledge and experience will continue to be published and made available over time.

Work to identify the wildcat Priority Areas was done before the SWA started, and you can find out more about this in SNH Commissioned Report number 768.

1. SURVEY AND MONITORING

 

• We collected data on cat populations within the designated SWA Priority Areas (PAs) and more widely outside of the PAs using baited camera-trap surveys, collection of road-casualty cats, live-trapping of cats during Trap, Neuter, Vaccinate, Return (TNVR, for non-wildcats) or for genetic screening and collaring (for putative wildcats), and through the collection of public sightings.
• We conducted standardised camera-trap surveys over winter in Year 1 (2015-2016) for all PAs, excluding Morvern, Year 2 (2016-2017) for Morvern, and repeated winter surveys in Year 3 (2017-2018) for the Angus Glens, Strathbogie, Strathpeffer and Strathspey. During standardised surveys in each PA, volunteers (between 3 and 46, mean 26 per survey), staff and contractors set out between 60 and 142 (mean 82 per survey) baited camera traps for over 60 days each.
• We conducted additional targeted surveys in all PAs at other times, with the main aim of locating cats for trapping.
• Outside the PAs, project partners and affiliates conducted camera-trap surveys at more than 268 camera locations in over thirty sites stretching from Torridon to the Trossachs.
• We used the method developed by Kitchener et al. (2005) to score seven characteristics of the fur, including tail shape and various stripes and spots on different parts of the body.  The resulting pelage score (called the ‘7PS’ score) can distinguish between wild-living cats; cats with a score of 19 or more are wildcats.  However, it is not always possible to be certain of scoring all the pelage characters correctly in certain situations, such as through camera-trap photographs and live-trapped cats in cage traps, so we set a conservative pelage score of 17 or more to distinguish wildcats in these field scenarios.
• Genetic samples provided additional complementary information regarding wildcat identification, but could not be used to direct the immediate decision-making required in the field, hence all population data reported for this survey and monitoring study are based on the pelage scoring and not genetic scoring.
• In total, we invested 50,644 camera-trap days on standardised PA surveys, 35,755 camera-trap days on targeted surveys and over 10,492 camera-trap days on surveys by partners and affiliates outside PAs (total = 96,891), collecting over one million images and videos.
• Images were assessed by Project Staff, and cat images stored and catalogued. Individual cats were visually identified based on appearance, given a unique identification number, and recorded in a spreadsheet for each PA. Individual cats were scored for pelage characters from photos (and sometimes video) by two trained independent observers, and classified as wildcat (pelage score of 17+ and no single character scores of 1), hybrid (pelage score of 10.5-16) or domestic cat (pelage score below 10.5 with no scores of 3). 
• Across the project years (including data collected outside standardised surveys) we detected a total of 31 individual wildcats, 162 individual hybrids and 151 individual domestic cats, excluding known pets (total 356 including 12 un-categorised cats). Overall, hybrid and domestic cats together outnumbered wildcats over 10:1. Only two wildcats were detected outside our PAs during camera-trap surveys.
• During standardised surveys and summed across sites, we detected one individual cat for every 201 camera-trap days and one individual wildcat for every 1,746 camera-trap days. Where we repeated surveys, effort per individual cat dropped by 43%, but effort per wildcat increased by 12%.
• In Strathavon, the first standardised PA survey in Year 1 and a follow-up PA Survey in Year 2, failed to detect any wildcats. In all other areas we detected between 1-5 wildcats during the first standardised PA survey and between 0 and 5 wildcats in subsequent years, with evidence of turnover in the wildcat population during the project.
• Spatially explicit capture-recapture (secr) modelling was run on the standardised survey data only. This modelling requires large numbers of cats to visit cameras more than once to generate robust population size estimates, with sufficient data only available for four of the surveys. Population estimates for all cats are given in table 1 below:

Table 1. Population size estimates for all types of cat (wildcat, hybrid and domestic cat) in some of the PAs that had a sufficient number of individuals and visits using SECR. Upper and lower confidence intervals are also shown.

PA	Survey year	Population size estimate of all cats using secr	Lower	Upper	Actual  number cats detected	Total visits
Angus	1	42	31	69	24	142
Strathbogie	1	31	29	35	27	249
Strathpeffer	1	48	35	74	21	228
Strathpeffer	3	60	48	82	30	88

 

• Overall we detected too few wildcats in our PAs (mean density of 0.01 (± 0.005) wildcats/km²) to be sustainable in the short term without contribution from any wildcats that might exist outside the PAs, and too few wildcats nationally to be viable in the long term.
• In areas where standardised PA surveys were repeated over more than one winter, we detected no change in the wildcat populations (below), except in Strathpeffer where the survey in Year 3 found no wildcats; this was potentially due to significant weather differences with a first very mild winter in Year 1, compared to a colder winter in Year 3, which affected cat movements.

• The robustness of repeated standardised PA surveys was affected to some degree by the conflicting needs of systematic monitoring, maximising detection for management intervention, and baiting/trapping/removing cats during the survey.
• Some individual cats survived the duration of the project, and we were able to identify several individuals detected during research conducted in 2013-2014 prior to SWA’s launch in 2015. However, only one of those cats was confirmed as surviving until the end of this project, with the oldest known individual surviving until at least seven years old.
• Organising camera-trap deployment, camera-trap and volunteer management, data handling, and data analysis took considerable staff time that, during full survey years, severely compromised the amount of time we could invest in TNVR. Balancing the requirement for robust monitoring with conservation actions needs to be considered carefully during future wildcat conservation work. Volunteer management for any field activity requires significant staff time. More efficient methods are needed for the handling of data collected by volunteers.
• We trapped a total of 228 cats alive, including 20 classified as wildcat based on their pelage score, and obtained 13 hair samples and two saliva samples through non-invasive means. We obtained a further 58 samples (mostly hair) from live cats not captured by the project, including five wildcats. We obtained samples from 61 cats killed on the roads.
• Road casualty rates display a seasonal pattern, with significantly more cats killed on roads in autumn and early winter than at other times (below).

• No cats scoring below the pelage threshold of 17 passed the genetic threshold for a wildcat (based on the measure of LBQ = 0.75, which is described further in the Genetics and Morphology section below and the associated technical report) and only five of 73 cats captured further than 50m from buildings tested as a domestic cat. These results suggest that in the PAs almost all wild-living cats are hybrids. The situation appears to be similar elsewhere in northern Scotland (see the Genetics and Morphology technical report for more details). 
• We collected 769 records through the sightings page on the SWA website and the app, of which 317 were described by the observer as a wildcat. We were only able to verify 45 of these as wildcat, with an additional 106 reports verified as ‘plausible’ wildcats. Excluding plausible wildcats, no wildcats were verified north of Lairg (below), indicating either very low numbers or their extirpation from the far north.

2. GENETICS & MORPHOLOGY

• We carried out this work to improve our knowledge of wildcat genetics and morphology, and to investigate the levels of hybridisation in wild populations. In this project ‘morphology’ refers to the study of the form of cats, i.e. how they look and compare, and how this can be measured.
• Various methods have been proposed in the past, in order to distinguish wildcats from domestic cats and their hybrids. However, correlation between different methods has often been poor, so that their reliability is uncertain. We examined skins, skulls, guts and genetics of wild-living cats in Scotland to find the best combination of methods to distinguish wildcats from other wild-living cats.  
• From 2015 to 2019, 118 whole dead cats were collected or made available from a wide variety of sources, including SWA, SNH, zoos and members of the public collecting carcasses, mostly from roads.  These were measured, weighed, prepared as skins and skeletons, and samples were taken for subsequent analyses.

 

• As with the survey and monitoring work used in the field (above), we used the 7PS pelage score to assess fur characteristics, but with wildcats having a score of 19 or more. 
• For road-killed cats, undamaged skulls were given a Skull Character Score (SCT) based on five characters of the skull. SCTs ranged from 5 (domestic cat) to 15 (wildcat).
• A total of 529 samples was available for genetic testing, including 60 historic museum skins, 153 dead wild-living cats from 2015-2019, 72 captive cats and 116 samples from cats that were part of the TNVR management scheme.
• A genetic test was developed, which uses 35 genetic markers (Single Nucleotide Polymorphisms (SNPs)), to give a hybrid score called ‘Q’.  Q estimates the proportion of wildcat ancestry of a wild-living cat, with 0=domestic cat and 1=wildcat. The certainty of Q ranges from a lower to upper boundary (described as LBQ and UBQ) based on a statistical calculation. For genetic management cats are considered to be wildcats if LBQ is greater or equal to 0.75, i.e. “they pass the genetic threshold”. Further details of the test can be found in a report describing the wildcat hybrid scoring protocol for conservation breeding.
• Pelage scores (‘7PS’ as referred to in the section above) correlated very weakly with skull scores (SCT) for wild-living cats collected between 2015 and 2019.  None was identified as a wildcat. This contrasts with wild-living cats collected between 1960 and 1995 in Scotland, where the correlation is much stronger and four were identified as wildcats.
• Genetic results show that hybridisation between wild-living wildcats and domestic cats in Scotland is extensive with all wild-living cats collected between 2015 and 2019 being hybrids or feral domestic cats. None was identified genetically as a wildcat.

• Both morphological and genetic data show that today’s wild-living cats in Scotland show high levels of introgressive hybridisation , with a continuum between domestic cats and the highest scoring hybrids.  This is known as a hybrid swarm. Historical analyses confirm that hybridisation has been occurring throughout the 20^th century, but has increased since around 1995.
• Future conservation management of the wildcat in Scotland relies on the successful release and establishment of captive-bred wildcats with high pelage (7PS) and genetic (LBQ) scores into areas where future hybridisation with domestic cats and hybrids can be minimised.

3. ECOLOGY

• As part of the SWA fieldwork to collect samples for disease screening and genetic testing of wild-living cats within the SWA Priority Areas (PAs), we fitted some of the sampled individuals with GPS radio-tracking collars. This was to improve our understanding of wildcat ecology and behaviour, and how these affect conservation management for wildcats.
• Camera-trap survey work carried out by SWA identified several putative wildcats based on pelage score. Trapping was carried out during February – March 2018, November 2018 - March 2019 and December 2019 - March 2020. Overall, 14 individuals were fitted with GPS collars, across four of the five PAs; Morvern, Strathpeffer, Strathbogie and Angus Glens. Mean pelage score was 18 (range = 17-21).
• All collared individuals had a genetic score (LBQ) less than 0.75, failing to meet the genetic threshold of a wildcat, and indicating they were hybrids. The mean Q score was 0.54 (range = 0.28-0.71. See the Genetics and Morphology section above for more details about what Q scores mean). Although identified genetically as hybrids, collared individuals exhibited behaviour similar to wildcats radio-tracked in mainland Europe and Scotland, suggesting wildcat hybrids may fulfil a similar ecological niche as wildcats today. Therefore, these data are still relevant for wildcat conservation management.
• Collars have lasted, up to the time the data was analysed for this report, an average of 281 days (range = 113 – 333, excluding data from two young males; Brodie, whose collar stopped working/came off after one month and Finley, who we lost track of during COVID-related fieldwork restrictions and for which we only collected 15 days of data to date). Two collars fell off earlier than anticipated and were re-deployed on different individuals. Two males (Bogie and Clash) caught in Feb/March 2018 were re-collared in the 2018/2019 trapping season. Clash was collared for a third time in Jan 2020 and a female (Doll) was also recollared in January 2020. Two individuals (Finley and Woody) were still collared at the time the analysis was carried out.

 

• Mean total home range size (95% Minimum Convex Polygons) was 13.94 km² for females (range = 4.42-28.59 km², N= 4) and 18.15 km² for males (range = 8.32-29.27 km², N=8). Home-range size for females was larger than previously reported and for both sexes, and larger than reported for some mainland European wildcat populations.
• Home-range size varied monthly with significant differences between the sexes in the pattern of variation, though not the overall size. Mean monthly home ranges were largest for males between February and August (12.79 km²) and for females between February and April (11.93 km²). Unlike for males, female home range size decreased from the end of April due to decreased ranging behaviour when rearing kittens (mean monthly home range size from May to October of 3.48 km²).
• Three individuals were collared more than once to examine home range size and use over a longer period. Annual home range size for Bogie (Strathbogie PA) did not appear to vary notably during two consecutive years. In comparison, Clash’s home-range size (Strathbogie PA) decreased in 2020 compared to 2018 and 2019 and also shifted to more open farmland in the third consecutive year, likely in relation to the establishment of a home range by another collared individual, Emma, within part of his home range. Doll’s (Angus Glens PA) home-range size was also notably smaller in 2020 compared to 2019. This could be explained in part by the presence of Finley, a new young male not previously seen in her home range and also, by the increased presence of visitors and wild campers to the area in 2020 as a result of COVID-related travel restrictions.
• Two individuals in the Angus Glens (Markie and Doll) showed some seasonal shift in home range from forested areas in winter to more open heathland and upper grasslands in spring/summer, possibly in relation to prey availability.
• Two females showed evidence of giving birth; Fairburn (Strathpeffer PA) gave birth around the 30^th April/1^st May 2018 in a derelict stone barn with piles of wood leaning against the walls. A second female, Emma (Strathbogie PA), probably gave birth  around the 5^th/6^th May 2019 on the edge of an area of clearfell (an area of plantation forest recently felled for timber extraction). Emma showed a marked change in activity patterns for several weeks after this period. As has been found in other studies, Emma moved kittens to other den sites after a few weeks (below). We think Fairburn lost her kittens after a few weeks.

• Wildcat hybrids exhibited habitat selection within their home ranges. Although habitat use varied by individual cat across the different PAs, overall, collared individuals were significantly more likely to be found close to edge habitats (except for Fairburn, where distance to edge was not included as a predictor variable in the model). Pooling the individual model data showed that wildcat hybrids in general were more likely to be found close to or in arable landscapes (especially areas with rabbits), broadleaved woodland, edge habitats and using low-use roads/forest tracks, but that differences also existed between different PAs. For example, in the Angus Glens wildcat hybrids were significantly likely to be found in or close to coniferous woodland, which was not the case in the other PAs. Here, collared individuals regularly used areas of wind-blown trees within coniferous woodland, forested tracks through the trees or along the edges of coniferous woodland up the glens. In Strathbogie all six collared individuals were often found close to or within farm buildings, or around the edges of rural properties, and showed a significant decrease in probability of occurrence further away from these structures. In Morvern two individuals (Sprat and Stonker) showed a significant decrease in occurrence further away from mixed woodland. In this PA, in particular, there is a relatively substantial amount of mature mixed woodland that is well connected and highly structured, providing excellent cover and prey for cats.

 

• Wildcat hybrids used many different habitats and structures as rest sites, including patches of gorse, hedgerows, wind-blown trees, stone cairns, hay barns, dense vegetation, and woody debris in clear-fell. Across all collared individuals several different rest sites were identified, where cats spent three hours or more. Many of these were used more than once (mean = 40, SD = 33), some of which were very regularly used on an almost daily basis (e.g. a favourite hay barn), while the remainder were only used once (mean = 75, SD = 45). Rest sites were spread across the individuals’ home ranges, often using several different “locations” within the same general area (e.g., a large area of gorse or farmyard). The number of rest sites increased as the number of days a cat was monitored for increased, suggesting that the number of rest sites are not a limiting factor. Note this may not be the case for natal dens, where areas suitable for rearing kittens could be limited.

• The wildcat is a European protected species within the Conservation (Natural Habitats, &c.) Regulations 1994 (as amended). Under this legislation it is an offence to deliberately or recklessly disturb a wildcat in, obstruct access to, damage or destroy a den or any other structure or place it uses for shelter or protection. It is also an offence to disturb a wildcat, while it is rearing or otherwise caring for its young. Current advice for developers and land managers is to look for dens, or other structures used by wildcats for resting, by initial walking over the site, then using camera traps or genetic testing of samples, such as hair or scats, at any potential dens or resting sites to identify whether a wildcat is using them or not. However, experience from this and other studies shows that identifying these sites is often difficult, even with collared individuals, particularly if the individual is using an area of wind-thrown trees or a large patch of gorse, which is difficult to access.  In addition, the large number of locations used means that it is likely that some are being missed by the current survey methods. Therefore, forestry guidance is currently being reviewed and will be updated taking into account this latest research and we recommend that current advice to developers and land managers is also revised.
• Regular use of farm structures by wildcat hybrids highlights the importance of neutering farm cats to reduce hybridisation, and to minimise the risk of potential secondary rodenticide poisoning through the consumption of rodents found in many farm buildings.
• Some individuals used heathland/grouse moorlands. Site visits to these areas indicated they were not intensively managed and consequently had relatively high vegetation heights and a higher diversity of plants, providing cover and potentially supporting more prey species (e.g., mountain hare, water voles, field voles and ground-nesting birds). However, wildcats on grouse moorlands are at risk of accidental persecution as a result of difficulties in wildcat identification.  Therefore, continued work with the game-keeping industry to improve identification techniques and increase wildcat-friendly practices is essential.
• Roads are a major cause of mortality for wildcats and the use of roads by wildcat hybrids here highlights an area of concern for wildcat populations in Scotland. Initiatives to identify areas of potential high risk for wildcats in Scotland should be undertaken to try and reduce road-related wildcat mortality.

4. DISEASE AND HEALTH

• We consider ‘significant’ diseases of wildcats to be those with the capacity to compromise the sustainability of the wild-living population and its ability to withstand change. Interactions between domestic cats and wildcats provide opportunities for infectious disease transmission. Prior to the start of SWA there was evidence that wild-living wildcats in Scotland have been exposed to a number of infectious agents that are well known – and can cause severe disease and mortality – in domestic cats.
• Infectious disease has negatively affected some endangered wild felid populations elsewhere in the world and has the potential to impact the wildcat, especially in the face of other, simultaneous threats, such as habitat loss, population fragmentation and/or food scarcity. Therefore, infectious disease threats to wildcats merit further investigation, as does the extent to which wildcats are exposed to environmental toxins, which is largely unknown.
• The aim of this study was to increase our knowledge base relating to disease threats to wildcats in Scotland, more specifically infectious agents and rodenticides (rodent poisons), and to explore relationships between exposure to these agents and potential risk factors, such as hybridisation status.
• From 2015-2020 a range of samples was collected from wild-living cats, including feral domestic cats, and cats of wildcat or hybrid, appearance (based on the pelage). The opportunities to access these cats were through the TNVR (Trap, Neuter, Vaccinate, Return) scheme for feral domestic cats and hybrid cats with low-scoring pelage, and through the targeted trapping of live presumed wildcats and hybrid cats with high-scoring pelage. Samples taken included blood samples, swabs from the mouth, inner eyelids and rectum, and faeces, to detect evidence of infection with the following feline infectious agents: feline immunodeficiency virus (FIV), feline leukaemia virus (FeLV), feline calicivirus (FCV), feline herpesvirus (FHV), Bordetella bronchiseptica, Chlamydia felis, Mycoplasma felis, bloodborne Mycoplasma species and Tritrichomonas foetus. In addition, post-mortem examinations were carried out on carcases of wild-living cats, particularly those found on roads, which were submitted by wildlife rangers and members of the public. A proportion of these carcases was radiographed for evidence of gun-shot. Other post-mortem samples were taken, where possible, to screen for infectious agents and the detection of rodenticides.
• To date, samples have been assessed from 207 live cats trapped through the TNVR scheme and from 17 live cats with wildcat or high-scoring hybrid pelage trapped through the wildcat-trapping scheme. Results are presented for all the trapped wild-living cats as a whole, since through the course of the project it became evident that there was a continuous spectrum of hybridisation within the sampled population.
• All 11 infectious agents we screened for were detected in the sampled population of live wild-living cats. FIV (antigen) was detected in 10 (7%) of 144 live cats and was significantly more prevalent in trapped individuals at the ‘domestic cat’ end of the genetic hybrid spectrum.
• Eighty-one presumed wild-living cat carcases were also examined and samples taken where feasible. FIV (antigen) was detected in the blood of two (11%) of 18 cat carcases tested, which were both domestic cat-wildcat hybrids based on genetic testing. There were abnormalities typical of collision with road traffic in at least 58% of the 81 carcases, and lead shot was detected in the carcases of six cats, of which all but one had domestic cat pelage. However, four were found to be genetic domestic cat-wildcat hybrids. Rodenticides were detected in 30 (61%) of 49 liver samples and in 13 (27%) of these 49 cases the residues were above a threshold level associated with toxic effects in birds and mammals.

 

• Our findings demonstrate that wild-living cats in Scotland are infected with a broad range of infectious agents and have accumulated rodenticides in their livers. Both infectious agents and rodenticides are potential threats to the health of domestic cat-wildcat hybrids and wildcats. We have also highlighted road traffic collision and shooting as causes of mortality in domestic cat-wildcat hybrids.
• We have expanded the range of infectious agents detected in wild-living cats across the hybrid spectrum in Scotland. To our knowledge, this is the first time FIV has been detected in domestic cat-wildcat hybrids and it is a concern, since like FeLV (which was detected in 3% of cats) in domestic cats, the virus typically persists in an individual long term, predisposing them to other disease conditions and ultimately hastening mortality. An apparent association of FIV with domestic cat genotypes, and its lack of detection in previous wildcat studies in Scotland, suggests that infection is likely to have originated in domestic cats. It is probable that increased interactions between domestic cats, hybrid cats and wildcats in recent decades, as evidenced by increasing levels of introgressive hybridisation, have in parallel led to greater exposure of wildcats and hybrid cats to FIV and other pathogens associated with domestic cats. The ability of these infectious agents to cause disease in wildcats and wildcat-domestic cat hybrids, and their potential to impact population dynamics, are still poorly understood and merit further investigation.
• Similarly, the current impacts of road-traffic collisions and shooting on the wild-living cat population are unknown; to better monitor the threat from shooting, we recommend that radiography is included as part of any future post-mortem surveillance scheme. We also recommend further investigations to explore the impacts of rodenticide exposure on cat health.
• Health monitoring will be an important component of continuing conservation management of wildcats, but both the live-sampling protocols and post-mortem schemes will require modifications to maximise their investigative value.
• The presence of a large feral domestic cat population in Scotland poses a challenge to restoration of the wildcat population, not just from the perspective of hybridisation, but also with respect to wildcats’ exposure to disease-causing infectious agents. In order to combat this, a high proportion of domestic cats should be neutered and vaccinated in and around areas where any wildcats are reintroduced in the future. We believe tighter national regulation of domestic cat ownership must be introduced if the wildcat’s long-term future in Scotland is to be secured at a broad geographical scale.
• We recommend that a detailed disease risk analysis is conducted to inform plans for any future reintroduction of wildcats, and that continued attention is paid to other key issues, such as community engagement and habitat management.

5. TRAP, NEUTER, VACCINATE, RETURN (TNVR)

• Hybridisation with, and disease transmission from, feral domestic cats have been identified as key threats to wildcats in Scotland. Domestic cats are popular companion animals for many people, but the feral (wild-living) domestic cats are legally managed in some situations, where they are considered an invasive non-native pest species. Therefore, the management of feral domestic cat populations can be controversial, with lethal control methods viewed as unacceptable by some parties, and non-lethal methods as unacceptable by others.

 

• SWA adopted a non-lethal approach to cat management known as TNVR. Under this method cats are captured alive in cage traps, neutered, vaccinated, and then released at the point of capture. TNVR was used for three reasons:
  
  (i) It has been shown as effective where used intensively in defined areas similar to the approach proposed by the project.
  
  (ii) There were concerns that territorial perturbations from lethal methods could lead to increased movement and conflict between cats, exacerbating disease transmission.
  
  (iii) It was judged that the use of lethal control could lose significant levels of public support, and the project’s ability to promote and encourage the associated campaign to neuter, vaccinate and chip the pet domestic cat population.
   
• In order to achieve long-term changes in the feral domestic/hybrid cat population, modelling suggested that an annual neutering rate of 75% of the population was needed.   
• Detailed camera-trap surveys during the first winter of the SWA project (2015/16) were used to identify feral domestic cats and obvious hybrid cats to target.  A full TNVR programme was launched in the second winter (2016-2017) of the project. This TNVR programme was managed by a contractor in the Angus Glens PA and by staff and volunteers in the other PAs.
• In most cases, camera traps were set up before live-trapping to identify target cats and to familiarise the cats to the bait. Camera traps were often managed by volunteers, with staff or contractors more heavily involved in the live-trapping. Most effort was focussed on trapping hybrid and feral domestic cats in more remote areas (i.e., those living at greater distance from humans), because these presented a greater potential risk to wildcats. However, farm cat colonies and feral domestic cats closer to communities were also targeted in some areas.
• In all the PAs neutering and vaccinating were conducted by qualified vets. Veterinary contractors were employed in the Morvern PA and local veterinary practices were used in all the other PAs. If no signs of ownership could be found, trapped cats were neutered, ear-tipped, and vaccinated against feline panleucopaenia virus, also known as feline parvovirus (FPLV), as well as feline herpesvirus (FHV) and feline calicivirus (FCV) which are both primary causes of the disease syndrome commonly called ‘cat flu’. Cats were screened for feline leukaemia virus (FeLV) and feline immunodeficiency virus (FIV) using ‘SNAP’ tests. The project policy, following veterinary advice, was to euthanase any cat failing the SNAP test or exhibiting other signs of disease or injury that would adversely affect its welfare. When euthanasia was necessary, it was carried out by a qualified vet and not by the SWA project staff.

 

• Overall, we invested 3,652 trap nights in the live-capture of cats. From the third winter (2017-2018) wildcats were also targeted for genetic screening and GPS-collar attachment. This meant there was significant overlap in trapping for both TNVR and wildcats in the third winter and therefore the effort was not recorded separately.
• In total we captured 228 cats on over 262 occasions, including 205 feral domestic and obvious hybrid cats (captured on at least 239 occasions). Another 12 feral domestic and obvious hybrid cats from other sources were processed for TNVR. Bycatch was rare due to camera-led targeting, but included 13 pet cats, pine martens, foxes, badgers, hedgehogs and buzzards. Trapping was avoided at locations visited regularly by badgers and pine martens. The use of a valerian lure reduced incidences of trapping non-target species.
• The average ‘effort’ was 14 trap nights per capture for all cats. This improved from 16 nights in the first trapping year to 12 nights in the second, before declining to 33-40 nights in the third and fourth year, when the focus was more on capturing wildcats than TNVR.  Ninety-nine (43%) of the captured feral domestic and obvious hybrid cats were farm cats, most of which were from the Strathbogie PA. Excluding farm cats, effort per target cat was 28 trap nights (a low of 19 in the second year and a high of 40 in the fourth year when the genetic screening and collaring of wildcats were the main focus).

• Thirty-nine volunteers contributed an estimated 544 days directly to TNVR. Recruiting volunteers to conduct trapping for TNVR was substantially more difficult than for camera-trap surveys.
• A significant effort was made by SWA staff to ensure that trapped domestic cats were not neutered until there was reasonable certainty that they were unowned animals, and not pet cats.  This was a challenging issue, and project staff and volunteers spent considerable time visiting houses around trapping areas to identify local pets. A ‘Mog Shot’ gallery was placed on the project website and promoted on social media, and posters displayed on community notice boards, to help identify pet cats ahead of trapping. However, there were limited responses from this. This experience has demonstrated the significant resources required to identify individual pet domestic cats and is judged unsustainable for future, long-term domestic cat management work that will be necessary to restore wildcats to Scotland. Therefore, a recommendation is that owners of domestic cats be required to microchip their animals, thereby allowing TNVR workers to be able to instantly determine if any trapped domestic cat is owned or not. 
• Getting permission from landowners to trap on their land was a significant part of the programme. However, not all landowners that allowed us to survey on their ground would allow trapping for TNVR. This was primarily the case for estates that conducted predator-control activity (including for feral domestic cats). Relationships need to be built with estates and with farmers who have cat colonies.  Farmers’ concerns (primarily ‘running out’ of cats for rodent control) could usually be addressed, particularly in discussion with other farmers who have directly observed the benefits.
• The majority of trapped feral domestic cats and hybrids were detected through volunteer camera-trap surveys.  Only a small proportion was reported to the project via the SWA website. Local knowledge was more helpful in identifying farm cat colonies and feral domestic cats around settlements.
• Informal collaboration with staff and volunteers of animal welfare organisations was extremely helpful to the project. For example, SWA teamed up with the SSPCA and others when neutering some of the large farm colonies. These welfare organisations also helped to hold kittens too young to neuter and cats that required genetic testing. In some areas, we coordinated trapping to avoid duplication of effort.
• Discounting farm colonies, the 75% annual neutering rate, based on the number of feral domestic and hybrid cats estimated from camera surveys, was not reached. On two occasions, approximately two-thirds of the population was neutered. Overall the 75% rate proved to be extremely difficult to attain with the resources available to the project. The target was also difficult to measure because surveys concentrated on wild-living cats, but farm cats were also neutered in some areas. Wild-living hybrids can behave similarly to wildcats, and the difficulty in identifying and trapping these was a key issue. However, important experience was gained and, with more resources, the 75% neutering rate can be achievable at local levels, at least for limited periods.  However, at wider geographical scales, and in longer timescales, the restoration of the wildcat through the widespread use of TNVR alone is unlikely to be sustainable. This emphasises the need for considering additional options, potentially including the future use of legislation that would encourage more responsible domestic cat ownership, thereby reducing the risk of the pet domestic cat population acting as a source for the feral domestic cat population.

6. LAND MANAGEMENT

• The management of the living and working landscape of Scotland has played, and will continue to play, an important role in the conservation status of the wildcat. 
• Hybridisation and the presence of feral domestic cats and hybrids in wildcat habitats are recognised as the main threats to the survival of the wildcat in Scotland. However, a combination of other threats linked to the management of the land have contributed to the wildcat’s problems, including habitat loss or fragmentation and the killing of wildcats, particularly prior to their legal protection. 
• Wildcats require a mosaic of suitable habitats with sufficient cover, in which to hunt, shelter and breed. Changes to land use, habitat fragmentation and degradation all pose potential threats to wildcat numbers, in addition to the continuing management activities carried out as part of estate, forestry or agricultural operations. Development in wildcat areas may potentially also affect the species.
• However, past or ongoing land management in some areas may help create habitat suitable for wildcats. For example, the rotational clear-felling of commercial conifer plantations can create a mixture of prey and cover habitat combinations that wildcats use, while traditional farming practices may have allowed wildcats to exist historically in some areas that were otherwise poor habitat.

Game management

• Wildcats are a protected species. Feral domestic cats, a non-native species, are not protected and can be shot legally by gamekeepers during predator control. However, hybridisation with domestic cats means that reliably distinguishing wildcats from feral domestic cats and hybrids is difficult and the accidental shooting of wildcats misidentified by gamekeepers is a recognised threat. Conversely, legitimate control of feral domestic cats by gamekeepers may play a role in mitigating the threat of further hybridisation and exposure to infectious disease, if conducted appropriately.
• A protocol for ‘wildcat-friendly’ predator control (WFPC) was first developed under the Cairngorm Wildcat Project and continued to be promoted during SWA. In essence, the protocol discourages the use of spot-lighting (or lamping) at night for cat control, encouraging instead the use of cage-trapping, so that cats can be more reliably identified and, if identified as a possible wildcat, released. We asked the Moorland Forum to review the WFPC, and an updated version was added to the Moorland Management best practice website.
• During SWA we identified an emerging threat to wildcats through the use of thermal-imaging technology to assist night-shooting. Wildcats cannot be identified reliably using this technology. Therefore, we produced a leaflet that highlighted the risks associated with lamping and thermal-image equipment used in night-time shooting, and distributed it at relevant events and to land management partners in 2018-2019.

• Project staff built relationships with estates within the PAs over the course of the project. We loaned trail cameras to five estates to enable identification of wildcats ahead of any predator control they were planning.
• Scottish Rural Development Programme (SRDP) funding was also available to any estates in the PAs to purchase and maintain their own trail cameras and cage traps. However, there was no successful take-up of this funding option. Following a questionnaire survey of 15 landowners, we identified that lack of awareness of the fund, its low value and the onerousness of scheme requirements were all factors in lack of take-up, but that landowners would consider applying to a revised scheme. Any future scheme would require more staff resources to promote it and review its effectiveness. Changes in land-use constraints may also help; the SRDP option is currently restricted to moorland, whereas wildcats will more often use woodland, rough pasture, linear habitats, etc., outside summer.

 

• Staff and volunteers set up project stands, hosted by project partners, at targeted events such as the Highland Field Sports Fair (Moy), the Scottish Game Fair (Scone), the Black Isle Show and local shows to engage with land managers. We published blogs and articles with land management partners (e.g., SLE’s magazine Land Business, BASCs magazine Shooting and Conservation, and the website of Shooting and Country Sports UK). During 2019, we also ran a regular column in Shooting Scotland magazine. We revamped the wildcat identification cards that had proved popular with gamekeepers during the Cairngorm Wildcat Project and distributed these at events and with partners.

• However, by 2018 we found that direct engagement with land managers over wildcat-friendly predator control measures remained relatively low. Reasons for this included lack of interest from some estates, an underestimate by the project of the staff time and resource needed to engage, build relationships and work with estates, or simply that some estates do not conduct predator control using methods that put wildcats at risk. To help improve engagement, we set up a land management subgroup towards the end of 2018. One task agreed from the subgroup was to have a group of gamekeepers trial the WFPC method to inform its future development and encourage uptake of the protocol. Seven gamekeepers in the Angus Glens were involved and the trial took place over the summer of 2019. Feedback indicated that some gamekeepers may not be confident enough to identify wildcats in traps – a recommendation for future work is to consider the production of a wildcat-identification training video.

Forestry

• We worked closely with Forestry and Land Scotland (FLS) to investigate a range of habitat management measures in commercial forestry areas and to promote wildcat-friendly forestry practice.
   
• FLS and SWA staff produced an awareness leaflet for those working in forests containing wildcats, which was distributed to all relevant forestry contractors and FLS operational staff.

• Surveys of rodent prey were carried out in Clashindarroch Forest in the Strathbogie PA.  The aims of this work were to determine the relative abundance of three common species of rodents in three typical habitats found in commercial plantations; unfelled stands of timber, restocked (i.e. replanted) areas, and clearfells.  Bank voles were most common in unfelled stands of timber and field voles in felled areas.  Wood mice were found in low numbers mainly in felled and unfelled stands.  Felled areas with high numbers of field voles were judged most likely to offer the best foraging areas for wildcats. This has been supported by the results of the WildCRU GPS-tagging study of wildcat hybrids in forest habitats, summarised in the ‘Ecology’ section above and in the associated technical report. Although bank voles are present in high numbers in forests, they occur in habitats that are less suited to wildcat hunting (e.g., unfelled and unthinned Sitka spruce stands).  Longer-term monitoring has now been established through FLS to track annual population fluctuations in field voles, which is thought to be the most important rodent prey species within commercial forests.

• Based on our current knowledge, it would appear that timber production and wildcat conservation may be compatible where plantations consist of a mosaic of habitats suitable for wildcats.  Mature single-species stands of conifers, such as Sitka spruce, have limited value for wildcats and their prey.
• FLS constructed a total of 63 artificial dens of varying design, including 30 in the Angus Glens PA and 33 in the Strathbogie PA. FLS staff and SWA volunteers used camera traps to monitor 17 of the artificial dens for 10,490 camera-trap days.  No wildcat activity was noted in any of the artificial dens, although cats were caught on camera traps passing dens.  Results so far indicate that the provision of artificial dens is not a useful mitigation or conservation action. However, these artificial dens may be used by reintroduced wildcats in the future as they settle into the landscape.

• FLS Environment Rangers worked with SWA field staff and wildcat biologists to develop and improve pre-operational surveys for wildcats, and to enhance mitigation measures to lessen the potential impact of forest operations on wildcats.  Enhanced pre-operational surveys were developed, with increased deployment of camera traps, and new restrictions on felling operations were introduced as a precautionary measure. 
• FLS approaches to pre-operational surveys and mitigation will be reviewed and revised to take into account the results of the WildCRU GPS-tagging project. An overall assessment and comparison of the benefits to wildcats and the implications for the local timber industry will need to be carried out by FLS, in order to judge if such measures are worth rolling out more widely. The Scottish Forestry (formerly FCS) Guidance Note 35d: Forest operations and wildcats in Scotland is currently being reviewed and revised based on our experiences and data collected during the camera-trap surveys and GPS-tagging.

 

Agriculture

• We worked with a placement student to conduct a questionnaire survey of 147 farmers and crofters on the management of their cats and farm pests. The responses were weighted towards smaller farms and crofts and showed there was a low use (20% of responses) of rodenticides. The most commonly used chemicals were ‘second generation rodenticides’ that present the greatest secondary poisoning risk (‘secondary poisoning’ occurs when one animal comes into contact with, or ingests, another animal or plant that has poison in its system). Most farms and crofts did not use professional pest controllers, although 90% took some measures to reduce poisoning risk. The most common non-chemical pest control methods were the use of cats. Cats were not regarded as a predation threat against livestock (e.g., poultry), unlike foxes and crows. Most respondents did not conduct predator control, although 30% used shooting and 11% lamping, the latter a potential threat to wildcats.
• We assessed the levels of rodenticide in the livers of 49 wild-living cats collected on roads through an MSc project (also see the ‘Disease and Health’ section above). 61% showed rodenticide residues, nearly half of which were above the toxic level recognised for other mammals and birds. The most prevalent types detected were second generation rodenticides.
• Following this study, we ran regular columns in Farming Scotland magazine during 2019-2020. These discussed the risks to wildcats from rodenticides and encouraged readers to follow the code set out by the Campaign for Responsible Rodenticide Use (see www.thinkwildlife.org). We also described wildcat use of barns for denning, and habitat management techniques farmers could adopt to benefit wildcats. These included farmers and crofters retaining scrub habitat, such as gorse, particularly on south-facing slopes, and ensuring a diverse connected agricultural landscape through the creation of hedgerows and uncultivated field margins.

Development

• We provided advice and wildcat records to ecological consultants involved in development projects over the course of the project. Information gained from the camera-trap surveys and GPS-collaring study will be used to update the guidance available on the NatureScot website in due course.

General

• We held a ‘Sharing Good Practice’ event in 2017 for consultant ecologists, developers, and foresters. The discussions focused on ecological surveys, developments and forestry; presentations are available for download.

7. CONSERVATION BREEDING

 

• For species whose wild populations are threatened by extinction, like that of the wildcat in Scotland, captive breeding programmes can act as a vital conservation support tool.
• A captive (conservation) breeding programme was identified as a valuable resource in the Scottish Wildcat Conservation Action Plan (SWCAP) (Anon, 2013) under objective 3: “taking forward work to underpin a wider conservation programme”. Two projects were launched as part of a long-term “ex situ” programme to ensure that captive populations of wildcats remained as an integral part of conservation actions.
• Since 2015 the Royal Zoological Society of Scotland (RZSS), an SWA partner, has managed the UK’s Wildcat Captive Breeding Programme, which is part of the wider European Associations of Zoos and Aquaria (EAZA) Felid Taxon Advisory Group (Felid TAG) as a Monitoring Programme (Mon-P).
• In recent years RZSS, working closely with captive holders across the UK and with specialists in Europe, has implemented a range of population management techniques to develop to the captive breeding programme and ensure it has the potential to support future wildcat population reinforcement.   

Assessing genetic status of the captive population

• RZSS took over management of the Scottish Wildcat Breeding Programme in 2015. Between 2015 and 2017 samples were acquired from all fertile captive wildcats held in UK collections that were included in the Scottish Wildcat Studbook. Following genetic screening, using the Wildcat Hybrid Scoring Protocol, results were used to support population management decisions. 

Acquisition of more wildcats of known genetic status to increase the genetic diversity of the captive population.

• RZSS engaged with landowners and gamekeepers from 22 estates outside the PAs to assess sites for potential wildcat presence and to collaborate with landowners in the event of accidental trapping of suspected wildcats. From these estates and other estates outside PAs that approached SWA staff, 13 suspected wildcats were screened to assess hybrid genetic status, pelage characters and disease. From the 13 suspected wildcats only two have been kept and included in the conservation breeding programme.

Update of captive husbandry guidelines

• In 2016 RZSS published Wildcat Captive Husbandry Guidelines and disseminated these to all wildcat holders. The guidelines identified the recommended standards for captive wildcat management, housing, breeding, veterinary care and welfare. Following the publication of these husbandry guidelines, RZSS undertook further work to explore the additional management challenges relating to conservation breeding for reintroduction. This work will be refined further with input from international specialists as a support tool for future conservation breeding for release.

 

• Following breeding programme best-practice guidance from the European Association of Zoos and Aquariums (EAZA) the captive wildcat breeding population increased by 67.2% from 64 animals (24 males and 40 females) to 107 animals (47 males and 60 females) during the period  from August 2015 to October 2019. This population growth was supported by new captive holders joining the breeding programme, which increased the enclosure space for new wildcat breeding pairs.
• Annual breeding success from the five breeding seasons between 2015 and 2019 resulted in an average annual offspring survival rate of 85%. By the end of the SWA project all wildcats in the breeding programme were available for continued breeding following the genetic and pelage assessments carried out in 2017 as detailed in the Wildcat Hybrid Scoring Protocol.
• Genomic analysis was used to reconstruct relatedness between all the wildcats within the captive breeding programme, so that the population could be managed to maximize its genetic diversity and minimise its inbreeding levels going forward. Population management is undertaken using the Single Population Analyses and Record Keeping System (SPARKS) and Pedigree Analyses and Management Software (PMx), following best-practice population management guidance.

8. RESPONSIBLE CAT OWNERSHIP

• We identified cat owners as a key stakeholder in the conservation of the wildcat because domestic cats can potentially transmit disease to, and interbreed with, wildcats. Pet cats can also enter the feral population though a number of routes, leading to added risks to wildcats. Reasons for cats entering the feral domestic cat population are further complicated by the varying statuses of cats along a spectrum ranging from owned house cats to unowned feral cats. For example, some cats are ‘strays’, relying on multiple households for food (indeed, some owned house cats visit multiple households too). Therefore, ownership is difficult to establish and responsibilities of ownership are consequently unclear. With hybridisation occurring within the ‘pet’ cat population too, the situation is even less clear. There is therefore a need to ‘turn off the taps’ of new recruits into the feral cat population through promoting responsible cat ownership, which helps by minimising direct interbreeding with wildcats and reducing disease transmission.
• At the core of our responsible cat ownership strategy was a campaign. The main aim of the campaign was to promote the neutering and vaccinating of pet cats, with the additional aim of encouraging the use of pet ID microchip implants to aid differentiation of owned cats from stray and feral domestic cats when conducting TNVR programmes.
• We identified three key messages to push in the campaign:

     (i) Wildcats in Scotland are really special.

     (ii) Cat owners can take action to protect the wildcats that they love.

     (iii) Happy healthy pet cats are neutered, vaccinated & microchipped.

• We branded the campaign #Supercat and worked with a designer to create a comic-book style campaign that featured a ‘normal’ domestic cat turning into a ‘supercat’ through vaccination, neutering and microchipping. The campaign was formally launched on the 28^th Feb 2017.

• From this core comic graphic, we created other marketing material, including stickers, facemasks, t-shirts, a Facebook filter, posters and a leaflet. The leaflet included an explanation as to why these actions were helpful to wildcats, some key facts about wildcats and ten benefits to neutering, vaccinating and microchipping.
• Stickers and facemasks were made available on our website. There we also included further information on the benefits of the procedures to pet cats and to wildcats, and offered cat owners the option of taking a pledge to make their cat a supercat. 

• We promoted the campaign on social media and our website, e.g., using the #supercat hashtag on Twitter, and asked project partners to promote the campaign too. We also asked high profile figures to help, including wildlife filmmaker Gordon Buchanan and the Scottish Government’s Cabinet Secretary for Environment, Climate Change and Land Reform Roseanna Cunningham MSP, alongside our Local Authority Wildcat Champions. This led to a four- to eight-fold increase in engagement through Twitter (1,600 engagements on Twitter in February 2017 compared with 200-400 per month before the launch), but less so for Facebook. Over the duration of the project, we published 40 tweets on Twitter with #supercat, along with 65 Facebook posts.
• During many media and press interviews and articles we promoted  the key message that cat owners could help us by getting their pets neutered, vaccinated and microchipped.
• We promoted responsible cat ownership at events in and around our PAs. We distributed leaflets around local veterinary practices and worked with partners to distribute them to other locations.
• We offered Supercat vouchers to cover the cost of the procedures for pet cats within our PAs. Over the project duration 47 vouchers were used, some for all three procedures and others for one or two (e.g., just microchipping and/or vaccination for pets that owners had neutered already).
• Through our partners at the University of Edinburgh’s Royal (Dick) School of Veterinary Studies (RDSVS) we conducted a questionnaire survey of cat owners. Out of 1,313 responses, 95% had neutered, 85% vaccinated, and 75% microchipped their cat(s) already. Overall two-thirds had done all of these, indicating a third of cats could potentially impact wildcats negatively. 53% had already heard about the project (84% for those living near one of our PAs), so this sample has an unrepresentatively high number of owners already engaged with responsible cat ownership, suggesting in reality that more than a third of pet cats could potentially impact wildcats negatively.

• A placement student helped us by conducting a questionnaire survey of 147 farmers and crofters. Smaller farms and crofts were highly represented in responses. Thirty nine per cent had cats living on their farm or croft with an average of 0.42 pet, 0.35 farm, and 0.27 unowned cats per farm or croft (of the 147 total). Neutering and vaccination rates for pet cats were similar to those reported above for the more general population (97% neutered and 75% vaccinated). Rates were lower for farm cats (70% and 20%) and lower still for unowned cats (15% and 0%), with 65% of farms or crofts not knowing the status of the unowned cats that lived at or visited the property. Linked to the lower neutering rates for farm and unowned cats, at least 10% of these had kittens in the past year. The largest barrier to neutering and vaccination reported was that it was not the farmers’ responsibility (49%), that they did not have time (16%), or that they could not catch the cats (14%). However, other research conducted by RDSVS has found higher compliance, with the main barrier to neutering quoted as practicality. Nevertheless, a significant proportion of cats on farms may present a risk to wildcats, where their ranges overlap. We followed this up in our regular column in the Farming Scotland magazine by encouraging farmers to have the cats on their farms neutered and vaccinated.
• We did not have the opportunity to assess changes in public attitudes and behaviour around responsible cat ownership over the project duration and would recommend a follow-up questionnaire. Unneutered and unvaccinated cats could present a problem for any future wildcat conservation projects. Legislation to enforce neutering and vaccinating would benefit wildcat conservation, but some cat welfare organisations have argued against such a mandatory approach. 
• However, there appears to be more support for compulsory microchipping. This would require owners to engage with responsible ownership, enable  a ‘census’ of owned cats in Scotland, and greatly assisting the  identification of owned pet cats during work to improve wildcat conservation (such as during TNVR)  or other land management activities.

9. COMMUNICATIONS

• The SWA project was the first coordinated, nationwide approach to wildcat conservation. We developed and implemented a national communications plan to support wildcat conservation actions.  Our key audiences included cat owners, land owners, farmers, gamekeepers as well as the wider public.
• We worked together with partner organisations to co-ordinate press releases and shared information to promote and deliver the project’s key messages.
• SWA was launched at a major press event at Holyrood in Edinburgh with the support of the Scottish Government’s Cabinet Secretary for the Environment, Land Reform and Climate Change, Roseanna Cunningham MSP. William the Wildcat, our adult-sized mascot, made his first appearance at the launch and was also used by partners to promote the project’s key messages at other events.
• Printed material was created and covered a range of key messages, such as responsible cat ownership, wildcat-friendly predator control and a camera-trapping guide. This was distributed at a variety of places, including veterinary practices and at events.
• One of our main aims was to raise awareness and carefully challenge and change public attitudes to domestic cat management through clear communications. We created the #Supercat campaign to promote responsible cat ownership and encourage neutering, vaccination and microchipping of pet and farm cats. The #GenerationWildcat movement encouraged people to take ownership of being the last generation who can save the wildcat from extinction in Scotland.

• Media coverage was steady throughout the project, ensuring our key messages and progress were publicised. Stories were picked up via our press releases, blogs published on our website and social-media posts. We worked with media students to create The Tigers of Scotland documentary, which was picked up by Netflix, iTunes and Amazon Prime. Our work was also broadcast on television (e.g. the BBC's Springwatch, Winterwatch and Countryfile) and radio (e.g., BBC Scotland’s Brainwaves and BBC Radio 4). Certain events triggered an increase in coverage, such as the birth of wildcat kittens in collections managed under the conservation breeding programme, which was picked up locally, nationally and internationally.

 

• There was occasional negative press promoted mainly from an outside interest group, the rebuttal of which took up valuable staff time that would have otherwise been invested in wildcat conservation.
• We developed a website, which provided a one-stop-shop for information on the wildcat and project updates. Our leaflets and posters were published on the website, along with educational materials, and blogs which provided updates and emphasised particular messages. The website also published general information about the wildcat and our various campaigns, such as #Supercat and the TNVR programme. Google Analytics was used to measure and track website usage, to ensure our content was engaging. New content, such as blogs, were uploaded regularly to maintain public interest.
• Social media was an important tool for circulating information, engaging the public and keeping the project current. Facebook and Twitter were the most popular, with LinkedIn and Instagram also used. We utilised the platforms’ analytics to measure engagement with posts and to learn the most engaging ways to communicate our key messages. Our social-media users engaged with fact-based information as well as more light-hearted posts. The #WildcatOrNot feature worked well as the public could actively participate. Using pictures and trending hashtags increased engagement.
• Email newsletters were distributed via the Campaign Monitor platform, which provided email updates to subscribers.
• An education package was developed by RZSS for schools in line with the Curriculum for Excellence. The aim was to engage younger audiences in the plight of the wildcat and the need to save it. Funding was obtained from the Gordon and Ena Baxter Foundation in addition to that from the National Lottery Heritage Fund.
• We worked with the Mammal Society on their established Mammal Tracker app and its replacement, Mammal Mapper.  Efforts were made to ensure accurate wildcat information was available and that people could record sightings.
• Our two most frequently highlighted messages were about responsible cat ownership and the reporting of cat sightings.

10. A SUMMARY OF KEY RESULTS

Population status and defining a wildcat

• It is difficult to estimate a precise number of wildcats remaining in Scotland, in part because of the difficulty in distinguishing wildcats from hybrids and in part because of their secretive nature. The project has amassed a large body of evidence, including detailed knowledge from within its PAs and indicative information elsewhere, that there are too few wildcats remaining to be viable as a population.
• Cats that look like wildcats are outnumbered in the wild in the PAs by at least 10:1. Of these cats that look like wildcats, the majority are hybridised at a level where more than a quarter of their ancestry is from domestic cats. Furthermore, we have no recent evidence of wildcats from public sightings, camera-trap surveys or road-killed cats in the Highlands north of Lairg and scant evidence of any in in Argyll and the Trossachs (although we recognise that there are  gaps in our data).
• This situation, where wildcats, wildcat hybrids and domestic cats have mated freely with each other over many generations (known as introgressive hybridisation), has led to a hybrid swarm, where the population of wild-living cats in in Scotland ranges across the spectrum from wildcat to domestic cat. The high degree of hybridisation that has affected the wildcat appears to have started since 1960 based on the results of morphological studies, and since the 1990s based on genetic studies, but there are few data available for the period 1960-1980.
• In reality, few cats living wild in the areas where we collected the most data can be considered wildcat and data submitted by the public from other areas within the former range of the wildcat give us no reason for hope that the situation elsewhere is any different. Instead, we appear to have a wild population consisting mostly of hybrids.
• ​​​​​​Introgressive hybridisation has furthermore led to a breakdown in a previously well-defined relationship between the cats’ external appearance and their underlying genetics. The situation is complex. Some cats that do not look like wildcats have significant wildcat ancestry as revealed through genetic screening, while a few cats that look like wildcats do not have a significant wildcat ancestry. However, there were few cats scoring very highly on pelage that scored poorly on genetics.
• ​​​​​Therefore, in most cases where animals can only be observed, or are handled for only a brief period, a cat’s appearance has to remain the appropriate guide for identification. This should therefore be applied to situations where access to genetic data is not possible or practical, such as during predator control by land managers, neutering programmes or surveys ahead of forestry operations or developments.  The simple definition of a wildcat referred to at the beginning of this report therefore applies in most situations “a striped tabby cat with a thick, ringed, blunt, black-tipped tail and no stripe down the middle of the tail”.
• Our research also highlights the importance of using both appearance and genetic testing in identifying wildcats where this is possible.  For example, there is an expectation that possible wildcats held for rehabilitation purposes for more lengthy periods of time should have the appropriate genetic test, as well as a pelage assessment verified by an appropriate wildcat specialist nominated by NatureScot.

Insights into wildcat ecology

• The wildcat hybrids collared in the PA’s have very large home ranges, some of which are similar in size to those of mainland European wildcats living in areas of low quality habitat and/or low prey population densities. Large home range sizes can also result from low population densities. 
• Each individual wildcat hybrid used many tens of resting sites comprised of a wide variety of features within their home range, including wood stacks, areas of wind-blow, clear-fell, brash piles, hedgerows, outbuildings, rocky crevices and gorse scrub. This is similar to wildcat den/rest site use in mainland Europe, and suggests that resting site availability may not limit wildcat populations. This may not however, be the case for breeding site choice which will be influenced by a variety of factors including proximity to prey and cover. This underlines the need to review guidance and advice to support the protection of specific dens or rest sites, and encompass other factors that may be as important to their survival.
• Wildcat hybrid habitat use is similar to that of wildcats in mainland Europe, with preferred habitats including broadleaf forest, scrub, young conifer plantations, and clear-felled forest patches. They also made use of linear features, such as roads, tracks, hedgerows and fence lines. Wildcat hybrids usually preferred to use areas closer to habitat edges, probably because they use some wooded habitats for cover and some open habitats for hunting. In the Angus Glens PA, some individuals showed seasonal shifts in their home ranges, moving to open moorland in the summer, probably attracted there by a variety of potential prey species (rather than specifically to predate grouse).
• The GPS tracking data also highlighted some potential risks to wildcat populations in Scotland. The regular use of farm structures by wildcat hybrids shows how important neutering feral farm cats is to reduce hybridisation and that there is a risk of potential secondary rodenticide poisoning through the consumption of rodents found in many farm buildings. The use of open heathland/grouse moors by some individuals stresses the need for continued work with the game keeping industry to improve identification techniques and increase wildcat-friendly practices. Road mortality is a significant issue for wildcats across their range and the regular use of roads by the collared individuals (and the number of road casualties across the project) suggests this is likely to be an issue for wildcats as well. Initiatives to identify areas of potential high risk for wildcats in Scotland should be undertaken to try and reduce road-related wildcat mortality.

Management of domestic and hybrid cats

• We found a broad range of infectious agents in wild-living cat populations and detected FIV in a hybrid wildcat for the first time. These diseases are a potential threat to wildcats.
• We learned important lessons in implementing TNVR as a management tool for feral domestic cats and hybrids. Camera trapping, to identify where the cats are, is an efficient method for detecting wild-living cats. Local knowledge is also essential when finding feral domestic cats around farms settlements.
• TNVR requires plenty of staff resources, and we found that few volunteers were able to commit sufficient time to such an intensive undertaking. Achieving the neutering rates needed to reduce the population size of feral domestic and hybrid cats may be possible in discrete areas, but the resources required to attain this in the long term at the larger scales needed to reduce their numbers across Scotland means that such aims are unlikely to be achievable . By itself, TNVR cannot be the only method for feral domestic and hybrid cat management in Scotland.
• Domestic cat neutering rates in Scotland are high and rates of neutering for farm cats only slightly lower. Vaccination rates are lower than neutering rates. Nevertheless, the fraction of pet and farm cats that are unneutered could make a significant contribution to recruitment to the feral domestic cat population and the spread of diseases. Our campaign to encourage cat owners to have their pets neutered and vaccinated will likely have led to more neutering around the PAs, but the fraction of pet and farm cats that are not neutered remains an unresolved issue that threatens the future of wildcat restoration and conservation in Scotland. Our finding that hybrid wildcats visit farms underlines this risk. Changes to the legislation around cat ownership are required to ‘turn off the taps’ of domestic cat recruitment into the wild. As a minimum this should involve compulsory neutering, possibly in some specific geographic areas (with appropriate exceptions for some domestic cats for breeding etc.) and microchipping. We are aware that the recommendation for compulsory neutering is likely to be questioned by some, but we would welcome further discussions with key stakeholder organisations to find a way forward to ensure we are able to restore a long term, viable wildcat population to Scotland. Compulsory microchipping by itself would, at least make the task of TNVR less difficult.

Lesson for land management

• Accidental killing remains a threat, especially with the emergence of new technologies, such as thermal imaging. Since so few wildcats remain this may lead to a risk of complacency from land managers, who may believe that the cat in front of them is unlikely to be a wildcat. Such complacency needs to be discouraged since there are individuals living in the wild in Scotland that have passed both pelage and genetic thresholds and therefore there remains a danger that a wildcat could be shot during predator control work.  
• Productive forestry is compatible with wildcats, but can be improved by employing certain forms of forest management. These include retaining scrub, brash and wind-blow where it is safe to do so, ensuring there is a network of open habitats suitable for wildcat prey and planting more broadleaf trees. In addition, reducing certain activities during the main breeding season for wildcats (March – August) will also help reduce the risk to female wildcats with dependent young. Such management is likely to help other species too and is already being adopted as part of wider policy by Forestry and Land Scotland.
• Surveys ahead of forestry operations could be improved to reduce the risk of disturbance. Dens and other resting places are very difficult to find and the use of detection dogs could be explored to improve detection rates, noting the restrictions set out within the Hunting with Dogs (Scotland) Bill (2023). This is probably most important during the breeding season when there is a need to avoid disturbing mothers with kittens. Since the use of detection dogs also risks disturbance, handlers should undergo appropriate training and certification.
• Resting places may not be limiting wildcat numbers and so it may be better to consider the requirements of wildcats across the landscape, whether it is forest, farmland or moorland. This means ensuring that a mixture of cover and good prey habitats is available, alongside retaining features in the landscape that may be used for breeding or resting.
• The use of rodenticides to control rodent pests is a potential threat to wildcats through secondary poisoning. The chief source of secondary poisoning is unclear and likely to be from a combination of farms, other industries and households. Therefore, advice on safe use of rodenticides needs to be targeted at all potential users.

Establishing a conservation breeding programme

• The captive wildcat population, sourced from Scotland before introgressive hybridisation was widespread, showed low levels of hybridisation, and notably lower than the levels of hybridisation now evident in the wild.  Screening of the captive population has allowed us to reduce hybridisation levels further. Detailed genetic analysis has allowed the development of a detailed studbook based on genetic pedigree, something that was not available prior to the project.
• During the project we increased the size of the captive population by over two thirds to more than 100 individuals, including two wildcats from the wild. Offspring survival in the programme has been high.

Public engagement

• Our engagement with the public through our website and social channels, media, campaigns, education and public events was undoubtedly effective in raising the profile of wildcat conservation and getting our messages across to the public. This has been helped by the fact that the wildcat is a charismatic species with wide public appeal. However, certain aspects of domestic cat management can be controversial, such as the debate over neutering versus lethal control for feral domestic cats and obvious hybrids, and whether neutering of domestic pet cats should be made compulsory, at least in some areas. It can also be difficult to set out simple messaging around hybridisation and the conservation value of cats that are not ‘100% pure’ wildcat. The significant expertise in science, conservation and land management held within the project partnership enabled us to address some of these complex issues. Nevertheless, communications around wildcat conservation remain challenging. Our communications represent the first step in a long conversation with the public and land managers on the protection of wildcats and the management of domestic cats and hybrids.

CHAPTER 3: MOVING FORWARD

 

The IUCN Review

As we gathered data on the status of wildcats in Scotland over the course of the Scottish Wildcat Action project, we became increasingly concerned about what the data showed. The SWCAP Steering Group decided to seek an independent review of the data and wildcat conservation in Scotland. In spring 2018, NatureScot on behalf of the Steering Group, commissioned members of the International Union for the Conservation of Nature (IUCN) Species Survival Commission (SSC) Cat Specialist Group to undertake a review, based on the scientific literature and available reports and data. The findings of this Review (Breitenmoser et al., 2019) were stark:

“All the robust information available indicates that the wildcat in Scotland is at the verge of extinction. Based on the available information, we consider the wildcat population in Scotland to be no longer viable. The number of wildcats is too small, the hybridisation too far advanced and the population too fragmented. We therefore conclude that it is too late to conserve the wildcat in Scotland as a stand-alone population.”

In arriving at this conclusion, the authors drew on much of the data made available in the technical reports and associated annexes that accompany this summary report. None of the data collected since then, much of which are also available in the accompanying technical reports, gives us reason to doubt this conclusion. Scottish Wildcat Action achieved a great deal during its work, but the Review concluded that halting the further decline of the wildcat in Scotland, as set out in the original conservation action plan (Anon, 2013), was now unattainable given the status of the wild population. The Review highlighted the achievements of Scottish Wildcat Action in providing the evidence and experience to inform this difficult conclusion:

“…this is not an indication of a failure of the programme, but simply the consequence of more reliable data and a more realistic understanding; we think that earlier assessments of the situation of the wildcat in Scotland were too optimistic. So an important achievement of SWA has been to present a more realistic picture of the situation of wild-living cats in Scotland.”

The Review proposed a solution:

“…we still recommend that efforts are made to save as much of the gene pool of the Scottish wildcat population as possible. We reason that (1) the recovery of the wildcat in Scotland will only be possible with the support of reintroduction/reinforcement projects, and that (2) the remaining “pure Scottish wildcats” (based on a rigorous genetic protocol) should be combined with wildcats from continental Europe. Both, reintroduction projects or reinforcement of remnant nuclei should be considered. This requires a rigorous suppression of feral cats and hybrids in areas of reinforcements, and the removal of feral cats in areas of potential reintroduction.”

And the Review further highlighted the risk that domestic and hybrid cats present:

“Considering the efforts and the achievements so far, responsible cat ownership, the control of feral cats and removing the hybrids especially in the remnant wildcat areas of Scotland, seems to be the major challenge in wildcat recovery across the UK. The history of continental [European] wildcat populations, some of which are presently expanding, demonstrates that wildcats can also maintain their genetic integrity also in landscapes with high densities of domestic cats. However, small remnant or initial reintroduced populations are likely to be much more vulnerable to hybridisation and may require stronger management interventions to prevent hybridisation than a large vital population.”

A workshop was then held in Battleby, October 2018 (at which point the IUCN review was completed although was not formally published until a few months later) facilitated by the IUCN SSC Conservation Planning Specialist Group. It involved SWCAP Steering Group members, partners of the wider Scottish Wildcat Conservation Action Plan and members of the IUCN SSC Cat Specialist Group. The findings of the Review formed the focus of the workshop, and participants were invited to discuss and identify ways forward.  These discussions helped to provide agreement and clarity on the best way forward for wildcat conservation in Scotland.

The SWCAP Steering Group welcomed the findings of the Review. While we believe that some wildcats may remain in the wild in Scotland, achieving a consensus that the population is no longer viable allows us to take the essential next steps to give the wildcat a future. This must include a conservation-breeding programme, drawing on the captive populations, the status of which was improved under Scottish Wildcat Action. A key part of any future work will also be a national conversation on the management of domestic pet, feral and hybrid cats.

Saving Wildcats

Following the recommendations set out in the IUCN Review and the wildcat workshop, and drawing on the expertise built during Scottish Wildcat Action, the Royal Zoological Society for Scotland led a successful bid to secure funding from EU LIFE for a major new project called Saving Wildcats www.savingwildcats.org.uk. Partners in this project include members of the SWCAP partnership (NatureScot, Cairngorms National Park Authority and Forestry and Land Scotland) alongside European partners with experience in conservation breeding and reintroductions (Nordens Ark and Junta de Andalucía). The aim of Saving Wildcats is to:

• Establish Britain's first large-scale dedicated conservation ‘breeding for release’ centre for wildcats.
• Grow the population of wildcats through release of wildcats into the wild.
• Continue to remove the threats facing wildcats in the Highlands to create other safe areas for wildcat reintroductions.
• Work with local communities to understand how best people can benefit from the presence of wildcats.

The projects central aim is to release 20 individual wildcats each year from the conservation breeding centre, although translocating cats from European populations to help boost the gene pool is considered as an option. These individuals will undergo an extensive pre-release training programme to ensure they are prepared for life in the wild. Such releases will begin from spring 2023. The project applies the IUCN’s One Plan approach, whereby the captive and wild population are managed as one unit and their conservation involves all responsible parties. This six-year programme represents the next, but certainly not final, step in restoring the wildcat in Scotland. Part of the Saving Wildcats project plan is to build partnerships, expertise and activities that can continue after the project finishes in 2027.

 

A long-term vision

The ultimate goal for wildcat conservation in Scotland must be to establish and restore a population of wildcats that does not require further intervention to secure the viability of the species, beyond ensuring that wildcats or their habitats are protected. This is likely to require action over several decades. As the work of Scottish Wildcat Action has shown, conservation interventions are extremely challenging when hybridisation has occurred. Wildcats are most at risk of hybridisation where populations are small and fragmented, and it is likely that once hybridisation occurs, the chance of it perpetuating increases. To achieve this goal, there must be very low levels of hybridisation at the outset and the population needs to be large, encompassing at least the Highlands, but potentially all of mainland Scotland and perhaps all of mainland Britain. This goal could be achieved though staged discrete population reintroductions or reinforcements, but the eventual aim is for conservation at the regional or national scale. Therefore, threats to the wildcat must also be addressed at these scales. The key threats revolve around population size, fragmentation and the consequent risk of hybridisation. Other hazards, such as exposure to disease, can also be reduced by our actions to counter these three key threats.

Hybridisation

In the Highlands, we have a well-adapted population of wild-living hybrid cats occupying much of the niche that should be inhabited by wildcats. Elsewhere, feral domestic cats may occupy at least part of this niche. Trapping for neutering is by itself unlikely to achieve sufficient population reductions in these cats to remove hybridisation risk at a regional or national scale, although it may do so locally where investment is high. Two key elements of this problem are 1) the continuing recruitment into the feral or hybrid population from domestic pet and farm cats and 2) the feral and hybrid cats living in remote locations that are difficult to access, where trapping takes more effort.

Tightening the legislation around cat ownership, to encourage more responsibility from cat owners, could help address the first element, as long as such legislation also covers farm cats.

To resolve the second element, the wild-living hybrid/domestic population must be reduced dramatically. A review, perhaps involving a public consultation, will be needed to identify the best approach to managing our wild-living feral and hybrid cats. Though this is a long-term goal, trapping for neutering is likely to remain an important tool in the short term, used in coordination with other activities, although it requires substantial resourcing. On-going monitoring and analysis of dead wild-living cats will continue to be important to assess progress in reduction of the feral and hybrid cat populations and offer valuable opportunities for assessing diet, reproduction, disease and toxicology.

Where any releases of wildcats are carried out in areas unlikely to contain hybrids, because there has been no recent history of wildcat presence, securing the threat from feral, farm and house domestic cats will still be essential. A coordinated neutering and vaccination programme will also reduce risks from disease.

However, it is unlikely that the threats from feral or hybrid cats can be completely nullified. Therefore, it is essential that the root causes that led to hybridisation are addressed and we learn why hybridisation rates appear to be very low in many other European countries with viable wildcat populations (and the causes in the few regions where this is not the case).       

Predator control

Wildcat populations are at risk of becoming small and fragmented when too many are shot accidentally during predator control. At the same time, in some locations predator control may be the only method by which feral cat numbers are currently being managed. Wildcats are also known to use heather moorland in the summer months, where it can be an important habitat for them, putting them at added risk from persecution. There is a continuing need to reduce the risks to the very few remaining wildcats from accidental killing. As wildcat populations re-establish, this risk will increase and the need to apply precautionary predator control practices will become more important.    

Habitat

Wildcat populations become small and fragmented where there is insufficient habitat and where suitable habitat is itself fragmented. During this project, we discovered a lot about how wildcats use their environment, but we still have a lot to learn. For example, it appears that some areas in the Highlands are not supporting the numbers of cat (of any type) that they once did and we are not sure why. It is possible that the decline in rabbit numbers has played a role in this. More likely, there has been a combination of factors that have reduced the resilience of the cat populations to the loss of a key prey resource, such as rabbits. Rabbits are a non-native species and therefore, historically, wildcats must have survived on other prey. But of course the landscape is very different today. Prey densities are greatly affected by deer density, forest type and management, farming practices and will have been affected by generational shifts in land-use, such as changes in farming or upland management. More research is needed to clarify what has driven these changes.

A landscape containing a diverse prey base and sufficient cover habitat for denning and resting, is likely to support a more stable wildcat population than one dominated by one or two prey species and where there is little cover. Activities, such as deciduous-tree planting and the provision of uncultivated field margins, have been included in various agri-environment incentive schemes and can benefit many species. Expanding and coordinating this approach so that such habitats form a contiguous network, could help create a less fragmented landscape that contains greater biodiversity and can support a more resilient population of wildcats. Research in mainland Europe has also shown that wildcats require forests and large areas of cover with high structural diversity in them to survive at high population numbers. In Scotland, much of our woodland is coniferous plantation and here structural diversity can be improved through retention of areas of deadwood and wind-throw, patches of gorse and scrub on forest and farmland edges and ensuring woodland management retains a mosaic of species and ages of trees. Land management practices that don’t negatively impact on prey species should be encouraged as well.  

Where next?

Solving these key threats will require a coordinated effort, with the first step towards this the creation of a new Action Plan for wildcats in Scotland before the Saving Wildcats project ends. Elements to address in this action plan will include much of what has been discussed above, as well as the need for ongoing monitoring, improved guidance and training for land managers, and reviewing the legislation around cat ownership. Meanwhile, the public can still contribute by submitting sightings of wildcats and ensuring their own pet cats are neutered and vaccinated.

It is encouraging to see a wider interest in restoring the wildcat in England and Wales. A British-wide wildcat population would provide the best insurance policy against future problems. However, the emergence of another hybrid population in one of the British nations could put at risk populations elsewhere in Britain. Inadequate population reinforcement or not mitigating against risks at the release sites and around the expanding population front could make hybridisation inevitable. To minimise this risk, projects in the different nations need to coordinate conservation breeding and to share knowledge and experience in reducing threats.

Finally, restoring the wildcat is a long-term undertaking and will require long-term commitments from government, non-governmental organisations and the public to succeed.  The risk of partial, short-term success is an eventual return to the current situation and the potential loss of the wildcat in Scotland. The benefit of complete success is the restoration of this iconic species, wider biodiversity and ecosystem gain, and less need for further intensive intervention.  

FURTHER READING

The SWA Specialist Reports:

Bacon, A., Beckmann, K.M., Anderson, N.E., Alves, B.S.G., Ogden, R., Meredith, A.L. (2023). Scottish Wildcat Action final report: Disease surveillance. NatureScot, Inverness.

Barclay, D. Senn, H. (2023). Scottish Wildcat Action final report: Conservation breeding, NatureScot, Inverness.

Campbell, R.D., Burns, V., Bacon, A. & Gaywood, M. (2023). Scottish Wildcat Action final report: Responsible cat ownership. NatureScot, Inverness.

Campbell, R.D., Kortland, K., Strong, R., Tallach, N., Cumberbirch J. and Mason, G.  (2023). Scottish Wildcat Action final report: Land management. NatureScot, Inverness.

Campbell, R.D., Langridge, K., Kilshaw, K., Carus. H., Hislop, C., Rawling, E., and Tallach, N. (2023). Scottish Wildcat Action final report: Survey and monitoring. NatureScot, Inverness.

Campbell, R.D., Rawling, E., Langridge, K.V. and Tallach, N. (2023). Scottish Wildcat Action final report: Trap Neuter Vaccinate Return Programme. NatureScot, Inverness.

Kilshaw, K., Campbell, R.D., Kortland, K. and Macdonald, D.W. (2023). Scottish Wildcat Action final report: Ecology. NatureScot, Inverness.

Kitchener, A.C, Senn H (2023). Scottish Wildcat Action final report: Wildcat genetics and morphology. NatureScot, Inverness.

Wemyss, J., McKenzie, D., Burns, V. and Campbell, R.D. (2023). Scottish Wildcat Action final report: Communications. NatureScot, Inverness. 

Other reading:

Anon (2013). Scottish Wildcat Conservation Action Plan. Scottish Natural Heritage, Inverness.

Breitenmoser U., Lanz T. and Breitenmoser-Würsten, C. (2019). Conservation of the wildcat (Felis silvestris) in Scotland: Review of the conservation status and assessment of conservation activities. Available at:

Howard-McCombe, J., Ward, D., Kitchener, A. C., Lawson, D., Senn, H. V., and Beaumont, M. (2021). On the use of genome-wide data to model and date the time of anthropogenic hybridisation: An example from the Scottish wildcat. Molecular Ecology. 30, 3688– 3702. 

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ACKNOWLEDGEMENTS

Scottish Wildcat Action (SWA) was supported by numerous professional and voluntary workers and organisations across a wide spectrum of society, too numerous to acknowledge here. Members of the public provided vital help through donating funds and time, and by ensuring their pet cats became ‘Supercats’. Their dedication and contributions to the vision of restoring the wildcat to Scotland has also helped to build the foundation for the next phase of work, the ‘Saving Wildcats’ project. Our thanks go to everyone who has helped, including the many individuals we have not been able to list here.

Scottish Wildcat Action (SWA) was supported by the National Lottery Heritage Fund.

The SWA team carried out and organised much of the project work, its members were David Barclay, Vicky Burns, Roo Campbell, Hebe Carus, Calum Hislop, Keri Langridge, Duncan McKenzie, Sarah O’Mullan, Emma Rawling, Nicola Tallach, Jenny Wemyss, Matt Wilson.

Several contractors, students and full-time volunteers also contributed significantly through field or technical assistance and data analysis. These included Phil Bacon, Laura Braunholtz, Roisin Campbell-Palmer, Jen Clark, Declan Duffy, Rhian Evans, Martha Gavan, Claire Hill, Sima Lionikaite, Ana Luisa Barros, Kirstin Mair, Ana Morales González, Fiona Pamplin, Lydia Peters, Héctor Ruiz, Lara Semple, Laura Sessions, James Walker, Barrie Williams, Kelsey Wilson, and Matt Wilson.

Additional funds were provided by the Scottish Government, NatureScot, the Royal (Dick) School of Veterinary Studies at the University of Edinburgh, the Royal Zoological Society of Scotland and through donations received from members of the public. SWA was led by NatureScot, with additional significant support and resources provided by the members of Steering Group. The Steering Group members, and their representatives were:

• Cairngorms National Park Authority – Will Boyd-Wallis,  David Hetherington
• Forestry and Land Scotland – Kenny Kortland [with additional support from Jackie Cumberbirch, Henry Dobson, Graeme Finlay, Gareth Mason, Philippa Murphy, Chris Nixon, John Thomson, Gareth Ventress]
• National Museums Scotland – Andrew Kitchener
• NatureScot – Jenny Bryce, Lynne Clark, Martin Gaywood, George Hogg, Kath Leys, Eileen Stuart [with additional support from Bruce Brown, Christian Christodoulou-Davies, Kieren Jones, Fergus Macneill, Alison McQuaker, Eleanor Meikle, Dominic Shann, Fiona Strachan, Cat Synnot]
• Scottish Land and Estates – Karen Ramoo, Rebekah Strong
• Scottish Wildlife Trust – Olly Davies, Pete Haskell
• The National Trust for Scotland – Stuart Brooks, Lindsay Mackinlay, Steven Reeves
• The Royal (Dick) School of Veterinary Studies, University of Edinburgh – Alice Bacon, Katie Beckmann, Rob Ogden, Anna Meredith
• The Royal Zoological Society of Scotland – Sarah Robinson, Helen Senn
• The Scottish Gamekeepers Association -  Bob Connelley, George Macdonald
• Wildlife Conservation Research Unit (WildCRU), University of Oxford – Kerry Kilshaw, David Macdonald
• Independent – Allan Bantick

The Scottish Government contact for the Steering Group was Hugh Dignon, with additional advice from Andrew Voas.

Additional sponsors, funders and contributors to SWA work included Aberdeenshire Council, Aigas Field Centre, Ailsa Black, British Association of Shooting and Conservation, Cairngorm Brewery Company, Chester Zoo, The Gordon and Ena Baxter Foundation, The Highland Council,  Highland Foundation for Wildlife, Highland Tiger Fling, John Muir Trust, Loch Lomond and Trossachs National Park, Lukas Ruiter Production, MyPetOnline, NFU Scotland, The Peoples Trust for Endangered Species, petDetect, RSPB, Scotland: The Big Picture, and Swift Ecology, The Robertson Foundation funded the majority of the GPS tracking.

Special thanks also go to Roseanna Cunningham MSP, former Cabinet Secretary for Environment, Climate Change and Land Reform for her support throughout the project. We are also very grateful to the many academics, animal welfare experts, land owners and managers, farmers, foresters, keepers, captive collection managers, veterinary specialists, consultants and others who contributed valuable time and effort, members of the public who contributed funds and support, pet owners who responded to the ‘Supercat’ campaign, and in particular the many volunteers who got involved in the project.

ISBN: 978-1-78391-975-8