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Research essayKnowledge & Research•Updated October 2026•UK Squirrel Accord and APHA programme pages, the 2024 field-trial papers (Pest Management Science, Journal of Wildlife Management), Rushton 2006, Sainsbury 2000, Sheehy 2014/2018, Bertolino & Genovesi 2003, Benson 2013, the 1937 Order and 2019 Order, the Eddas and Mooney 1902. Conflicting popular figures flagged in the sources.

October 2026 · Musing

On Squirrels

Britain is trying to put the eastern gray squirrel on birth control with hazelnut spread. How a park ornament became an invader on three continents, why the red squirrel loses, what the pine marten changes, and the lore that followed the animal everywhere it went.

Grays in Britain

2.7M

Reds in England

~15,000

Took the bait

49–56%

Sciurus carolinensis

The gray

Native to the eastern United States. 400 to 650 g in Britain. Carries squirrelpox without symptoms. Invited onto four continents by people who liked it.

Sciurus vulgaris

The red

Native to Britain and Europe. 100 to 350 g. Dies of squirrelpox within two weeks of symptoms. Down from 2.5 million to about 287,000 in a century.

Martes martes

The pine marten

The native predator that was hunted out, is coming back, and empties woods of grays wherever it does. Reds, which co-evolved with it, cope.

The squirrels on my block in Kalorama were black. I took that for a Washington thing and it was, but not an old one: in 1902 the National Zoo, a ten-minute walk from my door, received eight black squirrels from the parks superintendent of Ontario, swapped for some ordinary grays, and let them go in the northwest corner of the grounds; more came in 1906, and a century later the black morph is up to a quarter of the squirrels in parts of the city[1]. Now I live in Chicago, inside the animal’s native range, where grays and the bigger fox squirrels share the parks and nobody has an opinion about them. In Britain the same animal is on a kill list, outnumbers the native red seventeen to one, and is about to be offered contraceptive in hazelnut spread from a feeder that weighs it before opening the door[2][3]. That is the thesis of this piece: the eastern gray squirrel is the rare invader that was invited everywhere it went, by name, by dukes and park boards and a diamond magnate, and the hazelnut program is the first attempt in a hundred and fifty years to take the invitation back without a gun.

01

Invited everywhere

Start with the fact that American cities had no squirrels. By the middle of the nineteenth century they had been hunted out of urban areas, and the animal that now defines a park was put there on purpose. The first documented introduction was Philadelphia’s Franklin Square in 1847, three squirrels with food and nest boxes; Boston Common followed in 1855 and New Haven Green by the early 1860s. The large releases came with the Olmsted-era parks of the 1870s, and Central Park had about 1,500 squirrels by the mid-1880s[4]. The historian Etienne Benson, who traced this, found the reasoning in the record: the squirrel was “living proof of the rewards of extending charity and community beyond the bounds of humanity,” and feeding it was adopted as a way of encouraging humane behavior in children. People had to stop shooting squirrels and start feeding them[4][5].

Britain imported the ornament. The first verifiable release was a pair let go in Henbury Park, Cheshire, by Thomas Brocklehurst in 1876. In 1890 ten squirrels from New Jersey went to Woburn Park, and from Woburn the 11th Duke of Bedford gifted them around the country for thirty years, including a release in Regent’s Park that likely seeded London; 100 more went into Richmond Park in 1902, and the last documented release was Needwood Forest in 1929[6][7]. A 2016 genetic study of 381 squirrels from fourteen sites confirmed that the British population descends from a handful of deliberate human moves rather than from the animal spreading on its own: Aberdeen’s grays are closest kin to the New Forest’s, four hundred miles away[7]. Just over seventy percent of the successful introductions began with fewer than ten animals[6].

Ireland got a wedding present. In the summer of 1911 a wicker basket was opened on the lawn of Castle Forbes, County Longford, and about a dozen grays from the Woburn stock jumped out[8][6]. Cecil Rhodes put the first batch on his estate at Groote Schuur, Cape Town, soon after 1900, and a 1957 bounty of sixpence a tail paid out two guineas in total[9]. Italy’s came to Piedmont in 1948; when a trial eradication began at Racconigi in 1997 and the early results showed it was feasible, animal-rights groups sued, the biologists were acquitted three years later, and by then the window had closed[10]. Only Australia reversed it: Melbourne’s colony, introduced around 1900, was gone by 1973[11].

Figure 1

A century of invitations

1840186018801900192019401960US citiesBritainIrelandSouth AfricaAustraliaItaly1937: keeping one banned1847 Philadelphia: three squirrels in Franklin Square, with nest boxesPhiladelphia1855 Boston: Boston CommonBoston1884 Central Park: about 1,500 squirrels by the mid-1880sCentral Park1902 DC (black): 8 black squirrels from Ontario to the National Zoo, 1902; more in 1906DC (black)1876 Henbury: T. V. Brocklehurst releases a pair in Cheshire: first verifiable recordHenbury1890 Woburn: ten from New Jersey at Woburn Park; the Duke of Bedford then gifts them around BritainWoburn1902 Richmond Park: 100 releasedRichmond Park1929 last release: Needwood Forest, Staffordshire: the last documented releaselast release1911 Castle Forbes: about a dozen, Woburn stock, a wedding present opened on the lawn in Co. LongfordCastle Forbes1902 Cape Town: Cecil Rhodes, Groote Schuur, soon after 1900Cape Town1900 Melbourne: c. 1900; gone by 1973 (secondary source)Melbourne1948 Piedmont: first reported near Stupinigi; 1997 eradication halted by lawsuitPiedmontEvery dot is a person letting squirrels go on purpose. Britain banned keeping one eight years after the last.

Documented deliberate releases, one lane per country[4][6][8][9][10][11]. The American dots are civic: park boards buying squirrels as a moral ornament. The British dots are mostly one family, the Dukes of Bedford at Woburn, gifting them for thirty years. The black dot is Washington’s 1902 swap with Ontario[1]. The red tick is the 1937 Order that made keeping a gray squirrel in Britain a crime, eight years after the last release[24].

02

Why the red loses

The numbers first. Britain had around 2.5 million red squirrels a century ago and has about 287,000 now, three quarters of them in Scotland; England is down to roughly 15,000. Grays, from zero, number at least 2.7 million[2][12]. The easy explanation is size, and size matters: a British gray averages around 550 g against a red’s 280 to 300, and grays pilfer red caches so effectively that reds in shared woods go into spring lighter and breed less[2][13]. But competition alone is slow. What made it fast is a virus.

Squirrelpox is carried by grays without symptoms. A 2000 survey found antibodies in 61 percent of 223 apparently healthy grays and in 2.86 percent of reds, four animals out of 140, which is the signature of a reservoir host[14]. In reds the disease is almost invariably lethal, with death typically within two weeks of symptoms; an outbreak at Formby in 2007 killed an estimated 85 percent of the colony[15]. Stephen Rushton’s group, modelling a decade of Cumbrian data, found red declines “17–25 times higher in regions where SQPV is present in grey squirrel populations,” and that stopping the decline by culling would need a kill rate above 60 percent[16]. The gray does not have to outfight the red. It has to stand near it.

Figure 2

Britain’s two squirrels, a century apart

0M1M2M3M190019251950197520002025Britain, animals (millions) → endpoints are measured; the lines between are only joinedRed squirrels, 1900: about 2.5 millionRed squirrels, 1995: about 161,000Red squirrels, 2018: about 287,000Gray squirrels, 1900: about 0 millionGray squirrels, 1995: about 2.5 millionGray squirrels, 2018: about 2.7 millionreds, 2.5M a century ago287,000 nowgrays, 2.7M in 2018

Endpoints from Natural England’s 2018 count (2.7 million grays), the UK Squirrel Accord’s red estimate (287,000, down from 2.5 million), Harris et al. 1995 (161,000 reds, 2.5 million grays) and Defra’s 2019 figure for England (about 15,000)[2][12][17]. The lines between the points are only joined, not measured; the shape is the point. The crossover happened before anyone counted properly.

Figure 3

The virus does the work

0%25%50%75%100%Grays with antibodies, apparently healthy61% of 223 grays, Sainsbury et al. 200061%Reds with antibodies4 of 140 reds, Sainsbury et al. 20002.86%Reds killed, Formby 2007estimated 85% of the Formby reds85%Cull needed to halt red declinemore than 60% effective kill, Rushton et al. 200660%With the virus present, reds vanish 17 to 25 times faster than with competition alone.

Sainsbury et al. 2000 seroprevalence (61% of grays, 2.86% of reds)[14]; the 2007 Formby outbreak[15]; Rushton et al. 2006, the Cumbrian model that put red decline at 17 to 25 times faster where the virus is present and the required cull above 60%[16]. A reservoir host is an animal that carries a disease it does not suffer from; the gray is the textbook case.

03

The bill

Grays strip bark. In 2021 the Royal Forestry Society commissioned a model of what that costs in England and Wales, assuming damage on 15 percent of broadleaf and 5 percent of conifer area: £37 million a year, £1.1 billion over forty years, more than 90 percent of it on broadleaves, with sycamore taking 40 percent of the damage, beech 26, birch 9 and oak 6[18]. The figure is a model, not a count, and the report says so. It is also the number that made the Treasury care, which is why the contraceptive got funded.

Figure 7

The bill, by tree

0%10%20%30%40%SycamoreSycamore: 40% of the modelled damage (Royal Forestry Society, 2021)40%BeechBeech: 26% of the modelled damage (Royal Forestry Society, 2021)26%BirchBirch: 9% of the modelled damage (Royal Forestry Society, 2021)9%OakOak: 6% of the modelled damage (Royal Forestry Society, 2021)6%All other speciesAll other species: 19% of the modelled damage (Royal Forestry Society, 2021)19%£37 million a year in England and Wales on the probable scenario; more than 90% of it to broadleaves.

Royal Forestry Society 2021, modelled by RDI Associates with the Forestry Commission, National Forest Company, Natural Resources Wales and the Woodland Trust[18]. “Probable scenario” assumptions: 15% of broadleaf and 5% of conifer area damaged. Sycamore and beech, thin-barked and fast-growing, take two thirds. The carbon lost is costed separately at about £9 million a year.

04

Hazelnut spread

The UK Squirrel Accord is a partnership of 48 forestry and conservation bodies, government agencies and companies. Since 2017 it has paid the Animal and Plant Health Agency about £1 million, under Giovanna Massei, to turn an injectable immunocontraceptive vaccine into something a wild squirrel will eat[3][19]. The vaccine works on the immune system to stop the body producing sex hormones, making both sexes infertile; the active ingredient is sealed in plant-spore capsules, the same delivery used for vitamin D in human trials, so it survives the gut[3]. The bait is hazelnut butter. The feeder is the clever part: a hopper that only grays can open, one design with a weighing platform that unlocks the door only for an animal inside a set weight range. Reds are too light[3].

Everything published so far uses a dye instead of the drug, to measure who eats. In six trials across five English woods, 271 squirrels were trapped and tagged and 151 of them, 56 percent, turned up dye-positive, spring highest and summer lowest; no other animal got into the hoppers[20]. A parallel study in three woods found 49 percent, with sex, age and breeding status making no difference to who ate[21]. Earlier modelling had used a 75 percent efficacy threshold, and the preliminary finding is that at least 60 percent of animals given the oral dose could become infertile[22]. In 2025 the Accord announced the formula had produced infertility in rats by free feeding and “a reaction” in grays, with field-scale trials, registration and rollout still ahead; its chair called it the world’s first oral free-feeding immunocontraceptive[3]. A Clemson trial of the injectable version in 2009 shrank testes and raised antibody levels but also left abscesses at the injection site, which is the argument for a spread you eat[23].

Figure 4

The weight gate

0 g250 g500 g750 g1,000 g1,250 gthe gap the door usesRed squirrel100 to 350 g (UK Squirrel Accord)100350Gray squirrel, UK400 to 650 g (UK Squirrel Accord); Yorkshire sample 422 to 760 g400650Fox squirrel, US550 to 1,400 g (USGS)5501,400One hopper design weighs the animal and opens only within a set range: grays in, reds out.

Mass ranges from the UK Squirrel Accord (reds 100–350 g, grays 400–650 g)[2], a Yorkshire APHA sample (grays 422–760 g)[13], and the USGS for the American fox squirrel[25]. The hopper’s weighing platform works because the two British species barely overlap. The fox squirrel is here for Chicago’s sake: it is the bigger animal in my parks now, and it would walk straight through a British door.

Figure 5

How many take the bait

0%25%50%75%100%49% of assessed squirrels dye-positive; sex, age and breeding status made no difference49%Tiberi et al. 20243 woods, 94 assessed151 of 271 dye-positive; spring highest, summer lowest56%Beatham et al. 20245 woods, 271 taggedat least 60% of animals given an oral contraceptive could become infertile (Massei et al., modelling paper)60%Preliminary infertilityanimals given the oral dose75% efficacy threshold used in earlier modelling75%Modelled targetfor landscape effectBoth trials used dyed hazelnut butter as a stand-in for the drug. Half ate within days. Half is not three quarters.

Field trials used 0.18% Rhodamine B dye in hazelnut butter as a stand-in for the contraceptive, with PIT-tagged squirrels checked for the dye afterward[20][21]. The 60% is the preliminary infertility rate from the modelling paper and the 75% is the threshold that paper’s predecessor used[22]. The gap between the bars on the left and the line on the right is the open question: enough squirrels have to eat, and then enough of those have to respond.

05

The enemy of my enemy

There is one place the gray has lost on its own. The pine marten, a native predator trapped nearly to extinction in Britain and Ireland, has been recovering, and Emma Sheehy’s four-year study of the Irish midlands found that “where pine martens had returned to healthy numbers, grey squirrels had all but disappeared,” and reds were recolonising woods that had been gray for thirty years[26]. The 2018 follow-up in Scotland, 223 feeders across the Borders, Central Scotland and the Highlands, put it formally: gray occupancy “strongly negatively affected by exposure to pine martens,” with an indirect positive effect on reds[27]. The mechanism is history. Reds co-evolved with the marten over tens of thousands of years and stay off the ground; grays are heavier, less agile, feed on the forest floor and have never met one[26]. A 2020 all-Ireland survey found grays had all but disappeared from ten midland counties[28].

Where there is no marten, people have done it by hand once. On Anglesey, grays arrived in the mid-1960s, and by 1998 fewer than 40 reds remained, all of one female bloodline, among some 4,000 grays. A trapping program began that year, reached the coastal boundary in 2013, and the island was declared gray-free in 2015 with around 700 reds[29][30]. Seropositivity in the island’s remaining grays fell from 75 percent to 4 percent as the cull proceeded[30]. Anglesey is 710 square kilometres with a strait around it. England is not.

Figure 6

Anglesey: the only mainland-scale win, on an island

01,0002,0003,0004,0000200400600800graysreds199620002004200820122016Anglesey, 710 km², 1998–2015 → endpoints measured; lines only joined1998: some 4,000 grays on the island2013: eradication to the coastal boundary complete; declared gray-free 20151998: fewer than 40 reds, all one female bloodline2015: around 700 redsgrays trapped outreds return

Fewer than 40 reds and about 4,000 grays in 1998; eradication complete to the coastal boundary in 2013 and declared in 2015; around 700 reds by then[29][30]. Two y-axes because the two species are an order of magnitude apart. Since the declaration about ten grays have swum the Menai Strait, and the last lone arrival in 2024 took six weeks to catch[31].

06

The animal itself

It is worth saying what the gray is good at, since everything above is about what it does to others. It is a scatter-hoarder, burying several thousand caches a season one nut at a time, and it finds them again: hand-raised squirrels tested at two, four and twelve days dug up significantly more of their own caches than of strangers’[32][33]. Recovery runs from 95 percent in lean years down to 26 percent in heavy mast years, when there is no need to bother[32]. And it lies. Michael Steele’s group at Wilkes University, in more than 500 trials, watched grays dig a hole, mime burying, and walk off with the nut still in their mouths; the fake burials ran about 13 percent of caching events, rose when another squirrel was within twenty metres, and doubled after the researchers deliberately stole caches. Human observers found the acorn one time in ten when the deception was used[34]. An animal that keeps a mental map of thousands of points and salts it with decoys is not a pest in any interesting sense. It is a competitor that happens to be winning.

07

Lore, and law

The stories agree on one thing: the squirrel is a go-between, and not a trustworthy one. In the Norse Eddas, Ratatoskr runs the trunk of the world-tree Yggdrasil carrying words between the eagle at the top and the dragon Nidhogg at the roots; the prose version says plainly that he “tells slanderous gossip, provoking the eagle and Nidhogg”[35]. In the Cherokee story of the ball game between the birds and the animals, recorded by James Mooney in 1902, the birds take a small creature and stretch its skin until it becomes Tewa, the flying squirrel, who catches the ball on almost the first toss and carries it up a tree[36]. In Wabanaki telling the squirrel was once a giant until Glooscap shrank it; in a Choctaw one a black squirrel eats the sun, which is an eclipse[37]. The Victorian version is Beatrix Potter’s Squirrel Nutkin, 1903, a red, impertinent to an owl on an island in Derwentwater, who loses his tail for it[38].

The law is its own lore. Britain’s Grey Squirrels (Prohibition of Importation and Keeping) Order of 1937 made it an offence to import or keep “any animal of the species designated Sciurus Carolinensis,” eight years after the last release; a 2015 deregulation dropped the duty on landowners to report them, on the stated ground that eradication was “neither feasible nor widely supported”[24]. Since December 2019 a rescued gray cannot legally be released back into the wild, which puts wildlife hospitals in the position of euthanising an animal they have just saved[39]. The epithet “tree rat” is the everyday version of the same thing, and Jason Gilchrist’s defence of the gray in 2017 notes that it outnumbers the red seventeen to one and that tens of thousands are culled a year[40]. Benson’s point lands here: a century ago feeding the squirrel was a lesson in kindness, and by the late twentieth century it had become “almost a crime or a sin”[4]. Same animal.

08

What to make of it

The gray squirrel is guilty of nothing except being carried. Every colony on the timeline was a gift, a civic project or a rich man’s estate, and the people who made them were acting on the best idea of their time, that a tame wild animal in a park teaches children to be gentle. The idea was right and the ecology was wrong, and the bill came due in a country the animal never asked to visit, paid by a smaller red squirrel that dies of a virus the gray does not notice. The honest accounting runs through the whole piece: a modelled £37 million, a measured 61 percent, a measured 56 percent, a declared island.

What I find myself respecting about the hazelnut program is its temper. For ninety years the only tools were a trap, a gun and a law, and they worked on an island with a strait around it and nowhere else. The alternative is a feeder that weighs the animal, lets it in, and gives it something it wants to eat, and the whole bet is that half the squirrels eating is enough. It may not be: half is not three quarters, and nobody has yet run the drug at scale. But it is the first fix in a century and a half that does not require anyone to hate the squirrel, and in Chicago, where I watch the same animal bury a walnut under the same tree twice and pretend the second time, that seems like the only version of this story I could stand to tell at the end.

Sources40 referencesShow

[1]DCist, “Blame Canada for D.C.’s Black Squirrels,” April 2011; Washingtonian, “Black Squirrels in Washington,” November 2008. The 1902 exchange with Thomas W. Gibson of Ontario, the 1906 batch, the release in the zoo’s northwest corner, and the morph’s present share. dcist.com · washingtonian.com

[2]UK Squirrel Accord, “Ecology.” At least 2.7 million grays (Natural England JP025, 2018); about 287,000 reds, down from 2.5 million; body-mass ranges for both species. squirrelaccord.uk/ecology

[3]UK Squirrel Accord, “Fertility control: 2025 update.” The partnership, the immunocontraceptive mechanism, Sporomex spore encapsulation, the weight-platform hopper, the rat result, Lord Kinnoull and Bex Pinkham quotations, and the R&D-to-rollout stages. squirrelaccord.uk/fertility-control-2025

[4]Benson, Etienne. “The Urbanization of the Eastern Gray Squirrel in the United States.” Journal of American History 100, no. 3 (December 2013). As summarised with quotations in ScienceDaily (6 Dec 2013) and the Penn Program on the Environment. sciencedaily.com · pan-school.sas.upenn.edu

[5]Harvard Magazine, “Squirrely,” December 2013. Harvard Yard’s squirrels c. 1900 and the acorn bounty paid to children. harvardmagazine.com

[6]Wildlife Online, “What’s the story behind the grey squirrel in Britain?” Compiling Lever, Naturalized Animals of Britain and Ireland, and Shorten 1954: Henbury 1876, Woburn 1890, Richmond 1902, Needwood 1929, the share of introductions under ten animals, and Watt 1923 on the Castle Forbes release. wildlifeonline.me.uk

[7]Signorile, A. L., et al. “Using DNA profiling to investigate human-mediated translocations of an invasive species.” Diversity and Distributions (2016), 381 squirrels, 14 sites; Imperial College press release, 26 January 2016, on the Duke of Bedford’s gifts and the Aberdeen–New Forest kinship. imperial.ac.uk

[8]Viney, Michael. “Grey squirrel’s bite is worse for the bark.” Irish Times, 11 September 2004. The wicker basket on the lawn of Castle Forbes, 1911. Popular accounts differ on the count (“six pairs”) and the giver; this piece uses “about a dozen, Woburn stock” per Watt 1923. irishtimes.com

[9]Pinelands History, “Pinelands squirrels.” Rhodes at Groote Schuur soon after 1900 (citing Davis 1951), the spread dates, and the 1957 sixpence bounty. pinelandshistory.co.za

[10]Bertolino, Sandro, and Piero Genovesi. “Spread and attempted eradication of the grey squirrel in Italy, and consequences for the red squirrel in Eurasia.” Biological Conservation 109, no. 3 (2003): 351–358. Piedmont 1948, the 1997 Racconigi trial, the lawsuit and the 2000 acquittal. iris.unito.it

[11]“Eastern gray squirrel,” Wikipedia, citing the IUCN assessment for Melbourne c. 1900 and extirpation by 1973. Secondary; the primary (Seebeck 2002) was not consulted. en.wikipedia.org

[12]Defra, “March 2019 update: invasive non-native species and grey squirrels.” “Only about 15,000 red squirrels remain in England.” gov.uk

[13]APHA Science Blog, “Saving the red squirrel,” 20 September 2022 (Yorkshire mass samples); Wauters, Tosi & Gurnell on cache pilfering and red spring body mass, via PMC. aphascience.blog.gov.uk · pmc.ncbi.nlm.nih.gov

[14]Sainsbury, A. W., P. Nettleton, J. Gilray, and J. Gurnell. “Grey squirrels have high seroprevalence to a parapoxvirus associated with deaths in red squirrels.” Animal Conservation 3, no. 3 (2000): 229–233. 61% of 223 grays; 4 of 140 reds. cambridge.org

[15]Wildlife Online, “Squirrel survival and mortality.” Lethality and two-week course in reds; the Formby 2007 outbreak; carrier share of grays. wildlifeonline.me.uk

[16]Rushton, S. P., et al. “Disease threats posed by alien species: the role of a poxvirus in the decline of the native red squirrel in Britain.” Epidemiology & Infection 134, no. 3 (2006): 521–533. Declines 17–25 times higher with SQPV present; cull above 60% needed in Cumbria. cambridge.org

[17]Wildlife Online, “Squirrel population,” citing Harris et al. 1995 (161,000 reds, 2.5 million grays) and the charities’ 140–160,000 consensus. wildlifeonline.me.uk

[18]Royal Forestry Society. “The cost of grey squirrel damage to woodland in England and Wales,” 2021, modelled by RDI Associates. £37m a year, £1.1bn over 40 years, species shares. rfs.org.uk

[19]Massei, Giovanna. Staff page, University of York. “Developing and delivering oral contraceptives for grey squirrels (2017–2024). Project leader. £1M, funder: UK Squirrel Accord” and the earlier DEFRA grants. york.ac.uk

[20]Beatham, S. E., et al. “Delivering bait to grey squirrels” (title abbreviated). Pest Management Science 80, no. 11 (2024): 5597–5607, doi:10.1002/ps.8379. 271 tagged, 151 (56%) dye-positive; no non-target uptake. Summarised with figures by Faunalytics, 24 September 2025. faunalytics.org

[21]Tiberi, M., S. E. Beatham, J. Coats, et al. Journal of Wildlife Management (2024), doi:10.1002/jwmg.22687. 138 tagged, 94 assessed, 49% dye-positive; no effect of sex, age or reproductive status. Data on Dryad. datadryad.org

[22]Massei, G., et al. “Modelling the management of an invasive species at landscape scale: are oral contraceptives the missing ingredient for success?” Botstiber Institute for Wildlife Fertility Control. The “at least 60%” preliminary infertility figure and the 75% threshold of prior modelling. wildlifefertilitycontrol.org

[23]Pai, Murali. “Field Evaluation of the Immunocontraceptive GonaCon in Reducing Eastern Gray Squirrel Fecundity in Urban Areas.” PhD dissertation, Clemson University, 2009. 317 squirrels; reduced scrotal size (p = 0.001); six injection-site abscesses. open.clemson.edu

[24]Grey Squirrels (Prohibition of Importation and Keeping) Order 1937 (S.R. & O. 1937/478), under the Destructive Imported Animals Act 1932; Deregulation Act 2015, Schedule 13, and its explanatory notes on eradication being “neither feasible nor widely supported.” legislation.gov.uk

[25]USGS, fox squirrel and gray squirrel mass ranges and habitat partitioning (publication 70263100); Frontiers in Ecology and Evolution 2020 on fine-scale competition. pubs.usgs.gov

[26]Sheehy, Emma, and Colin Lawton. “Population crash in an invasive species following the recovery of a native predator.” Biodiversity and Conservation (2014); Sheehy in The Ecologist, 28 August 2015, on the mechanism and the quotation used here. theecologist.org

[27]Sheehy, E., C. Sutherland, C. O’Reilly, and X. Lambin. “The enemy of my enemy is my friend: native pine marten recovery reverses the decline of the red squirrel by suppressing grey squirrel populations.” Proceedings of the Royal Society B 285 (2018): 20172603. 223 feeders, 2014–2017. st-andrews.ac.uk · abdn.ac.uk

[28]Vincent Wildlife Trust, “Red squirrels making comeback as return of native pine marten spells bad news for invasive grey squirrel,” 17 June 2020, on the all-Ireland survey (Lawton et al., Irish Wildlife Manual 121). vincentwildlife.ie

[29]Shuttleworth, Craig. Anglesey red squirrel project slides, LIFE U-SAVEREDS closing event (ISPRA), citing Schuchert et al. 2014 and Jones et al. 2017; fewer than 40 reds among some 4,000 grays in 1998, eradication to the coastal boundary by 2013. isprambiente.gov.it

[30]Schuchert, P., et al. “Landscape scale impacts of culling upon a European grey squirrel population.” Biological Invasions 16 (2014): 2381–2391 (seropositivity 75% to 4%); HPB, 2015, on the declaration and the rise to around 700 reds. research.bangor.ac.uk · hpb.co.uk

[31]Wales on Sunday, 7 April 2024, on post-declaration immigrant grays and the six-week capture. pressreader.com

[32]Wildlife Online, “Squirrel food, feeding and larders,” citing Jacobs 1989 (up to 95% recovery) and Steele & Smallwood 1994 (as low as 26% in mast years). wildlifeonline.me.uk

[33]Jacobs, Lucia F., and Emily R. Liman. “Grey squirrels remember the locations of buried nuts.” Animal Behaviour 41 (1991): 103–110, as summarised in Scientific American. “Several thousand caches per season” is the IUCN Red List’s figure. scientificamerican.com

[34]Steele, Michael A., Sylvia L. Halkin, et al. “Cache protection strategies of a scatter-hoarding rodent: do tree squirrels engage in behavioural deception?” Animal Behaviour (2008). Figures from the National Wildlife Federation’s account (13% fake burials; one in ten found; doubling after pilferage). One summary gives 22% at a second site. nwf.org

[35]Grímnismál 32 (Poetic Edda) and Gylfaginning 16 (Prose Edda), the latter in Jesse Byock’s 2005 translation: “He tells slanderous gossip, provoking the eagle and Nidhogg.” en.wikipedia.org/wiki/Ratatoskr · germanicmythology.com

[36]Mooney, James. Myths of the Cherokee. Bureau of American Ethnology, 19th Annual Report, 1902. No. 36, “The Ball Game of the Birds and Animals.” sacred-texts.com

[37]Native Languages of the Americas, “Native American squirrel legends”: the Wabanaki giant squirrel shrunk by Glooscap and the Choctaw eclipse story. Secondary compilation; Leland’s 1884 text was not consulted. native-languages.org

[38]“The Tale of Squirrel Nutkin,” Wikipedia: Frederick Warne, August 1903; Derwentwater and St Herbert’s Island; the picture-letter origin (Lear 2007). en.wikipedia.org

[39]Invasive Alien Species (Enforcement and Permitting) Order 2019 (SI 2019/527), article 37; Defra’s March 2019 statement that release licences for grays would no longer be issued and that vets are not required to euthanise. legislation.gov.uk

[40]Gilchrist, Jason. “In defence of the grey squirrel: Britain’s most unpopular invader.” The Conversation, 7 March 2017. “Tree rat,” the 17:1 ratio, the yearly cull. theconversation.com

A note on what was not verified. Several popular figures were checked and found to conflict: Ireland’s “six pairs”[8], the Australian dates[11], and the fake-burial rate (13% in one account, 22% in another)[34]. Where sources disagree the figure in the text is the one traceable to a named paper or contemporary record, and the sources above say which. The hazelnut trials’ counties are mostly unnamed in the papers; only North Yorkshire is confirmed.