UOA05-19: Securing the future of the globally threatened Large Blue butterfly
Submitting InstitutionUniversity of Oxford
Unit of AssessmentBiological Sciences
Summary Impact TypeEnvironmental
Research Subject Area(s)
Environmental Sciences: Environmental Science and Management
Biological Sciences: Ecology, Genetics
Summary of the impact
The Large Blue butterfly, formerly extinct in the UK, was successfully
reintroduced over two decades to sites in south-west England. New research
at the University of Oxford has greatly improved its conservation status
and identified key factors that determine the ability of this extreme
specialist to survive, especially in the context of climate change. Since
2008 this has led directly to new, larger and more stable populations, to
significant expansion of the butterfly's range into cooler regions, and to
new `races' with greater environmental tolerance. The research has thus
contributed directly to the positive upgrading of this species' global
The Large Blue butterfly, Maculinea arion, is an extreme
specialist: larvae feed briefly on thyme flowers before living for 10
months underground in Myrmica ant colonies, preying on ant grubs
whilst being tended by the workers whose Queens and larvae they mimic1-3.
In the 20th century the species declined across Europe,
becoming extinct in the UK in 1979 despite huge efforts to protect its
dwindling colonies. In the 1980s Professor Jeremy Thomas, then at the
Centre for Ecology & Hydrology, led a major initiative to reintroduce
M. arion to restored UK habitats. His research established that M.
arion survival depended on sufficient densities of one ant, Myrmica
sabuleti, a warmth-loving species that requires a very short sward
(< 1.4 cm) to maintain optimum temperatures in its brood chambers.
These findings informed conservation efforts, and led to intensive regimes
of grazing to maintain critical grass lengths: by 2006 several sites held
colonies of the butterfly although with one exception, all were small,
isolated and unstable.
In 2007, Professor Thomas moved to the University of Oxford's Department
of Zoology where his greatly increased scope for research has not only
improved existing conservation practices for M. arion but also
identified important factors that determine its survival (and that of
other endangered species) both now and as future environments change.
At Oxford, Thomas' studies of the ecology, population dynamics, behaviour
and physiology of M. arion (and other Maculinea species)
confirmed, and for the first time quantified, the extreme host specificity
of this butterfly and its close relatives3. The research
revealed important new findings about habitat quality and the butterfly's
ability to disperse and persist in meta-populations across landscapes4.
New analyses of population dynamics1,4 quantified the need for
≥ 68% of initial larval foodplants to grow within foraging range of M.
sabuleti for a population of M. arion to persist. The
analysis also showed, remarkably, that habitat with 100% coexistence
supported c.100-fold higher densities of butterfly than other
apparently suitable, but sub-optimal, habitats. Similar micro-habitat
variation was shown to affect other butterfly species, providing the first
explanation as to why many species whose potential UK ranges had increased
by >100 km due to climate warming had failed to expand4.
Thomas' group also investigated how Maculinea larvae engage in
chemical3 and acoustical2 mimicry of their host
ants, showing that their specialisms were more extreme and host-specific
at local scales than was previously thought. Thomas' 2007-9 analyses also
tested mathematical models describing M. arion population dynamics
under different conditions, and showed that they closely predicted data
recorded from reintroduction sites. These validated models are now used to
adjust management on suites of sites across landscapes1, 4-6.
Research at Oxford University also showed that optimum conditions for M.
arion were changing. Weaker grazing, generating a taller 2.1 cm
sward, is now optimal under the warmer climates of the past 20 years.
Moreover, on UK sites with exceptionally warm microclimates (e.g. steep
sheltered embankments), Thomas found that the niche of M. sabuleti
shifts to turf that is too tall (10-15 cm) for Thymus to flower.
Instead the butterfly lays on Origanum, a related labiate that
flowers 10-14 days later and on which larval survival is even higher. This
has led to the evolution of a late-emerging race of Large Blue, coinciding
with peak Origanum flower production. In contrast, on cooler sites
Thomas found that races had adapted to match the brief window of
thyme-flowering; 20 generations after establishment, some populations
develop more quickly than the original stock used to re-establish UK
Thomas had long argued that the restoration of a declining type of
habitat for one threatened species was likely to enhance the status of
other rarities that had been declining for broadly similar reasons.
However, this was disputed, and empirical confirmation was lacking. His
group at Oxford University quantified the beneficial increases in other
species that were evident anecdotally, and explained the reasons why this
occurred. For example, they found that the rare Pale Heath Violet, which
has increased 80-fold on sites managed for M. arion, has seeds
adapted to dispersal by Myrmica ants, but not by ants of other
genera. On these sites increases were also found in other rare
butterflies, other insects, plants and birds5,6, confirming
References to the research
1. Thomas JA, Simcox DJ, Clarke RT. (2009) Successful conservation of a
threatened Maculinea butterfly. Science 325: 80-83. doi:
10.1126/science.1175726 First rigorous analysis of 20-years'
M. arion conservation data; first test of models of
population; first analysis of M. arion metapopulations
and niche shifts under warmer climates.
2. Barbero F, Thomas JA, Bonelli S, Balletto E, Schönrogge K. (2009)
Queen ants make distinctive sounds that are mimicked by a butterfly social
parasite. Science 323: 782-785. doi: 10.1126/science.1163583 First
demonstration in any ant that queens make distinctive sounds, and of
acoustical mimicry by any social parasite.
3. Thomas JA, Elmes GW, Sielezniew M, Stankiewicz-Fiedurek A, Simcox DJ,
Settele J, Schönrogge K. (2013) Mimetic host shifts in an endangered
cuckoo-species of butterfly social parasite. Proceedings of the Royal
Society B 280: 711-719. doi: 10.1098/rspb.2012.2336 First
demonstration of regional host switches and local chemical adaptation
in Maculinea larvae.
4. Thomas JA, Simcox DJ, Hovestadt T. (2011) Evidence-based conservation
of butterflies. Journal of Insect Conservation 15: 241-258. doi:
10.1007/s10841-010-9341-z First study of relationship between
metapopulation structure and habitat quality of the Large Blue and
other butterflies, revealing key conditions for successful expansion
5. Thomas JA, Simcox DJ, Bourn NAD. (2011) Large blue butterfly. In Global
Re-introduction Perspectives 2011, ed PS Soorae, IUCN, Gland, pp
10-14. Review of the successful application of new science to Large
Blue conservation in UK and across Europe.
6. CLIMIT Report to the EU, held on file at Oxford University and UFZ
(Helmholtz) Leipzig/Halle. Includes CLIMIT research results already
communicated and applied to conservation policy and practice while
under review or `in prep' for publication in ISI journals.
Funding for research: Grants of €1.4M have been received for this
work since 2007, through the EU FP6 Biodiversa programme CLIMIT (CLimate
change impacts on Insects and their MITigation).
Details of the impact
Since 2008, the new findings that emerged from Professor Thomas' research
at the University of Oxford have had a major beneficial impact on the
global conservation of M. arion. In the UK, where links were
already established, the rapid communication of emerging results provided
the essential foundation for new large-scale reintroduction programmes,
which have made significant progress towards restoring the butterfly's
former (19th century) range. Crucially, the strength of the
scientific underpinning for the conservation programme has made funding
more forthcoming for these efforts 7,8. As a result of the new information
on the precise criteria for optimum M. arion habitat, the
management of all current and potential new UK Large Blue sites has been
modified by the interested conservation bodies (National Trust (NT),
Natural England, Clarke Trust, Network Rail, Somerset Wildlife Trust
(SWT), Gloucestershire Wildlife Trust (GWT), and Butterfly Conservation)
to meet Thomas' specifications7-9. This resulted in the UK
Large Blue population more than doubling from 128,000 eggs in 2007 to
272,000 in 2012, with huge increases on some key sites, e.g. the NT's
Collard Hill, where a previously stabilised population of c.230
adults has grown ten-fold since 20099.
The research into M. arion's adaptation to warmer microclimates
has enabled the design and creation of new areas of high-quality habitat
adjacent to established sites, providing optimum conditions for
meta-population expansion and the evolution of more dispersive races. In
major constructions by Network Rail10, two new Origanum-using
sites have been built from scratch — a level of habitat manipulation never
previously attempted for an insect in any country. These and two other
newly managed Origanum sites supported around 20% of the UK Large
Blue numbers in 2012-13. In addition, management elsewhere (by Natural
England, SWT, GWT and the NT) has been modified to extend the breeding
area on eight Thymus-using sites to encompass Origanum.
This has enlarged the available habitat for Large Blues (and hence
increasing population size) and, equally importantly has dampened the
fluctuations of populations in years of extreme weather7,8.
Selection for a race of Large Blue adapted to use Origanum
anticipates a time in coming decades when all sites supporting Thymus
are predicted to become too hot to support the ant host, M. sabuleti,
and hence M. arion.
The experimental selection6,10 for early-emerging races
adapted to thyme-flowering on cooler sites was crucial to the
reintroduction of M. arion to the Cotswold hills, where all
previous attempts at reintroduction in the 1980s and 1990s had failed. It
also showed that the sites selected previously had microclimates too cool
for the UK populations. As a result, Natural England, GWT and the NT
switched management to sites with warmer terrains and used the
early-emerging races for new introductions in 2010. These coincide
precisely with the flowering of local thyme, and the butterfly already
thrives on two sites8. This provided the rationale for an
ambitious restoration programme (led by Butterfly Conservation and Natural
England, funded by Biffa through landfill taxes) on a further nine
Cotswold sites7. Two more introductions were made in 2013, and
it is predicted that the butterfly will now spread across the landscape
after an absence of 50 years.
Within landscapes, Thomas' research on adult Large Blue dispersal has
guided conservation bodies' decisions about which new sites to develop.
For example, it prompted the NT and Clarke Trust in 2009 to embark on
creating a new visitor site within range of the existing one at Collard
Hill7. By clearing secondary woodland from a potentially
suitable hillside and imposing a targeted grazing regime the site was
colonised by the butterfly within four years7. The research
also provided the rationale for a consortium of conservation bodies, led
by Butterfly Conservation, to bid successfully for a SITA (landfill) award
in 2011-14. The award funded the restoration of former sites, and the
creation of new ones in the West Polden Hills, adjacent to the East
Poldens landscape where the butterfly already thrives7.
Thomas' demonstration that targeted management of a single species leads
to collateral gains in other declining wildlife enabled conservation
bodies to argue that management for the Large Blue would benefit entire
communities of threatened species, attracting funding that would not
otherwise be forthcoming. Thus, the NT have imposed `Large Blue
management' on four Dartmoor sites, with the prime aim now being to
promote habitat for four declining species of violet-feeding fritillary
butterfly (capitalising on Thomas' discovery that the Pale Heath Violet
thrives in M. arion habitat due to seed dispersal by Myrmica
ants). This strategy has been highly successful9, with some of
the largest known UK populations of High Brown and Pearl-bordered
Fritillary breeding in 2012-13 on all four sites. In comparison, both
experienced c.80% extinctions elsewhere, making the High Brown the
most endangered UK butterfly species. These sites have also been colonised
by other UK Biodiversity Action Plan species, including other butterflies
and rare insects7, 9. A new programme is currently underway on
the Atlantic coast of Cornwall, with the same aim9.
Since 2008, grant funding for the development of new Large Blue sites has
totalled over £200,000, excluding local economic impacts (£13,000 to
nurseries to propagate thyme plants; £50,000 to contractors to undertake
practical management and surveys). The new programmes have resulted in
significant local engagement of the community (for example involving
volunteers in programmes clearing scrub, planting thyme and monitoring
butterflies). The support of major national charities such as the NT,
Natural England, Wildlife Trusts and Butterfly Conservation underlines the
breadth of impact of Thomas' research. There are today more than 100 sites
across the UK being carefully prepared for M. arion - at least
double the number in 2008 — of which 40 already support the butterfly,
making the UK a European and global stronghold of this iconic species.
Indeed, the Large Blue is one of just three UK butterflies to meet the
Convention of Biological Diversity's target to reverse species' declines
by 201011,12. Notably, the successful conservation of the other
two also derives directly from Thomas' research4,12. Natural
England states that Oxford based research `has provided essential
knowledge that has enabled Natural England to fulfil its statutory
duties in reversing the decline of the EU Habitats Directive Species
Maculinea arion'8. The impact has been felt across Europe too,
with several M. arion (and other species') sites in other nations
having implemented Thomas' findings and achieved significant growth in
numbers10. As a consequence, M. arion's global IUCN
listing has changed from `Vulnerable to `Near-threatened'.
The NT's site at Collard Hill (with a busy web-site and blog) now
attracts over 1000 visitors in June-July when Large Blues are on the wing,
making it one of NT's prime public sites for wildlife watching8,13.
Widespread media coverage of the reintroduction of the large blue
butterfly has reached a very large audience within the UK via a range of
media, including BBC's The One Show (4.48 million viewers) and Countryfile
(over 6 million viewers) in 2012, documentaries on local and national TV
and radio (e.g. Radio 4's Nature August 2009 and Saving
Species 201014), and many articles in the national
Sources to corroborate the impact
- Letter from the Chief Executive of Butterfly Conservation UK and Vice
Chair of Butterfly Conservation Europe (held on file), confirming
changes in site management, management of sites to mitigate the
effects of climate change, creation of sites to take advantage of
more mobile races of Large Blue, and gains in other threatened
- Letter from the Land Management Senior Adviser for the Cotswolds at
Natural England (held on file), confirming changes in site
management, management of sites to mitigate the effects of climate
change, establishment of new sites in the Cotswolds and ability to
fulfil its statutory duties under the EU Habitats Directive.
- Letter from the Wildlife Countryside Adviser for the National Trust
South West Region (held on file), confirming changes in site
management, increases in butterfly numbers, creation of sites to
take advantage of more mobile races of Large Blue, gains in other
threatened species as a result of `Large Blue management', and
greatly increased visitor numbers and public interest.
EU Biodiversa Policy Brief `Conservation of Threatened Insects in
Europe: Managing Habitats for land-use and Climate Change adaptation',
compiled in 2013 by IUCN from CLIMIT (largely UK M. arion)
research results and confirming increases in Large Blue numbers
elsewhere in Europe as a result of Thomas' research.
Double page article from the Guardian, confirming an increase in
Large Blue numbers against a general backdrop of plummeting
butterfly populations in the UK.
2009 Fourth UK National Report to the United Nations Convention on
Biological Diversity; confirms (p25) the increase in Large Blue numbers.
- National Trust Collard Hill: http://ntlargeblue.wordpress.com
Links to a popular blog about this visitor site.
Links to two Radio 4 broadcasts on the Large Blue.