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Monitoring of Odonata in Britain and possible insights into climate change 127
Monitoring of Odonata in Britain and possible
insights into climate change
Adrian J. Parr
10 Orchard Way, Barrow, Bury St Edmunds, Su olk IP29 5BX, Great Britain
Corresponding author: Adrian J. Parr (adrian.parr@btinternet.com)
Academic editor: Jürgen Ott|Received29 July 2010|Accepted 20 August 2010|Published 30 December2010
Citation: Parr AJ (2010) Monitoring of Odonata in Britain and possible insights into climate change. In: Ott J (Ed)
(2010) Monitoring Climatic Change With Dragon ies. BioRisk 5: 127–139. doi: 10.3897/biorisk.5.846
Abstract
e history of recording and monitoring of Odonata in Britain is brie y described. Results are then pre-
sented which suggest that the country’s Odonata fauna is currently in a period of ux, in a manner consist-
ent with the actions of a high-level regulatory factor such as climate change. e ranges of many resident
species are shifting. Leucorrhinia dubia has recently been lost from southern England, but many species are
presently expanding their ranges to the north and west, some (such as Aeshna mixta and Anax imperator)
with considerable speed. In addition to these changes, a number of ‘southern’ species have started to ap-
pear in Britain for the very rst time. ese include Lestes barbarus, Erythromma viridulum (which has now
become a locally-common resident in southeast England), Anax parthenope and Crocothemis erythraea. In
addition to these distributional changes, some recent trends in ight times are also discussed. Evidence
indicates that many species are now emerging signi cantly earlier than in the past, though trends relating
to the end of the ight period are less clear cut.
Keywords
Climate Change, Odonata, dragon ies, monitoring, distribution, range expansion, 'Mediterranean spe-
cies', migration, phenology, emergence dates
Introduction
e study of Odonata in the British Isles goes back a considerable time; several publi-
cations devoted at least in part to dragon ies appeared as early as the end of the 18th
century. ese included “An exposition of English insects” (Harris 1782) and “ e
natural history of British insects” (Donovan 1792–1813). Interest in Odonata was
BioRisk 5: 127–139 (2010)
doi: 10.3897/biorisk.5.846
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RESEARCH ARTICLE
BioRisk
A peer-reviewed open-access journal
Adrian J. Parr / BioRisk 5: 127–139 (2010)
128
however never as great as that for groups such as birds, owers or Lepidoptera, and it
was not until the time of W.J. Lucas that much in the way of centralised distributional
data became available. In his key book (Lucas 1900), Lucas listed the then known lo-
calities for each species on a county basis. Cynthia Long eld (Long eld 1937, 1949),
later assisted by Norman Moore and Philip Corbet (Corbet et al. 1960) continued to
update the available distributional data. Structured monitoring of dragon ies within
the UK was nally formalised in 1967, with the establishment of the Odonata Record-
ing Scheme under the auspices of the national Biological Records Centre at Monks
Wood, and the rst high-detail distribution maps started to become available (e.g.
Hammond 1977, 1983). ese events partly re ected, but to an even greater extent
induced, a substantial increase in the popular observation of dragon ies - a trend that
still continues today. In 1983 the British Dragon y Society (BDS) was formed, and
after a period of intense eld observations a national Atlas (Merritt et al. 1996) was
produced, dealing with observations up to, and a little beyond, 1990. At about this
time administration of the Odonata Recording Scheme was formally devolved to the
BDS, but the last year or two has then seen something of an extension into a more
centralised organisation with the development of the National Biodiversity Network
(NBN). is aims to facilitate the exchange of biological information of many sorts by
making a wide range of biological records readily available to all over the internet (see
the NBN Gateway at http://www.searchnbn.net/). e BDS has been closely involved
with this scheme from the start.
Clearly there is a long history of recording Odonata in the British Isles, and al-
though the volume of the older data is perhaps not as great as that for some other
biological groups it still provides much useful background information. Coupled to
the well-developed modern recording and monitoring scheme, su cient data is now
available to make it relatively easy to detect changes to the region’s dragon y fauna and
allow some investigation of the underlying causes. Indeed, with the current BDS mem-
bership standing at roughly 1500 (although admittedly not all will be highly active in
the eld), Britain must be one of the most intensively-recorded countries in Europe,
if not the world, as far as Odonata are concerned. In this article many recent develop-
ments are described and the possible role of climate change is discussed. In Britain,
climate change, whatever its underlying cause, would appear to be very much a reality,
with the standardized annual mean Central England Temperature (Manley 1974) hav-
ing exceeded the 1961–1990 average during nineteen of the last twenty years (Hadley
Centre 2008). In England, six of the seven warmest years on record (going back as far
as 1659) have also occurred since 1995 (Hadley Centre 2008). Currently, the averaged
CET anomaly stands at very nearly +1.0°C (Hadley Centre 2008).
Species recorded from the British Isles
One of the most obvious ways in which a country’s Odonata fauna can be a ected by
climatic conditions is by the direct in uence on what species can maintain a presence.
Monitoring of Odonata in Britain and possible insights into climate change 129
ree species have become extinct in Britain during the last 60 years, viz. Coenagrion
armatum (last seen 1958; Brownett 2005), C. scitulum (last seen 1952, Merritt et al.
1996), and Oxygastra curtisii (last seen 1963, Merritt et al. 1996). All three were at the
then limit of their European range, and had extremely localised distributions even at
their peak, so that they all eventually succumbed to habitat degradation of one form or
another. ese were thus probably ‘chance’ events. It is however worth noting that C.
scitulum was only ever recorded during the period 1946–1952, and it seems possible
that it was only a transient resident, initially colonising during the good migrant years
of the 1940s (Long eld 1949; Parr 1996) that also saw a number of unusual records
from the Channel Isles (Silsby and Silsby 1988).
With the exception of the extinctions listed above, the British Odonata fauna
remained relatively stable for an extended period of time. As with many other coun-
tries in northern Europe, the last few years have however seen the arrival of a sub-
stantial number of new species, many of them having their strongholds in the Medi-
terranean region. Erythromma viridulum was rst observed in 1999 and colonised
rapidly, with it now being a locally-common resident in SE England (Parr 2004,
2005). Lestes barbarus was rst observed at a single site in Norfolk during 2002,
with records from there and at another site in coastal SE England during 2003 and
2004. Despite thorough searches, the species was not however noted at these sites
in subsequent years, although a single individual was found at a third, inland, site
in 2006 (Parr 2007). As far as Anisoptera are concerned, there was an uncon rmed
report of Anax parthenope in the mid 1980s, though it had to wait until 1996 for a
substantiated record to be forthcoming. Since then the species has been seen annu-
ally, with several hundreds of individuals now having been recorded; during 2000
one even reached as far north as 59°17'N (Parr et al. 2004). Most sightings of A. par-
thenope currently seem to refer to migrants, but successful breeding has been proven
at two sites in the southwest (Jones 2000), and has been strongly suspected from
several further areas. Con rming that the situation is still evolving, during 2006
record numbers were seen, with ovipositing being reported from at least 5 sites (Parr
2007). Other dragon ies added to the British List in recent years include Sympetrum
pedemontanum - with one record in 1995 and another ‘possible’ seen by a member of
the public in 2003 (Parr 2004) - and Anax junius, where the rst European records
of this Nearctic species were made in southwest England during autumn 1998 (Pel-
low 1999). In addition, Crocothemis erythraea has also started appearing in Britain in
recent years, with 6 records in England since the rst sighting in 1995. ese records
principally refer to single males seen near the south coast (no females have as yet
been de nitely observed), though there was an unexpected sighting of a sub-mature
male in Cumbria, at very nearly 55°N, during 2004 (Parr 2005). Finally, although
not strictly directly related to the British mainland, it is worth noting that Orthetrum
brunneum was recorded for the rst time ever from the Channel Isles (on Guernsey)
during 2001 (Parr 2002).
In addition to the occurrence of new species, the frequency of appearance of cer-
tain more ‘normal’ migrant species also appears to be changing. Sympetrum fonsco-
Adrian J. Parr / BioRisk 5: 127–139 (2010)
130
lombii was for many years regarded as only an erratic visitor to British shores, often
apparently being absent for several years in a row (Long eld 1949). In the late 1980s
and early 1990s fairly regular records started to come from southwest England, and the
species is currently regarded as an expected and often not-uncommon migrant to Brit-
ain, with major in uxes having been noted in 1996, 1998, 2000, 2002 and particularly
2006 (Parr 2003, 2007). Successful breeding is now also regularly reported (e.g. Parr
2002, 2004, 2007). Although the recent increase in numbers of active recorders must
complicate interpretation, observed changes do seem to be real. us the related S. a-
veolum, a migrant of more easterly rather than southerly origin, shows a very di erent
trend to its pattern of records (Parr 1996, 2003).
Range expansions and contractions for resident species
One of the advantages of the relatively long history of dragon y monitoring in Brit-
ain, and its focus on distributional aspects, is that it is now feasible to sensibly analyse
range changes of the resident species. As with other countries in northern Europe, it
is clear that many species are currently undergoing signi cant changes to their range.
ese are mostly expansions, typically to the north and west, but a few species do
seem to be contracting their range. Hickling et al. (2005) have provided an overview
of changes seen over the period 1960–1995, but there are problems in interpreting
some of the data and events have also progressed since the time period they cover. It
may thus be worthwhile describing further certain species for which range changes
are being seen:
Calopteryx splendens. ere is evidence for a recent strengthening and range ex-
pansion of the more northerly sub-populations in Britain (Clarke 1999; Ward and
Mill 2004), especially since 1990. e role of possible improvements in water quality
however remains to be evaluated.
Brachytron pratense. is species declined or disappeared in several areas during
the post-World War II period, perhaps in part due to pollution and changes in land
use associated with an intensi cation of agricultural practices. A recovery described
by Merritt et al. (1996) and Perrin (1999) has continued, and there have been records
from many new areas in recent years. To some extent there has simply been an in lling
of range, but in addition there seems to have been an extension into western parts of
Wales. e species has also been recorded from a few southern and western areas of
Scotland since 1984 (Smith and Smith 1984; Merritt et al. 1996), though may per-
haps have been overlooked there previously, since Sélys (1846) reported being shown
a specimen from the region.
Aeshna mixta. In older times this species was essentially a scarce immigrant to
Britain, though there is circumstantial evidence that it may have been temporarily es-
tablished in East Anglia at the end of the 19th century (Mendel 1992). A large invasion
during 1935 saw records from several areas of southeastern England, and somewhere
around this time it apparently started to become increasingly established as a resident
Monitoring of Odonata in Britain and possible insights into climate change 131
(Mendel 1992). Merritt et al. (1996) describe the gradual expansion of the species
throughout southeast and central England up until 1990. Since then range expansion
has continued, and seems to be increasing in rate. e species reached Ireland in small
numbers during 2000 (Nelson and ompson 2004), and is now established there,
while the rst con rmed records for Scotland were in 2004, with reports from at least
three sites, one of which was well to the north (Parr 2005).
Aeshna isoceles. In the 19th century this species was found in a few widely-scat-
tered fenland localities throughout East Anglia, but became restricted to the Broadland
area of Norfolk and the immediately adjacent part of Su olk as a result of habitat loss
(Merritt et al. 1996). is highly localised distribution warranted the species being
a orded legal protection under the Wildlife and Countryside Act 1981. In the 1990s,
individuals began to be seen on the Su olk coastal strip, and ovipositing has been ob-
served here in the last few years. Although these sites are not a great distance from prior
strongholds (up to 30 km), the move into a new habitat is of some signi cance for such
a rare and localised species in Britain. Interestingly, while at most of its Norfolk sites
the species is apparently associated with the presence of Water Soldier Stratiodes aloides,
this is not the case in coastal Su olk. e relationship between this range expansion
and the occasional record on the east coast of what might be immigrants of Continen-
tal origin (Parr 2004) remains to be established.
Anax imperator. As with Aeshna mixta, this is another common species that is
now rapidly expanding its range (see Fig. 1A). Before 1982 there were very few re-
cords north of 52°55'N, and those mostly referred just to isolated individuals. By the
early 1990s the species had however been found in Cheshire and Lancashire, north to
about 53°45'N (Merritt et al. 1996), and good populations now exist in these areas.
By 1995 it had reached Cumbria (Clarke 2004), and by 2003 records for Scotland (at
ca. 55°00'N) were forthcoming (Parr 2004). It was rst seen in Ireland during 2000
(Nelson and ompson 2004), and now appears to be established there.
Libellula depressa. Long established in southern Britain, this species has been
increasingly noted further north in recent years. e northern limit of its range was at
roughly 53°30'N in 1990 (Merritt et al. 1996), but a wanderer or migrant reached very
nearly 56°N in Scotland during 2003 (Parr 2004).
Libellula fulva. is species has always been very local in Britain, where it has
the vernacular name of Scarce Chaser. Merritt et al. (1996) describe a distribution
centred on roughly six scattered river systems and nearby still-water sites in southern
and southeast England. e last few years have however seen a number of new areas
being discovered (Parr 2005, 2006), including records from an additional four coun-
ties. Some of these new sites are up to 100 km away from previously known sites, and
extend the overall range to the west and northwest.
Orthetrum cancellatum. is was once at best a rare and localised species in
southern England, but in the 20th century O. cancellatum became increasingly more
widespread and common, perhaps in part due to the spread of ooded gravel workings,
which provide excellent habitat (Merritt et al. 1996). Range expansion in both Britain
and Ireland has since continued, and a hot weather-related movement during late July/
Adrian J. Parr / BioRisk 5: 127–139 (2010)
132
early August 2006 saw the most northerly British records to date, including one from
ca. 56°15'N in south-central Scotland (Parr 2007).
Sympetrum sanguineum. Speculation during the 1970s that this species might
have been in decline in eastern England was probably exaggerated, earlier records hav-
ing been biased somewhat by a period of good immigration in the 1930s and 1940s
(Merritt et al. 1996). By the late 1980s it was clear that the species was in fact doing
well in several areas, and recently the species has been expanding its range rapidly to
the north (Fig. 1B). From a northern limit of ca. 53°
30'N in 1982, the species has
now been recorded right up to the Scottish border, beyond 55°
30'N. Curiously, it still
remains rather rare in the far southwest of Britain.
Leucorrhinia dubia. Although the British stronghold of this species has always
been in Scotland, there are (or, in some cases, were) outposts in the north of England,
the northwest Midlands and the Surrey heaths of southern England. In Surrey the spe-
cies had been recorded over the years from at least three 10 × 10 km grid squares
(Mer-
ritt et al. 1996). is southern population is however now extinct, the last individual
having been noted at ursley Common during 1999 (Perrin 2000). Certain other
sub-populations have also experienced a decline, in part due to habitat loss (Merritt et
al. 1996), though fortunately in places the species continues to do satisfactorily.
Figure 1. Distribution maps for A Anax imperator and B Sympetrum sanguineum. ● First recorded before
1980, ● First recorded 1981–1990, ○ First recorded 1991–2004.
Monitoring of Odonata in Britain and possible insights into climate change 133
Phenology
e British Odonata database now contains nearly 500,000 records, which in addition
to allowing the monitoring of distribution, will also contain extensive phenological
information. Basic summary data and a preliminary analysis of the e ects of latitude
and year-to-year variation on phenology have been available for some while (Merritt
et al. 1996). More recently, the appointment of a salaried BDS sta -member to work
with the database (to ensure that its structure is compatible with the evolving National
Biodiversity Network and to extract key data) has facilitated a more extensive analysis
of phenological data. It is clear that since 1960 there has been a signi cant shift in the
phenology of the British Odonata as a whole, with species experiencing a continuing
advance in the leading edge of the ight period - i.e. they are starting to emerge earlier
(Hassall et al. 2007). is correlates strongly with changes in British climate during
the same period. On average the forward shift amounts to 1.5 days per decade, though
there are indications that spring-emerging species are more strongly a ected than those
that emerge in summer (Hassall et al. 2007). No general trends relating to the end of
the ight period were apparent, perhaps because di erent species respond di erently
(Hassall et al. 2007).
In addition to the detailed statistical analysis of the British Odonata database,
other approaches to monitoring phenology are also available. For over 15 years, many
BDS members have been routinely recording the dates of the rst and last sightings of
di erent species in their area, and a database of national ‘ rst’ and ‘lasts’ has been estab-
lished. Such data has its limitations, but it does have an immediacy that has proved at-
tractive, both for encouraging recorders and for attracting public attention. Calopteryx
splendens is a useful species to examine in detail. Figure 2 illustrates changes in the dates
of reported rst and last sightings in Britain over recent years. Although there are a few
historic records that are similar to recent extremes, there is a clear trend for the species
to now be seen earlier in spring and later into the autumn than before. Indeed in 2005
individuals were recorded between 28 March and 10 October, which is unprecedented
for Britain. Detailed analysis of the data is di cult as extreme dates are by de nition
somewhat atypical, and they are also sensitive to recording e ort, with the increasing
number of recorders in recent years being more likely to pick up individual rare events.
C. splendens is however common and relatively large and showy, and is thus readily
spotted. It is also easy to identify from a distance. First and last records are therefore
less likely to be in uenced by observer e ects than those for many other species, and
it would seem likely that there is indeed an increasing disturbance to what was consid-
ered the normal phenology, with both unusually early emergences and either increased
adult survival or unusually late emergences now taking place.
Although the analysis of trends in rst dates can be problematic, there is one situ-
ation where interpretation is easier, namely when one is comparing observations at a
single site that has been regularly monitored over many years. Chartley Moss National
Nature Reserve, Sta ordshire, is one of the major sites for Leucorrhinia dubia in Eng-
land, and the ight period and numbers of individuals seen each year has been closely
Adrian J. Parr / BioRisk 5: 127–139 (2010)
134
Figure 2. Yearly A ‘ rst’ and B ‘last’ dates for Calopteryx splendens in Britain during 1991–2005.
monitored for two decades. Figure 3A plots the recorded ‘ rst dates’ for L. dubia during
this period; there is a statistically signi cant (p<0.05) trend towards earlier emergences
in recent years. Although meteorological data for Chartley Moss itself is not readily
available, it lies almost midway between the weather stations of Shawbury and Sutton
Bonnington, for which extensive historical data sets exist (Met O ce 2006). Analysis
shows that over the period of observations at Chartley, both weather stations showed a
slight trend towards increases in spring temperature, but this was small in comparison
to yearly variation, and was not statistically signi cant. In itself, such a trend cannot
therefore account for the phenological changes. More obvious was an increase in mean
annual temperature, with both the mean annual maximum temperature (Tmax) and the
mean annual minimum temperature (Tmin) increasing with time. Over the period in
question, Tmax showed a mean linear increase of ca. +0.06°C per year (p<0.05 at Sutton
Bonnington, p<0.01 at Shawbury - see Fig. 3B), while Tmin showed a mean linear in-
Monitoring of Odonata in Britain and possible insights into climate change 135
crease of ca. +0.05°C per year (p<0.05). ese trends may well account for the observed
shifts in spring emergences. In contrast to the results for rst dates, the data for late
dates was more scattered, principally due to sporadic very late sightings (the latest being
12 October) apparently resulting from unexpectedly late emergences in certain seasons.
is could sometimes be linked to poor weather in the early part of the season, but this
was not always the case, and perhaps these observations are also of some signi cance.
In addition to the situation with L. dubia alluded to above, other instances of
autumn emergences that seem at odds with accepted life-cycle strategies in northern
Europe have been documented in recent years. ese include September emergences
of Anax imperator (Parr 1999), the larvae of which are normally considered to have an
Figure 3. Phenological trends and climate: A ‘First dates’ for Leucorrhinia dubia at Chartley Moss NNR,
Sta ordshire, England, during the period 1987–2005. In most years the site was visited at 2–3 day inter-
vals during the appropriate time period. In 1991–1994 it was only possible to visit weekly, and to avoid
bias towards later dates, ‘ rst dates’ are here taken as midway between the visit on which L. dubia was rst
recorded and the preceding visit. B Mean annual maximum temperatures at Shawbury Weather Station.
Adrian J. Parr / BioRisk 5: 127–139 (2010)
136
autumn diapause to ensure synchronous emergence the following spring (Corbet et
al. 1960). In this particular instance the presence of an immigrant population with a
di erent phenology might be one explanation, but further investigation of unexpected
autumn emergences might help throw light on changing phenologies in the region.
Although such emergences are still rare events, it is, for example, not inconceivable that
certain species are starting to show partial changes in voltinism.
Conclusions
It is clear that the British Odonata fauna is currently going through a period of con-
siderable ux:
– Several resident species are currently expanding their ranges, though a few are
in regression. ose expanding their range are often species of ‘southern’ a nity, and
are expanding to the north and west, some with considerable speed. e ‘northern’
Leucorrhinia dubia is however contracting its range, having recently become extinct in
its southern English outpost.
– Species never before observed in the British Isles are now starting to turn up in
the region as they expand their ranges. Again, it is principally those somewhat ther-
mophilic species with a ‘southern’ distribution in Europe that are involved, although
there are exceptions (e.g. the more eastern Sympetrum pedemontanum and the Ameri-
can Anax junius).
– ere is evidence for shifts in phenology, with earlier emergences in spring and at
least for some species the suggestion of unusual late season records as well.
In addition to the Odonata, changes are also being observed in the UK for a very
wide range of other taxa. Particularly in the case of birds, changes in range and phe-
nology have already been extensively documented (e.g. omas and Lennon 1999;
Musgrove 2002; Cotton 2003; Eaton et al. 2005). Within the Insect order, it is clear
that in addition to dragon ies, several butter ies with a southerly distribution in the
UK are also expanding their ranges, though there are signs that habitat limitation may
be restricting the number of species involved (Warren et al. 2001). e spread of the
warmth-loving sphecid wasp Philanthus triangulum has similarly attracted attention
(Edwards 2000), as has the spread of the Orthopterans Metrioptera roeselii and Cono-
cephalus discolor ( omas et al. 2001). is latter situation is particularly interesting,
with the dramatic north-westerly range expansions seen in the last 20 years being fa-
cilitated by the increased occurrence of long-winged forms with enhanced dispersive
capabilities ( omas et al. 2001). In addition to these range changes, a wide variety of
insect species have recently been recorded in Britain for the very rst time. ese are
too numerous to list fully, but include various moths such as aumetopoea processio-
nea, Ochropleura leucogaster, Dryobota labecula, Spodoptera cilium, Platyperigea kadenii
and Pechipogo plumigeralis (Waring and Townsend 2003; Clancy 2006), the bee Bom-
bus hypnorum (Goulson and Williams 2001), the social wasps Dolichovespula saxonica
and D. media (Edwards 2000), a good variety of Coleoptera including the coccinellids
Monitoring of Odonata in Britain and possible insights into climate change 137
Rhyzobius chrysomeloides, Epilachna argus and Harmonia axyridis (Biological Records
Centre 2008), and also the hemipteran Nezara viridula (Shardlow and Taylor 2004).
Although some of these records may represent accidental introductions, many species
appear to have reached Britain unaided, in an extension of wider trends.
Since changes being seen in the UK are qualitative as well as quantitative, they can-
not simply be artifacts of the increased interest in Odonata and many other areas of wild-
life that has taken place in recent years. Rather, the types of change, and their widespread
and consistent nature, seem to imply that some high-level controlling variable such as
climate is involved, though of course this might operate in a variety of way (e.g. either di-
rectly or via e ects on things such as habitat quality). Although it is di cult to be precise,
recent changes seem to have started in earnest during the 1980s. e early 1980s roughly
coincide with the onset (or, more strictly, intensi cation) of a signi cant and continu-
ing increase in British, Northern Hemisphere and Global mean temperature (Jones &
Moberg 2003; Climate Research Unit, UEA 2008; Hadley Centre 2008). Indeed, for
Odonata, the closest historical situation in Britain to that which is currently taking place
appears to be the good migrant years of the 1930s and particularly the 1940s (Long eld
1949; Parr 1996). is period was also associated with a phase of steeply rising tempera-
ture, though in this case it was not sustained (Jones and Moberg 2003; Hadley Centre
2008). Many current trends would thus appear likely to be linked to climate change.
It will be of considerable interest to continue monitoring changes to the British
Odonata in years to come, and the BDS intends to strengthen its resident and migrant
recording schemes, develop more detailed studies of dragon y abundance, and also
consolidate its studies on phenology.
Acknowledgements
e author wishes to thank Graham French, the then BDS Key Sites O cer, for help
in producing the distribution maps shown here.
References
Biological Records Centre (2008) UK Ladybird Survey. http://www.ladybird-survey.org/de-
fault.htm
Brownett A (2005) A re-examination of the status of the Norfolk Damsel y Coenagrion arma-
tum (Charpentier): a species of Odonata now presumed extinct in Britain. J. Br. Dragon y
Soc. 21: 21–26.
Clancy S (2006) Occurrences of the rarer immigrant moths in 2005. Atropos 27: 24–31.
Clarke D (1999) e outpost populations of the Banded Demoiselle Calopteryx splendens (Har-
ris) in the Solway Firth area, Cumbria: historical perspective and recent developments. J.
Br. Dragon y Soc. 15: 33–38.
Clarke D (2004) ‘Southern’ dragon ies make headway in Cumbria in 2003. Darter 21: 6–7.
Adrian J. Parr / BioRisk 5: 127–139 (2010)
138
Cotton PA (2003) Avian migration phenology and global climate change. Proc. Natl. Acad. Sci.
USA 100: 12219–12222.
Donovan E (1792–1813) e natural history of British insects (in 16 volumes). F. & C. Riv-
ington, London.
Climate Research Unit, University of East Anglia (2008) Data: temperature: hemispheric and
global averages. http://www.cru.uea.ac.uk/cru/data/temperature.
Corbet PS, Long eld C, Moore NW (1960) Dragon ies. Collins, London, 260pp.
Eaton MA, Noble DG, Hearn RD, Grice PV, Gregory RD, Wotton S, Ratcli e N, Hilton
GM, Reh sch MM, Crick HQP, Hughes J (2005) e state of the UK’s birds 2004. BTO,
RSPB, WWT, CCW, EN, EHS and SNH, Sandy, 28pp.
Edwards M (2000) BWARS: mapping the distribution of British Aculeates, and much more.
National Federation for Biological Recording Newsletter 26.
Goulson D, Williams PH (2001) Bombus hypnorum (Hymenoptera: Apidae), a new British
bumblebee? Br. J. Ent. Nat. Hist. 14: 129–131.
Hadley Centre for Climate Prediction and Research (2008) Mean Central England Tempera-
ture, annual anomalies, 1772–2007. http://hadobs.meto ce.com/hadcet/
Hammond CO (1977) e dragon ies of Great Britain and Ireland. Curwen Press, London,
115pp.
Hammond CO (1983) e dragon ies of Great Britain and Ireland. 2nd edition, revised R.
Merritt. Harley Books, Colchester, 116pp.
Harris M (1782) An exposition of English insects. White & Robson, London, 166pp.
Hassall C, ompson DJ, French GC, Harvey IF (2007) Historical changes in the phenology
of British Odonata are related to climate. Global Change Biology 13: 933–941.
Hickling R, Roy DB, Hill JK, omas CD (2005) A northward shift of range margins in Brit-
ish Odonata. Global Change Biology 11: 502–506.
Jones PD, Moberg A (2003) Hemispheric and large-scale surface air temperature variations: An
extensive revision and an update to 2001. J. Climate 16: 206–223.
Jones SP (2000) First proof of successful breeding by the Lesser Emperor Anax parthenope
(Sélys) in Britain. J. Br. Dragon y Soc. 16: 20–23.
Long eld C (1937) e dragon ies of the British Isles. F. Warne and Co., London, 220pp.
Long eld C (1949) e dragon ies of the British Isles; second edition. F. Warne and Co.,
London, 255pp.
Lucas WJ (1900) British Dragon ies (Odonata). L. Upcott Gill, London, 356pp.
Manley G (1974) Central England Temperatures: monthly means 1659 to 1973. Quarterly
Journal of the Royal Meteorological Society 100: 389–405.
Mendel H (1992) Su olk dragon ies. Su olk Naturalists’ Society, Ipswich, 159pp.
Merritt R, Moore NW, Eversham BC (1996) Atlas of the dragon ies of Britain and Ireland.
HMSO, London, 149pp.
Met O ce (2006) Past Weather: UK: Historic station data. http://www.meto ce.gov.uk/cli-
mate/uk/stationdata/index.html
Musgrove AJ (2002) e non-breeding status of the Little Egret in Britain. British Birds 95: 62–80.
Nelson B, ompson R (2004) e Natural History of Ireland’s Dragon ies. e National
Museums and Galleries of Northern Ireland, Belfast, 454pp.
Monitoring of Odonata in Britain and possible insights into climate change 139
Parr AJ (1996) Dragon y movement and migration in Britain and Ireland. J. Br. Dragon y
Soc. 12: 33–50.
Parr AJ (1999) Late season records of Emperor Dragon y Anax imperator. Atropos 6: 33–34.
Parr AJ (2002) Migrant and dispersive dragon ies in Britain during 2001. J. Br. Dragon y Soc.
18: 40–45.
Parr AJ (2003) Migrant and dispersive dragon ies in Britain during 2002. J. Br. Dragon y Soc.
19: 8–14.
Parr AJ (2004) Migrant and dispersive dragon ies in Britain during 2003. J. Br. Dragon y Soc.
20: 42–50.
Parr AJ (2005) Migrant and dispersive dragon ies in Britain during 2004. J. Br. Dragon y Soc.
21: 14–20.
Parr AJ (2006) Migrant dragon ies in 2005. Atropos 27: 33–38.
Parr AJ (2007) Migrant and dispersive dragon ies in Britain during 2006. J. Br. Dragon y Soc.
23: 40–51.
Parr AJ, De Knijf G, Wasscher M (2004) Recent appearances of the Lesser Emperor Anax par-
thenope (Sélys) in northwestern Europe. J. Br. Dragon y Soc. 20: 5–16.
Pellow K (1999) An in ux of Green Darner Anax junius (Drury) into Cornwall and the Isles of
Scilly: the rst European records. Atropos 6: 3–7.
Perrin VL (1999) Observations on the distribution, ecology and behaviour of the Hairy Drag-
on y Brachytron pratense (Müller). J. Br. Dragon y Soc. 15: 39–45.
Perrin VL (2000) Dragon y news for 1999 - resident’s round-up. Dragon y News (Newsletter
of the British Dragon y Society) 37: 11–12.
Sélys-Longchamps E de (1846) Revision of the British Libellulidae. - Annals and Magazine of
Natural History 18: 217–227.
Shardlow MEA, Taylor R (2004) Is the Southern green shield bug Nezara viridula (L.) (Hemip-
tera: Pentatomidae) another species colonising Britain due to climate change? Br. J. Ent.
Nat. Hist. 17: 143–146.
Silsby JD, Silsby RI (1988) Dragon ies in Jersey. J. Br. Dragon y Soc. 4: 31–36.
Smith EM, Smith RWJ (1984) Brachytron pratense (Müller) and other Odonata of the Black
Lochs, Argyll. J. Br. Dragon y Soc. 1: 51–54.
omas CD, Lennon JJ (1999) Birds extend their ranges northwards. Nature 399: 213.
omas CD, Bodsworth EJ, Wilson RJ, Simmons AD, Davies ZG, Musche M, Conradt L
(2001) Ecological and evolutionary processes at expanding range margins. Nature 411:
577–581.
Ward L, Mill PJ (2004) Distribution of the Banded Demoiselle Calopteryx splendens (Harris) in
northern England: an example of range expansion? J. Br. Dragon y Soc. 20: 61–69.
Waring P, Townsend M (2003) Field guide to the moths of Great Britain and Ireland. British
Wildlife Publishing, Rotherwick, 432pp.
Warren MS, Hill JK, omas JA, Asher J, Fox R, Huntley B, Roy DB, Telfer MG, Je coate
S, Harding P, Je coate G, Willis SG, Greatorex-Davies JN, Moss D, omas CD (2001)
Rapid responses of British butter ies to opposing forces of climate and habitat change.
Nature 414: 65–69.
