Rapid declines of common, widespread British moths provide evidence of an insect biodiversity crisis

Abstract

A fundamental problem in estimating biodiversity loss is that very little quantitative data are available for insects, which comprise more than two-thirds of terrestrial species. We present national population trends for a species-rich and ecologically diverse insect group: widespread and common macro-moths in Britain. Two-thirds of the 337 species studied have declined over the 35 yr study and 21% (71) of the species declined >30% 10 yr−1. If IUCN (World Conservation Union) criteria are applied at the national scale, these 71 species would be regarded as threatened. The declines are at least as great as those recently reported for British butterflies and exceed those of British birds and vascular plants. These results have important and worrying implications for species such as insectivorous birds and bats, and suggests as-yet undetected declines may be widespread among temperate-zone insects.

Full text

Rapid declines of common, widespread British moths
provide evidence of an insect biodiversity crisis
Kelvin F. Conrad
a,
*, Martin S. Warren
b
, Richard Fox
b
, Mark S. Parsons
a
, Ian P. Woiwod
a
a
Rothamsted Research, Plant and Invertebrate Ecology, West Common, Harpenden, Hertfordshire AL5 2JQ, UK
b
Butterfly Conservation, Manor Yard, East Lulworth, Wareham, Dorset BH20 5QP, UK
ARTICLE INFO
Article history:
Received 23 November 2005
Received in revised form
31 March 2006
Accepted 14 April 2006
Available online 30 June 2006
Keywords:
Biodiversity
Population trends
Population dynamics
Abundance
Occupancy
Lepidoptera
ABSTRACT
A fundamental problem in estimating biodiversity loss is that very little quantitative data
are available for insects, which comprise more than two-thirds of terrestrial species. We
present national population trends for a species-rich and ecologically diverse insect group:
widespread and common macro-moths in Britain. Two-thirds of the 337 species studied
have declined over the 35 yr study and 21% (71) of the species declined >30% 10 yr
1
. If IUCN
(World Conservation Union) criteria are applied at the national scale, these 71 species
would be regarded as threatened. The declines are at least as great as those recently
reported for British butterflies and exceed those of British birds and vascular plants. These
results have important and worrying implications for species such as insectivorous birds
and bats, and suggests as-yet undetected declines may be widespread among temperate-
zone insects.
Ó2006 Elsevier Ltd. All rights reserved.
1. Introduction
Insects are a vital component of terrestrial ecosystems and
form a substantial proportion of terrestrial biodiversity. De-
spite this, knowledge of endangered insects lags behind that
of vertebrates and vascular plants (New, 2004; Thomas
et al., 2004). Whether recent extinction rates of insects are
as great as for other groups has been debated keenly (Thomas
and Morris, 1994; Lawton and May, 1995; McKinney, 1999).
Most early estimates of insect extinction rates were much
lower than those of birds, large mammals and plants, but at-
tempts to quantify losses amongst insects were hampered by
a lack of suitable data (Thomas and Morris, 1994; McKinney,
1999; New, 2004; Thomas et al., 2004).
Recently, Thomas et al. (2004) compared similarly mea-
sured changes in native butterfly, bird, and plant species
and concluded that butterflies had declined more rapidly
than these other groups in Britain; the first time such a com-
parison has been achieved for an insect taxon at the national
scale. They proposed that if other insect groups are similarly
sensitive to recent environmental change, then the unmea-
sured or under-recorded extinction rates of insects may rival
or exceed those documented for vertebrates and plants
(McKinney, 1999; Thomas et al., 2004). Furthermore, Thomas
et al. (2004) argued that such high rates of extinction for in-
sects would signal the ‘sixth great extinction’ (Wilson, 1992).
Here, we report severe national population declines
among another intensively recorded insect group: the larger
British moths, or ‘macro-moths’. Thomas (2005) noted that
long time series of species abundance should provide sensi-
tive indicators of environmental change and cited the British
marco-moths as one of three long-term datasets suitable for
this purpose. In a previous paper (Conrad et al., 2004)
we have described and validated our methodology for
0006-3207/$ - see front matter Ó2006 Elsevier Ltd. All rights reserved.
doi:10.1016/j.biocon.2006.04.020
*Corresponding author: Tel.: +44 1582 763133; fax: +44 1582 760981.
E-mail addresses: kelvin.conrad@bbsrc.ac.uk,conradkf@hotmail.com (K.F. Conrad).
BIOLOGICAL CONSERVATION 132 (2006) 279–291
available at www.sciencedirect.com
journal homepage: www.elsevier.com/locate/biocon

estimating long-term population trends for British macro-
moths and outlined some general patterns in the trends
based on ecological characteristics of the moth species. In
this paper we apply IUCN (IUCN World Conservation Union,
2001) criteria to identify nationally threatened species and
compare macro-moth species declines to those reported
for UK butterflies (Thomas and Clarke, 2004; Thomas,
2005). While the utility of butterflies as indicators of insect
biodiversity has been questioned (Hambler and Speight,
2004; but see Thomas and Clarke, 2004; Thomas, 2005),
moths form a much more ecologically diverse and species-
rich group and are thus more likely to represent a greater
range of terrestrial insects in Britain. We suggest, therefore,
that declines in common and widespread moths provide fur-
ther evidence of wider declines in British terrestrial insects.
2. Methods
2.1. Data source and selection criteria
Population data on British macro-moths were extracted from
the Rothamsted Insect Survey (RIS, Woiwod and Harrington,
1994), one of the longest-running and spatially extensive
datasets of a species-rich insect group anywhere in the world
(Conrad et al., 2004). Established in the early 1960s to provide
information on the spatial variation of insect abundance, the
RIS has operated a national network of approximately 100
standard light-traps (Williams, 1948) annually since 1968.
These traps provide standardized, nightly counts of individ-
ual moth species from a wide range of habitats (Woiwod
and Harrington, 1994; Conrad et al., 2004). Catches are small,
but consistent and representative, making the traps suitable
for long-term monitoring of common and widespread species
without affecting the moth populations being sampled (Wil-
liams, 1952; Taylor and French, 1974; Conrad et al., 2004).
We analysed data for 337 species, each of which was repre-
sented by more than 500 individuals captured over the 35-yr
sampling period (1968–2002), and derived annual national
indices of abundance from the 199 sites that operated for a
minimum of 48 weeks a year for 5 yr (Conrad et al., 2004).
2.2. Estimates of abundance and population change
We estimated indices of annual abundance, allowing for dif-
ferences between sites, by fitting a generalised linear model
with Poisson errors and logarithmic link, using version 3.2
of the TRIM (TRends and Indices for Monitoring data) soft-
ware package (Pannekoek and Van Strien, 2001). By conven-
tion, the estimated abundance in the first year is set to one
and each annual index, A
i
, for year i, is calculated relative to
the first, A
1
.T, the ‘TRIM trend index’ is the overall slope of
the regression of annual indices on a logarithmic scale
(Pannekoek and Van Strien, 2001). Tis a reliable and robust
estimator of long-term trends that is suitable for comparisons
across a range of species (Van Strien et al., 2001; Conrad et al.,
2004). Annual rates of population change were calculated
from Tand 10-yr percentage declines were estimated from
the annual rates of change (Van Strien et al., 2001).
We considered species population decline rates >30%
10 yr
1
to be of significant conservation concern. We further
divided these rapidly declining species into two categories:
vulnerable (30–50% 10 yr
1
) and endangered (>50% 10 yr
1
),
according to the criteria and time period used to identify
globally Vulnerable and Endangered species (IUCN World Con-
servation Union, 2001). Following the guidelines of Gardenfors
et al. (2001), we applied the IUCN thresholds unaltered at the
national level because the British populations can be regarded
as effectively isolated, insular populations and their extinction
risk is unlikely to be affected by populations in continental Eur-
ope (i.e., there is unlikely to be any significant ‘rescue effect’).
2.3. Regional variation
In order to assess geographical variation in population trends
for common macro-moths we divided Great Britain into two
regions along the 4500 N gridline of the British national grid
system. The region to the north of 4500 N was called ‘North’
(N), and the region to the south of 4500 N was called ‘South’
(S). This division into regions was arbitrary but gave a reason-
able number and distribution of sites for analysis in each re-
gion. More importantly, it provides the first steps in
examining a number of species trends for the influences of
climate change and changes in land-use already demon-
strated to affect the decline of the once-common moth, Arctia
caja (Conrad et al., 2002, 2003).
2.4. Comparison of short-term and long-term trap data
While the core number and geographical distribution of traps
never changes significantly from year to year, there has been
turnover of trapping sites during the 35 yr of our study (Con-
rad et al., 2004). In order to examine the effect of this turnover
on our population trend estimates we calculated 10-yr per-
centage population changes using only traps that operated
for 15 or more years and compared the results with those
from our standard ‘all sites’ analysis, which used trapping
sites that had operated for five or more years.
2.5. Light competition
‘Astronomical light pollution’ results from the cumulative ef-
fects of artificial lighting sources increasing the illumination
of the night-time sky (Longcore and Rich, 2004) and may com-
pete with light-traps and decrease their effectiveness. An in-
crease in astronomical light pollution during our study period
could thus decrease trap catches and lead to overestimates of
downward population trends.
To examine the effects of ‘light competition’ on our trap
catches, we obtained ‘world change pair’ images of the
night-time sky from the Defense Meteorological Satellite Pro-
gram Operational Linescan System (DMSP-OLS) dataset, pro-
vided by the US The National Oceanic and Atmospheric
Administration’s (NOAA) National Geophysical Data Centre
(NGDC) (http://dmsp.ngdc.noaa.gov/html/download_world_
change_pair.html). These images provide estimates of aver-
age annual night-time illumination of the earth’s surface for
the years 1992/93 and 2000. Illumination is recorded as pixels
on a linear scale from 0 (dark) to 63 (instrument light satura-
tion) (Elvidge et al., 2001). We selected the 116 RIS light-traps
running between 1992 and 2000, and extracted the night-time
280 BIOLOGICAL CONSERVATION 132 (2006) 279–291

illumination of the 1km
2
pixel containing each trap in 1992/
93 and 2000. We divided the traps into two groups: ‘dark’,
which included 35 trapping sites which scored 0 in 1992/93
and remained 0, or scored >0 in 1992/93 but were darker in
2000, and ‘light’ which comprised 81 sites that were >0 in
1992/93 and were lighter in 2000 (no sites initially >0 remained
unchanged). We then estimated, for each of the two groups,
the annual rate of change in total trap catch of the 337 moth
species in this study for the period 1992–2000.
3. Results
3.1. Rates of change of moth abundance and regional
variation
We found alarming declines in the overall abundance of wide-
spread marco-moths. The annual total number of all macro-
moths caught by the RIS light-trap network decreased by
31% over the 35-yr sampling period (Fig. 1). The majority of
this decrease occurred in southern Britain, while the north
showed no significant trend over time (Fig. 1). Year-to-year
fluctuations in abundance are very similar in both the north
and south despite the difference in overall trends (Fig. 1).
Two-thirds (0.66 ± 0.05, proportion ±95% CI) of the 337 indi-
vidual moth species declined (Fig. 2). The median 10-yr popu-
lation change was a decrease of 12% with a greater median
decrease in the south (17%) than in the north (5%; Fig. 2). Of
even greater concern, 21% (N= 71) of species displayed de-
clines placing them in the vulnerable or endangered categories
(Fig. 2). The total catch of each species and the trend index, T,
were not correlated (r= 0.020, N= 337, P= 0.714; Fig. 3), so the
total number of individuals captured did not affect whether a
species was likely to increase ordecline. Overall, 75% of species
in the south declined compared to 55% in the north (Fig. 2).
3.2. Land-use categories represented
Although the light-trap network originated from an agricul-
tural research station (Woiwod and Harrington, 1994), it was
not intended to monitor agricultural pest species and a wide
range of land-use categories have been sampled (Fig. 4). Be-
cause of trap turnover, the relative numbers of different types
of biotope sampled each year varies over time (Fig. 4). The
mean annual proportions of sites used corresponded with
the following categories: coastal (8.9%); farmland (13.5%);
mixed (15.3%); moorland (3.1%); parkland (22.8%); scrubland
(2.6%); urban (15.9%) and woodland (17.8%). Only the propor-
tion of scrubland changed significantly over time
(F
1,33
= 30.34, P< 0.001), and this is largely because no traps
were sited in areas that were categorised as scrubland in
the early years of the study. Annual variation in biotopes sam-
pled was not systematically biased in any way.
3.3. Comparison of short-term and long-term trap data
Estimates for 222 decreasing species were obtained from sites
that ran 15 or more years. These estimates were highly corre-
lated with those from the ‘all sites’ analysis (r= 0.95, N= 222,
P= <0.001), suggesting that light-trap turnover did not bias
the results. Using only long-term trap sites to calculate trends
had little impact on assigning species to the vulnerable and
endangered categories (Fig. 5). A similar result was obtained
when sites running 20 or more years were used (Conrad
et al., 2004). Therefore, the all-sites analysis was used because
it provides greater spatial coverage, larger sample sizes for
individual species and enables estimates for a greater number
of less common species.
3.4. Light competition
Contrary to expectation, the annual index of total trap catch
(slope ± SE) at ‘dark’ sites (0.044 ± 0.007) decreased margin-
ally more than at ‘light’ sites (0.035 ± 0.005) although the dif-
ference between these slopes was not significant (t
38
= 0.97,
P= 0.34). The decrease in total macro-moths captured was
therefore as great or greater at sites that remained dark or be-
came darker than at those where night-time illumination in-
creased between 1992 and 2000. In addition, annual estimates
-1.0
-0.8
-0.6
-0.4
-0.2
0.0
0.2
0.4
1965 1975 1985 1995 2005
-1.0
-0.8
-0.6
-0.4
-0.2
0.0
0.2
0.4
year
1965 1970 1975 1980 1985 1990 1995 2000 2005
TRIM annual index
-1.0
-0.8
-0.6
-0.4
-0.2
0.0
0.2
0.4
Britain
north
south
N
S
Fig. 1 – Decreases in total annual trap catches for all species. The decrease for Great Britain is significant (t
33
= 8.83, P< 0.001),
as is the decrease in the south (t
33
= 10.9, P< 0.001), and represent 31% and 44% decreases in total macro-moths caught,
respectively. Trap catches have increased by 5% in the north, but this trend is not significant (t
33
= 0.67, P= 0.51).
BIOLOGICAL CONSERVATION 132 (2006) 279–291 281

of abundance were very similar between groups. This indi-
cates that the declines in moth abundance observed over
the course of our study are not caused by decreased effective-
ness of RIS light-traps due to increasing light competition, but
does not preclude the possibility that light pollution has been
a cause of moth population declines (Frank, 1988).
4. Discussion
This study has, for the first time, shown that the so-called
‘‘common and widespread’’ macro-moth species in Britain
are undergoing severe population declines. These estimates
of population change represent a wide variety of biotopes,
are robust to trap turnover, are not affected by light competi-
tion and are independent of total catches for individual
species.
The overall pattern of decline for so many species points to
a widespread deterioration of suitable environmental condi-
tions across the country. The deterioration has been most se-
vere in the south of England where the rapid intensification of
agriculture and forestry already has been implicated in the
decline of butterflies, especially in the southeast (Warren
et al., 2001). However, the fact that a large proportion of spe-
cies are declining rapidly in both north and south Britain
(Fig. 2) indicates that adverse environmental changes are
impacting moth populations across the country.
The IUCN categories of threat are widely used to prepare
‘Red lists’ of threatened species and have become an impor-
tant tool to identify ecological problems and guide conserva-
tion action (Mace and Lande, 1991; IUCN World Conservation
Union, 2001; Dunn, 2002). While the quantitative data on pop-
ulation dynamics demanded by IUCN categories are lacking
for almost all moths and other insects that are currently of
conservation concern around the world (New, 2004), the
extensive RIS dataset did allow us to determine, quantita-
tively, 10-yr rates of population change of a large group of
British macro-moths. Following the criteria of the IUCN cate-
gories in our study provides a well-recognized scale of the
severity of moth population declines.
In this study we found 71 common moth species that are
declining at rates that should see them designated as endan-
gered or vulnerable if the quantitative IUCN criteria are ap-
plied at the national scale (Gardenfors et al., 2001; Eaton
et al., 2005). None of the threatened species is known
for long-distance migrations and it is unlikely that the declin-
ing populations can be ‘‘rescued’’ by continental migrants.
-50
-40
-30
-20
-10
0
10
20
30
40
50
60
70
80
90
100
>100
0
20
40
60
80
vulnerable
20
40
60
80
percent change over 10 years
-50
-40
-30
-20
-10
0
10
20
30
40
50
60
70
80
90
100
>100
number of species
0
10
20
30
40
50
60
endangered
Vulnerable
Britain
north
south
N = 337
N = 274
N = 298
N
S
endangered
Fig. 2 – Frequency distributions of changes in abundance of British macro-moths. The figures plotted are the percentage
changes over a 10-yr period, calculated from the annual rate of change estimated from long-term trends from 1968–2002. The
vertical dashed line shows the median 10-yr change. X-axis labels are the upper limits of each class. Shaded areas
correspond with the criteria thresholds for threatened species in the vulnerable and endangered categories.
ln (total catch)
6 7 8 9 10 11 12 13
TRIM trend index
-0.2
-0.1
0.0
0.1
0.2
Fig. 3 – TRIM trend index versus the natural logarithm of
total trap catch for each of the 337 species in the study.
Frequently captured species are no more or less likely to
decline or increase than less common species.
282 BIOLOGICAL CONSERVATION 132 (2006) 279–291

Even so, it is more important that the magnitude of the de-
clines are sufficient that the species could be considered for
threatened status. The number of potentially threatened spe-
cies in this study is more than double the published British
Red Data Book list of 33 species (Shirt, 1987), none of which
was included in our analysis. This finding suggests we may
be seriously underestimating the proportion of threatened
British insects.
Designation of threatened status for common and wide-
spread species on the basis of population decline rates alone
has been criticized (Dunn, 2002) and the method of applying
IUCN criteria at national rather than global scales is still being
formalised, although their utility has been recognised (Gar-
denfors et al., 2001; Dunn, 2002; Eaton et al., 2005). Neverthe-
less, it is important that monitoring effort is directed toward
understanding population changes among common species
as well as rare ones (Conrad et al., 2002; Dunn, 2002). Com-
mon species may undergo dramatic population changes that
go largely unnoticed by recorders and conservation manag-
ers, but which could provide valuable information for conser-
vation and ecological studies (Thomas and Abery, 1995;
Cowley et al., 1999; Leon-Cortes et al., 1999). Common species
should represent a greater variety of habitats and species
interactions and therefore play an important role in ecosys-
tem functioning.
A brief examination of moth population trends in relation
to ecological and life-history traits identified few significant
associations and declines are taking place in a wide variety
of biotopes (Conrad et al., 2004). While widely distributed spe-
cies are more likely to be declining, increasing species are
likely to be those that are expanding their range as well as
increasing in abundance, and are often species apparently
benefiting from human activity, such as those feeding on or-
namental conifers (Conrad et al., 2004). The causes of long-
term trends identified in this study are yet to be assessed in
detail, and are likely to be a complex mixture of factors influ-
encing the quantity, quality and spatial distribution of suit-
able habitat (e.g., land management, chemical and light
pollution, climatic conditions). Causes of decline will also
undoubtedly vary from species to species.
All of the moth species in our study are common and
widespread. Truly specialised species, such as have been de-
scribed for British butterflies (Warren et al., 2001) are too
uncommon and too locally distributed (Quinn et al., 1997)to
have been caught in sufficient numbers to be used in our
analysis and are therefore under-represented. If, like special-
ist butterflies (Warren et al., 2001), these species are more
Year
1970 1975 1980 1985 1990 1995 2000
% in land-use category
0
20
40
60
80
100
coastal
farmland
mixed
moorland
parkland
scrubland
urban
woodland
Fig. 4 – Annual proportions of land-use categories for light-traps used in the study.
declining
vulnerable
endangered
percent change, "all sites" analysis
-80 -60 -40 -20 0 20
percent change, sites that ran at least 15 years
-80
-60
-40
-20
0
20
Fig. 5 – Comparison of 10-yr trends estimated by analysis of
all light-trap sites and only using sites that operated for at
least 15 yr. The four areas shaded pale grey delineate
regions of assignment of rapid-decline categories between
the two methods of estimating trends.
BIOLOGICAL CONSERVATION 132 (2006) 279–291 283

likely to be declining, then we have underestimated the over-
all proportions of declining macro-moths.
Half of the species we studied experienced a 10-yr de-
cline of at least 12%, and while the precise comparison of
trends between different sampling methods is difficult and
may give misleading results (Thomas, 1996) our results sug-
gest that British macro-moths have undergone declines at
least as severe as British butterflies (Thomas et al., 2004).
Moreover, the percentage of moth species declining (66%)
was similar to the proportion of butterflies declining
(71%), and greater than the proportion of birds (54%) or
plants declining (28%) (Thomas et al., 2004; Eaton et al.,
2005). Thus, our findings support the view that insect biodi-
versity is declining very rapidly in Britain and probably at a
greater rate than vertebrates and vascular plants (Thomas
et al., 2004), with potentially serious consequences for eco-
system services.
Common macro-moths have undergone widespread and
serious declines in Britain. Environmental changes that affect
common and widespread herbivores, such as the macro-
moths reported here, signal strong impacts on the wider
ecosystem and at higher trophic levels such as predacious in-
sects, insectivorous spiders, birds and bats (Pollard and Yates,
1993; Ormerod and Watkinson, 2000; Donald et al., 2001;
Wickramasinghe et al., 2004). Compared to UK butterflies
(Thomas et al., 2004), the macro-moths in this study include
a greater number of species from a wider range of habitats
and, therefore are more likely to be representative of terres-
trial insect biodiversity. However, the observed declines of
macro-moths, taken together with those of butterfly species,
signal a biodiversity crisis for Britain and are a strong indica-
tor that insects may be facing great losses in other temperate-
zone industrialised countries. As yet, even correlative evi-
dence of factors driving long-term moth population trends
is lacking, but having identified so many decreasing trends,
the next priority is to examine the relative roles of climate,
chemical and light pollution, and changes in land-use in
greater detail.
Acknowledgements
We wish to acknowledge the efforts of the numerous volun-
teers who help run and maintain the light-traps of the
Rothamsted Insect Survey. Joe Perry, Suzanne Clark and Pe-
ter Rothery offered statistical advice and discussion. Arco
van Strien provided excellent advice and support for TRIM.
Marie Castellazzi extracted the UK light-change data from
world maps kindly provided by Chris Elvidge from the US
National Geophysical Data Centre. Georgina Mace and Erica
Dunn advised on the use of IUCN criteria. This study was
funded by the Esme
´e Fairbairn Foundation and the UK Bio-
technology and Biological Sciences Research Council
(BBSRC), from which Rothamsted Research receives grant-
aided support.
Appendix A. List of species studied with rates of annual population change
Number = ‘‘Bradley number’’, from Checklist of Lepidoptera recorded from the British Isles (Bradley, 2000); annual change
rate = annual rate of population change estimated from the 35-yr time series (see methods); 95% CI = 95% confidence interval for
the annual change rate; change status: increasing = change rate >0, declining = change rate <0, vulnerable = greater than
30% Æ10 yr
1
decline, endangered = greater than 50% 10 yr
1
decline.
Number Vernacular name Species Annual
change rate
95% CI Change
status
14 Ghost Swift Hepialus humuli 0.036 0.027, 0.046 Vulnerable
15 Orange Swift Hepialus sylvina 0.023 0.031, 0.015 Increasing
17 Common Swift Hepialus lupulinus 0.005 0.003, 0.013 Declining
18 Map-Winged Swift Hepialus fusconebulosa 0.014 0.007, 0.022 Declining
1631 December Moth Poecilocampa populi 0.030 0.025, 0.034 Declining
1632 Pale Eggar Trichiura crataegi 0.054 0.042, 0.065 Vulnerable
1634 The Lackey Malacosoma nuestria 0.063 0.044, 0.082 Vulnerable
1640 The Drinker Euthrix potatoria 0.007 0.000, 0.015 Declining
1645 Scalloped Hook-Tip Falcaria lacertinaria 0.021 0.013, 0.028 Declining
1646 Oak Hook-Tip Drepana binaria 0.047 0.033, 0.061 Vulnerable
1648 Pebble Hook-Tip Drepana falcataria 0.020 0.012, 0.027 Declining
1651 Chinese Character Cilix glaucata 0.018 0.011, 0.024 Declining
1652 Peach Blossom Thyatira batis 0.028 0.020, 0.036 Declining
1653 Buff Arches Habrosyne pyritoides 0.034 0.026, 0.043 Declining
1657 Common Lutestring Ochropacha duplaris 0.031 0.044, 0.018 Increasing
1658 Oak Lutestring Cymatophorima diluta 0.048 0.023, 0.072 Vulnerable
1659 Yellow-Horned Achlya flavicornis 0.015 0.022, 0.008 Increasing
1663 March Moth Alsophila aescularia 0.013 0.008, 0.019 Declining
1665 Grass Emerald Pseudoterpna pruinata 0.030 0.016, 0.044 Declining
1666 Large Emerald Geometra papilionaria 0.009 0.016, 0.002 Increasing
1667 Blotched Emerald Comibaena bajularia 0.008 0.013, 0.029 Declining
1669 Common Emerald Hemithea aestivaria 0.008 0.002, 0.014 Declining
284 BIOLOGICAL CONSERVATION 132 (2006) 279–291

Appendix A – continued
Number Vernacular name Species Annual
change rate
95% CI Change
status
1673 Small Emerald Hemistola chrysoprasaria 0.049 0.023, 0.074 Vulnerable
1674 Little Emerald Jodis lactearia 0.002 0.007, 0.010 Declining
1677 Birch Mocha Cyclophora albipunctata 0.020 0.002, 0.038 Declining
1680 Maiden’s Blush Cyclophora punctaria 0.028 0.046, 0.011 Increasing
1682 Blood-Vein Timandra griseata 0.043 0.037, 0.049 Vulnerable
1689 Mullein Wave Scopula marginepunctata 0.040 0.021, 0.059 Vulnerable
1690 Small Blood-Vein Scopula imitaria 0.028 0.021, 0.035 Declining
1692 Lesser Cream Wave Scopula immutata 0.003 0.023, 0.029 Declining
1693 Cream Wave Scopula floslactata 0.009 0.003, 0.015 Declining
1694 Smoky Wave Scopula ternata 0.006 0.017, 0.030 Declining
1699 Least Carpet Idaea vulpinaria 0.188 0.248, 0.128 Increasing
1702 Small Fan-Footed Wave Idaea biselata 0.006 0.001, 0.011 Declining
1705 Dwarf Cream Wave Idaea fuscovenosa 0.048 0.062, 0.034 Increasing
1707 Small Dusty Wave Idaea seriata 0.013 0.022, 0.003 Increasing
1708 Single-Dotted Wave Idaea dimidiata 0.013 0.019, 0.007 Increasing
1709 Satin Wave Idaea subsericeata 0.012 0.001, 0.023 Declining
1711 Treble Brown-Spot Idaea trigeminata 0.104 0.117, 0.090 Increasing
1712 Small Scallop Idaea emarginata 0.009 0.001, 0.017 Declining
1713 Riband Wave Idaea aversata 0.005 0.009, 0.001 Increasing
1715 Plain Wave Idaea straminata 0.043 0.079, 0.008 Increasing
1716 The Vestal Rhodometra sacraria 0.060 0.120, 0.000 Increasing
1719 Oblique Carpet Orthonama vittata 0.050 0.034, 0.065 Vulnerable
1722 Flame Carpet Xanthorhoe designata 0.018 0.026, 0.010 Increasing
1723 Red Carpet Xanthorhoe munitata 0.046 0.035, 0.057 Vulnerable
1724 Red Twin-Spot Carpet Xanthorhoe spadicearia 0.016 0.010, 0.022 Declining
1725 Dark-Barred Twin-Spot Xanthorhoe ferrugata 0.069 0.062, 0.076 Endangered
1726 Large Twin-Spot Carpet Xanthorhoe quadrifasiata 0.010 0.001, 0.021 Declining
1727 Silver-Ground Carpet Xanthorhoe montanata 0.015 0.010, 0.020 Declining
1728 Garden Carpet Xanthorhoe fluctuata 0.033 0.028, 0.038 Declining
1732 Shaded Broad-Bar Scotopteryx chenopodiata 0.037 0.029, 0.045 Vulnerable
1738 Common Carpet Epirrhoe alternata 0.004 0.002, 0.010 Declining
1739 Wood Carpet Epirrhoe rivata 0.001 0.017, 0.014 Increasing
1740 Galium Carpet Epirrhoe galiata 0.040 0.019, 0.062 Vulnerable
1742 Yellow Shell Camptogramma bilineata 0.019 0.029, 0.009 Increasing
1744 Grey Mountain Carpet Entephria caesiata 0.044 0.024, 0.064 Vulnerable
1745 The Mallow Larentia clavaria 0.009 0.001, 0.020 Declining
1746 Shoulder Stripe Anticlea badiata 0.032 0.026, 0.038 Declining
1747 The Streamer Anticlea derivata 0.019 0.012, 0.026 Declining
1748 Beautiful Carpet Mesoleuca albicillata 0.004 0.024, 0.016 Increasing
1749 Dark Spinach Pelurga comitata 0.085 0.061, 0.108 Endangered
1750 Water Carpet Lampropteryx suffumata 0.005 0.012, 0.002 Increasing
1751 Devon Carpet Lampropteryx otregiata 0.069 0.118, 0.020 Increasing
1752 Purple Bar Cosmorhoe ocellata 0.007 0.001, 0.012 Declining
1753 Striped Twin-Spot Carpet Nebula salicata 0.010 0.010, 0.030 Declining
1754 The Phoenix Eulithis prunata 0.012 0.026, 0.002 Increasing
1755 The Chevron Eulithis testata 0.015 0.007, 0.022 Declining
1756 Northern Spinach Eulithis populata 0.019 0.023, 0.015 Increasing
1757 The Spinach Eulithis mellinata 0.084 0.060, 0.108 Endangered
1758 Barred Straw Eulithis pyraliata 0.020 0.014, 0.026 Declining
1759 Small Phoenix Ecliptopera silaceata 0.042 0.035, 0.049 Vulnerable
1760 Red–green Carpet Chloroclysta siterata 0.057 0.067, 0.047 Increasing
1761 Autumn Green Carpet Chloroclysta miata 0.014 0.005, 0.023 Declining
1762 Dark Marbled Carpet Chloroclysta citrata 0.012 0.019, 0.005 Increasing
1764 Common Marbled Carpet Chloroclysta truncata 0.019 0.014, 0.024 Declining
1765 Barred Yellow Cidaria fulvata 0.010 0.003, 0.018 Declining
(continued on next page)
BIOLOGICAL CONSERVATION 132 (2006) 279–291 285

Appendix A – continued
Number Vernacular name Species Annual
change rate
95% CI Change
status
1766 Blue-Bordered Carpet Plemyria rubiginata 0.049 0.065, 0.032 Increasing
1767 Pine Carpet Thera firmata 0.038 0.051, 0.025 Increasing
1768 Grey Pine Carpet Thera obeliscata 0.005 0.011, 0.002 Increasing
1769 Spruce Carpet Thera britannica 0.067 0.090, 0.044 Increasing
1771 Juniper Carpet Thera juniperata 0.077 0.120, 0.034 Increasing
1773 Broken-Barred Carpet Electrophaes corylata 0.007 0.004, 0.018 Declining
1775 Mottled Grey Colostygia multistrigaria 0.026 0.019, 0.034 Declining
1776 Green Carpet Colostygia pectinataria 0.026 0.033, 0.018 Increasing
1777 July Highflyer Hydriomena furcata 0.012 0.018, 0.006 Increasing
1778 May Highflyer Hydriomena impluviata 0.005 0.010, 0.020 Declining
1781 Small Waved Umber Horisme vitalbata 0.014 0.033, 0.005 Increasing
1782 The Fern Horisme tersata 0.015 0.003, 0.032 Declining
1784 Pretty Chalk Carpet Melanthia procellata 0.056 0.038, 0.074 Vulnerable
1789 Scallop Shell Rheumaptera undulata 0.017 0.002, 0.031 Declining
1792 Dark Umber Philereme transversata 0.034 0.021, 0.048 Declining
1794 Sharp-Angled Carpet Euphyia unangulata 0.031 0.019, 0.042 Declining
1795 November Moth Epirrita dilutata 0.031 0.027, 0.036 Declining
1797 Autumnal Moth Epirrita autumnata 0.011 0.001, 0.020 Declining
1798 Small Autumnal Moth Epirrita filigrammaria 0.022 0.040, 0.084 Declining
1799 Winter Moth Operophtera brumata 0.004 0.003, 0.012 Declining
1800 Northern Winter Moth Operophtera fagata 0.011 0.001, 0.020 Declining
1802 The Rivulet Perizoma affinitata 0.015 0.006, 0.024 Declining
1803 Small Rivulet Perizoma alchemillata 0.003 0.009, 0.014 Declining
1807 Grass Rivulet Perizoma albulata 0.090 0.067, 0.113 Endangered
1808 Sandy Carpet Perizoma flavofasciata 0.005 0.003, 0.013 Declining
1809 Twin-Spot Carpet Perizoma didymata 0.028 0.036, 0.019 Increasing
1858 V-Pug Chloroclystis v-ata 0.009 0.022, 0.004 Increasing
1864 The Streak Chesias legatella 0.042 0.033, 0.051 Vulnerable
1865 Broom-Tip Chesias rufata 0.052 0.022, 0.081 Vulnerable
1867 Treble-Bar Aplocera plagiata 0.032 0.021, 0.044 Declining
1873 Welsh Wave Venusia cambrica 0.005 0.021, 0.010 Increasing
1874 Dingy Shell Euchoeca nebulata 0.020 0.065, 0.026 Increasing
1875 Small White Wave Asthena albulata 0.001 0.030, 0.028 Increasing
1881 Early Tooth-Striped Trichopteryx carpinata 0.032 0.041, 0.022 Increasing
1882 Small Seraphim Pterapherapteryx sexalata 0.033 0.015, 0.051 Declining
1883 Yellow-Barred Brindle Acasis viretata 0.023 0.036, 0.011 Increasing
1884 The Magpie Abraxas grossulariata 0.033 0.025, 0.040 Declining
1887 Clouded Border Lomaspilis marginata 0.004 0.001, 0.010 Declining
1888 Scorched Carpet Ligdia adustata 0.020 0.011, 0.029 Declining
1889 Peacock Semiothisa notata 0.091 0.132, 0.050 Increasing
1890 Sharp-Angled Peacock Semiothisa alternaria 0.013 0.001, 0.027 Declining
1893 Tawny-Barred Angle Semiothisa liturata 0.002 0.012, 0.008 Increasing
1894 Latticed Heath Semiothisa clathrata 0.058 0.048, 0.067 Vulnerable
1897 The V-Moth Semiothisa wauaria 0.097 0.072, 0.122 Endangered
1902 Brown Silver-Lines Petrophora chlorosata 0.005 0.000, 0.009 Declining
1903 Barred Umber Plagodis pulveraria 0.021 0.031, 0.011 Increasing
1904 Scorched Wing Plagodis dolabraria 0.002 0.010, 0.005 Increasing
1906 Brimstone Moth Opisthograptis luteolata 0.013 0.009, 0.017 Declining
1907 Bordered Beauty Epione repandaria 0.008 0.000, 0.016 Declining
1910 Lilac Beauty Apeira syringaria 0.031 0.023, 0.040 Declining
1912 August Thorn Ennomos quercinaria 0.047 0.033, 0.061 Vulnerable
1913 Canary-Shouldered Thorn Ennomos alniaria 0.030 0.024, 0.036 Declining
1914 Dusky Thorn Ennomos fuscantaria 0.103 0.088, 0.119 Endangered
1915 September Thorn Ennomos erosaria 0.068 0.056, 0.080 Endangered
1917 Early Thorn Selenia dentaria 0.026 0.022, 0.030 Declining
1918 Lunar Thorn Selenia lunularia 0.015 0.005, 0.026 Declining
286 BIOLOGICAL CONSERVATION 132 (2006) 279–291

Appendix A – continued
Number Vernacular name Species Annual
change rate
95% CI Change
status
1919 Purple Thorn Selenia tetralunaria 0.032 0.024, 0.041 Declining
1920 Scalloped Hazel Odontopera bidentata 0.004 0.001, 0.009 Declining
1921 Scalloped Oak Crocallis elinguaria 0.031 0.026, 0.035 Declining
1922 Swallow-Tail Moth Ourapteryx sambucaria 0.024 0.018, 0.031 Declining
1923 Feathered Thorn Colotois pennaria 0.024 0.019, 0.029 Declining
1926 Pale Brindled Beauty Apocheima pilosaria 0.022 0.012, 0.032 Declining
1927 Brindled Beauty Lycia hirtaria 0.046 0.038, 0.055 Vulnerable
1930 Oak Beauty Biston strataria 0.003 0.004, 0.011 Declining
1931 Peppered Moth Biston betularia 0.027 0.018, 0.035 Declining
1932 Spring Usher Agriopis leucophaearia 0.010 0.034, 0.015 Increasing
1933 Scarce Umber Agriopis aurantiaria 0.028 0.018, 0.039 Declining
1934 Dotted Border Agriopis marginaria 0.022 0.017, 0.027 Declining
1935 Mottled Umber Erannis defoliaria 0.000 0.012, 0.012 Increasing
1937 Willow Beauty Peribatodes rhomboidaria 0.015 0.009, 0.022 Declining
1940 Satin Beauty Deileptenia ribeata 0.111 0.153, 0.069 Increasing
1941 Mottled Beauty Alcis repandata 0.010 0.015, 0.005 Increasing
1942 Dotted Carpet Alcis jubata 0.062 0.077, 0.048 Increasing
1944 Pale Oak Beauty Serraca punctinalis 0.007 0.022, 0.009 Increasing
1945 Brussels Lace Cleorodes lichenaria 0.011 0.011, 0.034 Declining
1947 The Engrailed Ectropis bistortata 0.003 0.009, 0.003 Increasing
1950 Brindled White-Spot Paradarisa extersaria 0.008 0.014, 0.029 Declining
1951 Grey Birch Aethalura punctulata 0.000 0.019, 0.020 Declining
1954 Bordered White Bupalus piniaria 0.011 0.004, 0.027 Declining
1955 Common White Wave Cabera pusaria 0.016 0.021, 0.011 Increasing
1956 Common Wave Cabera exanthemata 0.006 0.011, 0.000 Increasing
1957 White-Pinion Spotted Lomographa bimaculata 0.010 0.031, 0.011 Increasing
1958 Clouded Silver Lomographa temerata 0.018 0.012, 0.025 Declining
1961 Light Emerald Campaea margaritata 0.007 0.011, 0.002 Increasing
1962 Barred Red Hylaea fasciaria 0.003 0.010, 0.005 Increasing
1981 Poplar Hawk-Moth Laothoe populi 0.007 0.001, 0.012 Declining
1994 Buff-Tip Phalera bucephala 0.022 0.012, 0.031 Declining
2000 Iron Prominent Notodonta dromedarius 0.012 0.001, 0.025 Declining
2003 Pebble Prominent Eligmodonta ziczac 0.021 0.011, 0.031 Declining
2005 Great Prominent Peridea anceps 0.016 0.028, 0.003 Increasing
2006 Lesser Swallow Prominent Pheosia gnoma 0.019 0.013, 0.026 Declining
2007 Swallow Prominent Pheosia tremula 0.012 0.027, 0.003 Increasing
2008 Coxcomb Prominent Ptilodon capucina 0.025 0.019, 0.030 Declining
2011 Pale Prominent Pterostoma palpina 0.009 0.002, 0.017 Declining
2014 Marbled Brown Drymonia dodonaea 0.011 0.000, 0.023 Declining
2015 Lunar Marbled Brown Drymonia ruficornis 0.022 0.039, 0.006 Increasing
2020 Figure of Eight Diloba caeruleocephala 0.081 0.071, 0.090 Endangered
2028 Pale Tussock Calliteara pudibunda 0.015 0.005, 0.024 Declining
2030 Yellow-Tail Euproctis similis 0.006 0.000, 0.013 Declining
2033 Black Arches Lymantria monacha 0.020 0.036, 0.005 Increasing
2035 Round-Winged Muslin Thumatha senex 0.013 0.039, 0.014 Increasing
2037 Rosy Footman Miltochrista miniata 0.040 0.054, 0.026 Increasing
2038 Muslin Footman Nudaria mundana 0.022 0.034, 0.010 Increasing
2040 Four-Dotted Footman Cybosia mesomella 0.004 0.014, 0.005 Increasing
2044 Dingy Footman Eilema griseola 0.063 0.076, 0.049 Increasing
2047 Scarce Footman Eilema complana 0.091 0.112, 0.070 Increasing
2049 Buff Footman Eilema deplana 0.065 0.104, 0.027 Increasing
2050 Common Footman Eilema lurideola 0.010 0.016, 0.004 Increasing
2057 Garden Tiger Arctia caja 0.062 0.054, 0.071 Vulnerable
2059 Clouded Buff Diacrisia sannio 0.028 0.007, 0.050 Declining
2060 White Ermine Spilosoma lubricipeda 0.041 0.035, 0.046 Vulnerable
(continued on next page)
BIOLOGICAL CONSERVATION 132 (2006) 279–291 287

Appendix A – continued
Number Vernacular name Species Annual
change rate
95% CI Change
status
2061 Buff Ermine Spilosoma luteum 0.037 0.031, 0.042 Vulnerable
2063 Muslin Moth Diaphora mendica 0.007 0.015, 0.001 Increasing
2064 Ruby Tiger Phragmatobia fuliginosa 0.007 0.015, 0.001 Increasing
2069 Cinnabar Tyria jacobaeae 0.049 0.035, 0.063 Vulnerable
2077 Short-Cloaked Moth Nola cucullatella 0.021 0.011, 0.030 Declining
2078 Least Black Arches Nola confusalis 0.061 0.082, 0.040 Increasing
2081 White-Line Dart Euxoa tritici 0.069 0.051, 0.088 Endangered
2082 Garden Dart Euxoa nigricans 0.097 0.067, 0.126 Endangered
2085 Archer’s Dart Agrotis vestigialis 0.031 0.016, 0.046 Declining
2087 Turnip Moth Agrotis segetum 0.032 0.022, 0.042 Declining
2088 Heart & Club Agrotis clavis 0.002 0.012, 0.016 Declining
2089 Heart & Dart Agrotis exclamationis 0.031 0.023, 0.040 Declining
2091 Dark Sword-Grass Agrotis ipsilon 0.025 0.003, 0.047 Declining
2092 Shuttle-Shaped Dart Agrotis puta 0.009 0.019, 0.001 Increasing
2098 The Flame Axylia putris 0.021 0.014, 0.029 Declining
2102 Flame Shoulder Ochropleura plecta 0.001 0.005, 0.007 Declining
2107 Large Yellow Underwing Noctua pronuba 0.025 0.030, 0.019 Increasing
2109 Lesser Yellow Underwing Noctua comes 0.017 0.024, 0.011 Increasing
2110 Broad-Bordered Yellow Underwing Noctua fimbriata 0.070 0.094, 0.046 Increasing
2111 Lesser Broad-Bordered Yellow Underwing Noctua janthe 0.008 0.015, 0.002 Increasing
2114 Double Dart Graphiphora augur 0.097 0.084, 0.110 Endangered
2117 Autumnal Rustic Paradiarsa glareosa 0.070 0.060, 0.079 Endangered
2118 True Lover’s Knot Lycophotia porphyrea 0.029 0.023, 0.036 Declining
2120 Ingrailed Clay Diarsia mendica 0.031 0.026, 0.036 Declining
2121 Barred Chestnut Diarsia dahlii 0.033 0.045, 0.021 Increasing
2122 Purple Clay Diarsia brunnea 0.018 0.012, 0.025 Declining
2123 Small Square-Spot Diarsia rubi 0.052 0.045, 0.060 Vulnerable
2126 Setaceous Hebrew Character Xestia c-nigrum 0.004 0.010, 0.003 Increasing
2127 Triple-Spotted Clay Xestia ditrapezium 0.020 0.002, 0.041 Declining
2128 Double Square-Spot Xestia triangulum 0.014 0.008, 0.019 Declining
2130 Dotted Clay Xestia baja 0.014 0.007, 0.021 Declining
2132 Neglected or Grey Rustic Xestia castanea 0.047 0.029, 0.065 Vulnerable
2133 Six-Striped Rustic Xestia sexstrigata 0.021 0.012, 0.029 Declining
2134 Square-Spot Rustic Xestia xanthographa 0.005 0.011, 0.001 Increasing
2135 Heath Rustic Xestia agathina 0.052 0.029, 0.074 Vulnerable
2136 The Gothic Naenia typica 0.032 0.012, 0.051 Declining
2138 Green Arches Anaplectoides prasina 0.019 0.031, 0.007 Increasing
2139 Red Chestnut Cerastis rubricosa 0.021 0.014, 0.029 Declining
2145 The Nutmeg Discestra trifolii 0.017 0.001, 0.035 Declining
2147 The Shears Hada plebeja 0.010 0.020, 0.001 Increasing
2150 Grey Arches Polia nebulosa 0.015 0.001, 0.029 Declining
2154 Cabbage Moth Mamestra brassicae 0.015 0.006, 0.025 Declining
2155 Dot Moth Melanchra persicariae 0.059 0.044, 0.073 Vulnerable
2158 Pale-Shouldered Brocade Lacanobia thalassina 0.003 0.011, 0.005 Increasing
2160 Bright-Line Brown-Eye Lacanobia oleracea 0.011 0.004, 0.018 Declining
2163 Broom Moth Ceramica pisi 0.041 0.032, 0.049 Vulnerable
2173 The Lychnis Hadena bicruris 0.024 0.010, 0.037 Declining
2176 Antler Moth Cerapteryx graminis 0.031 0.024, 0.038 Declining
2177 Hedge Rustic Tholera cespitis 0.098 0.087, 0.110 Endangered
2178 Feathered Gothic Tholera decimalis 0.065 0.052, 0.077 Vulnerable
2179 Pine Beauty Panolis flammea 0.044 0.057, 0.032 Increasing
2182 Small Quaker Orthosia cruda 0.008 0.021, 0.004 Increasing
2186 Powdered Quaker Orthosia gracilis 0.040 0.030, 0.050 Vulnerable
2187 Common Quaker Orthosia cerasi 0.006 0.013, 0.002 Increasing
2188 Clouded Drab Orthosia incerta 0.008 0.002, 0.014 Declining
2189 Twin-Spotted Quaker Orthosia munda 0.001 0.009, 0.011 Declining
288 BIOLOGICAL CONSERVATION 132 (2006) 279–291

Appendix A – continued
Number Vernacular name Species Annual
change rate
95% CI Change
status
2190 Hebrew Character Orthosia gothica 0.011 0.006, 0.015 Declining
2192 Brown-Line Bright-Eye Mythimna conigera 0.023 0.012, 0.035 Declining
2193 The Clay Mythimna ferrago 0.009 0.004, 0.015 Declining
2198 Smoky Wainscot Mythimna impura 0.000 0.006, 0.006 Declining
2199 Common Wainscot Mythimna pallens 0.029 0.021, 0.036 Declining
2205 Shoulder-Striped Wainscot Mythimna comma 0.036 0.024, 0.048 Vulnerable
2225 Minor Shoulder-Knot Brachylomia viminalis 0.037 0.025, 0.048 Vulnerable
2227 The Sprawler Brachionycha sphinx 0.049 0.040, 0.057 Vulnerable
2229 Brindled Ochre Dasypolia templi 0.063 0.040, 0.085 Vulnerable
2231 Deep-Brown Dart
a
Aporophyla lutulenta 0.064 0.044, 0.084 Vulnerable
2232 Black Rustic Aporophyla nigra 0.032 0.019, 0.044 Declining
2237 Grey Shoulder-Knot Lithophane ornitopus 0.072 0.101, 0.044 Increasing
2240 Blair’s Shoulder-Knot Lithophane leautieri 0.165 0.243, 0.087 Increasing
2241 Red Sword-Grass Xylena vetusta 0.013 0.002, 0.028 Declining
2243 Early Grey Xylocampa areola 0.004 0.013, 0.005 Increasing
2245 Green-Brindled Crescent Allophyes oxyacanthae 0.044 0.038, 0.050 Vulnerable
2247 Merveille Du Jour Dichonia aprilina 0.005 0.020, 0.009 Increasing
2248 Brindled Green Dryobotodes eremita 0.040 0.058, 0.023 Increasing
2250 Dark Brocade Mniotype adusta 0.043 0.027, 0.058 Vulnerable
2254 Grey Chi Antitype chi 0.023 0.005, 0.041 Declining
2255 Feathered Ranunculus Eumichtis lichenea 0.007 0.003, 0.018 Declining
2256 The Satellite Eupsilia transversa 0.024 0.035, 0.014 Increasing
2258 The Chestnut Conistra vaccinii 0.012 0.017, 0.007 Increasing
2259 Dark Chestnut Conistra ligula 0.019 0.009, 0.029 Declining
2262 The Brick Agrochola circellaris 0.028 0.021, 0.035 Declining
2263 Red-Line Quaker Agrochola lota 0.007 0.016, 0.001 Increasing
2264 Yellow-Line Quaker Agrochola macilenta 0.014 0.020, 0.007 Increasing
2265 Flounced Chestnut Agrochola helvola 0.058 0.043, 0.072 Vulnerable
2266 Brown-Spot Pinion Agrochola litura 0.039 0.031, 0.048 Vulnerable
2267 Beaded Chestnut Agrochola lychnidis 0.064 0.057, 0.072 Vulnerable
2269 Centre-Barred Sallow Atethmia centrago 0.038 0.029, 0.046 Vulnerable
2270 Lunar Underwing Omphaloscelis lunosa 0.020 0.027, 0.013 Increasing
2272 Barred Sallow Xanthia aurago 0.017 0.005, 0.029 Declining
2273 Pink-Barred Sallow Xanthia togata 0.018 0.012, 0.025 Declining
2274 The Sallow Xanthia icteritia 0.048 0.040, 0.056 Vulnerable
2275 Dusky-Lemon Sallow Xanthia gilvago 0.070 0.036, 0.104 Endangered
2284 Grey Dagger Acronicta psi 0.041 0.028, 0.054 Vulnerable
2289 Knot Grass Acronicta rumicis 0.045 0.035, 0.054 Vulnerable
2293 Marbled Beauty Cryphia domestica 0.051 0.062, 0.039 Increasing
2299 Mouse Moth Amphipyra tragopogonis 0.037 0.030, 0.044 Vulnerable
2302 Brown Rustic Rusina ferruginea 0.015 0.010, 0.019 Declining
2303 Straw Underwing Thalpophila matura 0.031 0.022, 0.040 Declining
2305 Small Angle Shades Euplexia lucipara 0.019 0.011, 0.027 Declining
2306 Angle Shades Phlogophora meticulosa 0.011 0.018, 0.004 Increasing
2312 The Olive Ipimorpha subtusa 0.031 0.061, 0.001 Increasing
2318 The Dun-Bar Cosmia trapezina 0.000 0.007, 0.008 Declining
2319 Lunar-Spotted Pinion Cosmia pyralina 0.026 0.010, 0.042 Declining
2321 Dark Arches Apamea monoglypha 0.009 0.004, 0.014 Declining
2322 Light Arches Apamea lithoxylaea 0.035 0.026, 0.043 Declining
2326 Clouded-Bordered Brindle Apamea crenata 0.003 0.007, 0.014 Declining
2330 Dusky Brocade Apamea remissa 0.039 0.028, 0.051 Vulnerable
2333 Large Nutmeg Apamea anceps 0.058 0.034, 0.081 Vulnerable
2334 Rustic Shoulder-Knot Apamea sordens 0.027 0.018, 0.036 Declining
2335 Slender Brindle Apamea solopacina 0.016 0.038, 0.006 Increasing
2340 Middle-Barred Minor Oligia fasciuncula 0.013 0.006, 0.019 Declining
(continued on next page)
BIOLOGICAL CONSERVATION 132 (2006) 279–291 289

REFERENCES
Bradley, J.D., 2000. Checklist of Lepidoptera Recorded from the
British Isles, second ed. Bradley & Bradley, Fordingbridge.
Conrad, K.F., Woiwod, I.P., Perry, J.N., 2002. Long-term decline in
abundance and distribution of the garden tiger moth (Arctia
caja) in Great Britain. Biological Conservation 106, 329–337.
Conrad, K.F., Woiwod, I.P., Perry, J.N., 2003. East Atlantic
teleconnection pattern and the decline of a common arctiid
moth. Global Change Biology 9, 125–130.
Conrad, K.F., Woiwod, I.P., Parsons, M., Fox, R., Warren,
M.S., 2004. Long-term population trends in widespread
British moths. Journal of Insect Conservation 8,
119–136.
Cowley, M.J.R., Thomas, C.D., Thomas, J.A., Warren, M.S., 1999.
Flight areas of British butterflies: assessing species status and
decline. Proceedings of the Royal Society of London Series
B-Biological Sciences 266, 1587–1592.
Donald, P.F., Green, R.E., Heath, M.M., 2001. Agricultural
intensification and collapse of Europe’s farmland bird
populations. Proceedings of the Royal Society of London Series
B-Biological Sciences 268, 25–29.
Appendix A – continued
Number Vernacular name Species Annual
change rate
95% CI Change
status
2341 Cloaked Minor Mesoligia furuncula 0.022 0.032, 0.012 Increasing
2342 Rosy Minor Mesoligia literosa 0.047 0.035, 0.058 Vulnerable
2343 Common Rustic Mesapamea secalis 0.004 0.009, 0.002 Increasing
2345 Small Dotted Buff Photedes minima 0.020 0.015, 0.025 Declining
2350 Small Wainscot Photedes pygmina 0.024 0.016, 0.031 Declining
2352 Dusky Sallow Eremobia ochroleuca 0.009 0.022, 0.004 Increasing
2353 Flounced Rustic Luperina testacea 0.019 0.013, 0.024 Declining
2357 Large Ear Amphipoea lucens 0.019 0.006, 0.044 Declining
2360 Ear Moth Amphipoea oculea 0.035 0.019, 0.051 Vulnerable
2361 Rosy Rustic Hydraecia micacea 0.054 0.047, 0.060 Vulnerable
2364 Frosted Orange Gortyna flavago 0.012 0.002, 0.022 Declining
2367 Haworth’s Minor Celaena haworthii 0.062 0.045, 0.079 Vulnerable
2368 The Crescent Celaena leucostigma 0.048 0.030, 0.066 Vulnerable
2375 Large Wainscot Rhizedra lutosa 0.054 0.042, 0.066 Vulnerable
2380 Treble Lines Charanyca trigrammica 0.007 0.019, 0.004 Increasing
2381 The Uncertain Hoplodrina alsines 0.002 0.009, 0.005 Increasing
2382 The Rustic Hoplodrina blanda 0.039 0.030, 0.048 Vulnerable
2384 Vine’s Rustic Hoplodrina ambigua 0.048 0.077, 0.019 Increasing
2387 Mottled Rustic Caradrina morpheus 0.037 0.030, 0.044 Vulnerable
2389 Pale Mottled Willow Caradrina clavipalpis 0.023 0.038, 0.007 Increasing
2394 The Anomalous Stilbia anomala 0.075 0.052, 0.097 Endangered
2410 Marbled White-Spot Protodeltote pygarga 0.018 0.032, 0.005 Increasing
2422 Green Silver-Lines Pseudoips prasinana 0.027 0.040, 0.015 Increasing
2425 Nut-Tree Tussock Colocasia coryli 0.015 0.023, 0.007 Increasing
2434 Burnished Brass Diachrysia chrysitis 0.024 0.018, 0.030 Declining
2439 Gold Spot Plusia festucae 0.018 0.036, 0.000 Increasing
2441 Silver Y Autographa gamma 0.019 0.014, 0.024 Declining
2442 Beautiful Golden Y Autographa pulchrina 0.009 0.002, 0.015 Declining
2443 Plain Golden Y Autographa jota 0.004 0.009, 0.017 Declining
2444 Gold Spangle Autographa bractea 0.002 0.016, 0.012 Increasing
2450 The Spectacle Abrostola tripartita 0.012 0.019, 0.005 Increasing
2473 Beautiful Hook-Tip Laspeyria flexula 0.029 0.016, 0.041 Declining
2474 Straw Dot Rivula sericealis 0.031 0.046, 0.016 Increasing
2475 Waved Black Parascotia fuliginaria 0.004 0.009, 0.016 Declining
2477 The Snout Hypena proboscidalis 0.006 0.000, 0.012 Declining
2489 The Fan-Foot Herminia tarsipennalis 0.013 0.006, 0.021 Declining
2492 Small Fan-Foot Herminia grisealis 0.016 0.011, 0.021 Declining
– Lead/July Belle Aggregate
b
Scotopteryx spp 0.035 0.024, 0.045 Declining
a Deep-brown dart Aporophyla lutulenta, and Northern deep-brown dart A. luenerbergensis were not initially recorded as sep-
arate species and appear in the table as an aggregate of counts of both species.
b After compiling the data we determined that Lead Belle (Scotopteryx mucronata, 1733) and July Belle (S. luridata, 1734) could
not be reliably distinguished on the basis of external appearance, gross morphology, phenology or distribution, so the catches
of the two species were combined.
290 BIOLOGICAL CONSERVATION 132 (2006) 279–291

Dunn, E.H., 2002. Using decline in bird populations to identify
needs for conservation action. Conservation Biology 16,
1632–1637.
Eaton, M.A., Gregory, R.D., Noble, D.G., Robinson, J.A., Hughes, J.,
Procter, D., Brown, A.F., Gibbons, D.W., 2005. Regional IUCN red
listing: the process as applied to birds in the United Kingdom.
Conservation Biology 19, 1557–1570.
Elvidge, C.D., Imhoff, M.L., Baugh, K.E., Hobson, V.R., Nelson, I.,
Safran, J., Dietz, J.B., Tuttle, B.T., 2001. Night-time lights of the
world: 1994–1995. Isprs Journal of Photogrammetry and
Remote Sensing 56, 81–99.
Frank, K.D., 1988. Impact of outdoor lighting on moths: an
assessment. Journal of the Lepidopterists Society 42, 63–93.
Gardenfors, U., Hilton-Taylor, C., Mace, G.M., Rodriguez, J.P., 2001.
The application of IUCN red list criteria at regional levels.
Conservation Biology 15, 1206–1212.
Hambler, C., Speight, M.R., 2004. Extinction rates and butterflies.
Science 305, 1563.
IUCN World Conservation Union. 2001. IUCN Red List categories
and criteria: Version 3.1. IUCN Species Survival Commission,
Gland, Switzerland and Cambridge, UK.
Lawton, J.H., May, R.M., 1995. Extinction Rates. Oxford University
Press, oxford.
Leon-Cortes, J.L., Cowley, M.J.R., Thomas, C.D., 1999. Detecting
decline in a formerly widespread species: how common is the
common blue butterfly Polyommatus icarus? Ecography 22,
643–650.
Longcore, T., Rich, C., 2004. Ecological light pollution. Frontiers in
Ecology and the Environment 2, 191–198.
Mace, G.M., Lande, R., 1991. Assessing extinction threats: toward
re-evaluation of IUCN threatened species categories.
Conservation Biology 5, 148–157.
McKinney, M.L., 1999. High rates of extinction and threat in poorly
studied taxa. Conservation Biology 13, 1273–1281.
New, T.R., 2004. Moths (Insecta: Lepidoptera) and conservation:
background and perspective. Journal of Insect Conservation 8,
79–94.
Ormerod, S.J., Watkinson, A.R., 2000. Editor’s introduction: birds
and agriculture. Journal of Applied Ecology 37, 699.
Pannekoek, J., Van Strien, A.J. 2001. TRIM 3 Manual (TRends &
Indices for Monitoring data) Statistics Netherlands, Voorburg.
Pollard, E., Yates, T., 1993. Monitoring Butterflies for Ecology and
Conservation, first ed. Chapman and Hall, London.
Quinn, R.M., Gaston, K.J., Blackburn, T.M., Eversham, B.C., 1997.
Abundance-range size relationships of macrolepidoptera in
Britain: the effects of taxonomy and life history variables.
Ecological Entomology 22, 453–461.
Shirt, D.B., 1987British Red Data Books, vol. 2. Insects Nature
Conservancy Council, Peterborough.
Taylor, L.R., French, R.A., 1974. Effects of light-trap design and
illumination on samples of moths in an English woodland.
Bulletin of Entomological Research 63, 583–594.
Thomas, L., 1996. Monitoring long-term population change: why
are there so many analysis methods? Ecology 77, 49–58.
Thomas, J.A., 2005. Monitoring change in the abundance and
distribution of insects using butterflies and other indicator
groups. Philosophical Transactions of the Royal Society
B-Biological Sciences 360, 339–357.
Thomas, C.D., Abery, J.C.G., 1995. Estimating rates of butterfly
decline from distribution maps – the effect of scale. Biological
Conservation 73, 59–65.
Thomas, J.A., Clarke, R.T., 2004. Extinction rates and butterflies.
Science 305, 1563–1564.
Thomas, J.A., Morris, M.G., 1994. Patterns, mechanisms and rates
of extinction among invertebrates in the United Kingdom.
Philosophical Transactions of the Royal Society of London
Series B-Biological Sciences 344, 47–54.
Thomas, J.A., Telfer, M.G., Roy, D.B., Preston, C.D., Greenwood,
J.J.D., Asher, J., Fox, R., Clarke, R.T., Lawton, J.H., 2004.
Comparative losses of British butterflies, birds, and plants and
the global extinction crisis. Science 303, 1879–1881.
Van Strien, A.J., Pannekoek, J., Gibbons, D.W., 2001. Indexing
European bird population trends using results of national
monitoring schemes: a trial of a new method. Bird Study 48,
200–213.
Warren, M.S., Hill, J.K., Thomas, J.A., Asher, J., Fox, R., Huntley, B.,
Roy, D.B., Telfer, M.G., Jeffcoate, S., Harding, P., Jeffcoate, G.,
Willis, S.G., Greatorex-Davies, J.N., Moss, D., Thomas, C.D.,
2001. Rapid responses of British butterflies to opposing forces
of climate and habitat change. Nature 414, 65–69.
Wickramasinghe, L.P., Harris, S., Jones, G., Vaughan Jennings, N.,
2004. Abundance and species richness of nocturnal insects on
organic and conventional farms: effects of agricultural
intensification on bat foraging. Conservation Biology 18,
1283–1292.
Williams, C.B., 1948. The Rothamsted light trap. Proceedings of
the Royal Entomological Society of London A 23, 80–85.
Williams, C.B., 1952. Some notes on killing insects for collections
and for scientific research. The Entomologist 85, 271–279.
Wilson, E.O., 1992. The Diversity of Life. Penguin Press, London.
Woiwod, I.P., Harrington, R., 1994. Flying in the face of change:
The Rothamsted Insect Survey. In: Leigh, R., Johnston, A. (Eds.),
Long-Term Experiments in Agricultural and Ecological
Sciences. CAB International, Wallingford, pp. 321–342.
BIOLOGICAL CONSERVATION 132 (2006) 279–291 291