Ecological Applications, 24(7), 2014, pp. 1561–1568
Ó2014 by the Ecological Society of America
LED lighting increases the ecological impact of light pollution
irrespective of color temperature
S. M. PAWSON
AND M. K.-F. BADER
Scion, P.O. Box 29-237, Fendalton, Christchurch, New Zealand
Scion, 49 Sala Street, Rotorua, New Zealand
Abstract. Recognition of the extent and magnitude of night-time light pollution impacts
on natural ecosystems is increasing, with pervasive effects observed in both nocturnal and
diurnal species. Municipal and industrial lighting is on the cusp of a step change where energy-
efﬁcient lighting technology is driving a shift from ‘‘yellow’’ high-pressure sodium vapor lamps
(HPS) to new ‘‘white’’ light-emitting diodes (LEDs). We hypothesized that white LEDs would
be more attractive and thus have greater ecological impacts than HPS due to the peak UV-
green-blue visual sensitivity of nocturnal invertebrates. Our results support this hypothesis; on
average LED light traps captured 48%more insects than were captured with light traps ﬁtted
with HPS lamps, and this effect was dependent on air temperature (signiﬁcant light 3air
temperature interaction). We found no evidence that manipulating the color temperature of
white LEDs would minimize the ecological impacts of the adoption of white LED lights. As
such, large-scale adoption of energy-efﬁcient white LED lighting for municipal and industrial
use may exacerbate ecological impacts and potentially amplify phytosanitary pest infestations.
Our ﬁndings highlight the urgent need for collaborative research between ecologists and
electrical engineers to ensure that future developments in LED technology minimize their
potential ecological effects.
Key words: biodiversity; high-pressure sodium lamp; light pollution; spectra; street lighting;
Since the invention of the ﬁrst practical incandescent
light bulb in the late 1870s, night-time light pollution has
now become almost ubiquitous in the populated regions
of developed countries (Bogard 2013). The extent of
artiﬁcial light pollution continues to expand swiftly
(;6%annual increase, range 0–20%[Ho
¨lker et al.
2010a]), especially in newly industrialized economies.
Several aspects of light pollution are clear: (1) there is
extensive evidence that artiﬁcial lighting that exceeds
natural background levels has signiﬁcant ecological and
biological impacts (Longcore and Rich 2004, Rich and
Longcore 2005, Ho
¨lker et al. 2010b, Davies et al. 2012,
Gaston et al. 2013, Le Tallec et al. 2013, Perkin et al.
2014), (2) the spectral composition of light pollution can
alter the magnitude of these impacts (van Langevelde et
al. 2011, Davies et al. 2013), and (3) the spectral
composition of light pollution has changed, and will
continue to change over time with the advent and
subsequent adoption of more energy-efﬁcient lighting
technologies (Schubert and Kim 2005).
The current trend in global lighting is a shift from
‘‘yellow’’ sodium lamps toward a new generation of
broad spectrum, energy-efﬁcient, ‘‘white’’ light-emitting
diodes (LEDs) for municipal and industrial lighting
(Schubert and Kim 2005, Anonymous 2012). The
biological effect of a shift toward a more ‘‘white-light
night’’ (sensu Gaston et al. 2012) has not been studied in
detail, but direct evidence of biological impacts is
mounting (Stone et al. 2012), and comparisons of visual
pigment absorbance spectra with the emission spectra of
municipal light sources suggests indirectly that such
impacts may be widespread among terrestrial animals
(Davies et al. 2013).
Gaston et al. (2012) proposed that the ecological
consequences of light pollution could potentially be
reduced by avoiding critical regions within the spectrum.
Currently available municipal and industrial-scale white
LED lights are based on monochromatic blue LEDs
Manuscript received 6 March 2014; revised 29 May 2014;
accepted 12 June 2014. Corresponding Editor: M. P. Ayres.
coated by a single yellow, or multiple yellow-green,
phosphor coatings that absorb blue light and reemit
longer wavelength emissions (Krames et al. 2007). The
phosphor coating can be manipulated to produce a
range of white LEDs that differ in the proportion of
blue (435–495 nm) wavelengths emitted. This range of
white LEDs is normally referred to by their color
temperature (degrees Kelvin [K]) with higher tempera-
tures having a greater proportion of emitted blue light.
Given the peak UV, blue, and green photoreceptors of
many invertebrates (Briscoe and Chittka 2001), we
hypothesize that low color temperature LED lights will
have less ecological impact than high color temperature
LED lights due to the lower intensity blue spectral
To test this hypothesis we ﬁrst compared the relative
attraction of ﬂying invertebrates to 4000 K white LEDs
and high-pressure sodium lamps at a scale equivalent to
current industrial/municipal site-lighting practices.
This comparison provided an assessment of the
potential impact of white LEDs on nocturnal inverte-
brates if adopted for industrial and municipal lighting.
We then compared the relative attraction of ﬂying
invertebrates to different color temperature white
LEDs at an experimental scale to identify opportunities
for minimizing the ecological impact of white LED
lighting. For both experiments we used the attraction
of nocturnal ﬂying invertebrates as a proxy measure of
Data collection: Comparison of LED and HPS
An industrial-scale lighting comparison of white
LEDs and high-pressure sodium (HPS) lamps was
conducted using ﬁve replicate pairs of 4000 K white
LEDs and HPS lamps (see Appendix, Table 1). The key
difference in the spectral composition of the LED and
HPS lamps was that the LED had greater relative
intensity from the blue-green portion of the spectra than
the HPS lamp (Fig. 1). An A2-sized sheet of Perspex
(Evonik Industries, Darmstadt, Germany) mounted 0.5
m below and directly between each pair of LED and
HPS lamps was used to sample ﬂying invertebrates
attracted to the lights (see Appendix, Fig. 1). Each night,
an A2-sized sheet of Tanglefoot-coated (Contech,
Victoria, British Columbia, Canada) Mylar (Fuji Xerox,
Connecticut, USA) was attached to both sides of the
Perspex pane to snare ﬂying invertebrates. Mylar sheets
were identiﬁed as ‘‘facing’’ and ‘‘away,’’ depending on
their orientation with respect to the location of the lamp
that was activated on that particular night. Trapping
was conducted between 21:00 and 00:00 on 10 suitable
nights from 21 January and 3 February with each
sampling location randomly assigned to either LED or
HPS lighting on the ﬁrst night. The active light in each
pair was then alternated on subsequent nights. This
resulted in a ﬁnal design of ﬁve independent replicates
(sampling locations) that compare the two light treat-
ments that were sampled on 10 different nights. The 10
sampling occasions cannot be considered as truly
independent, hence tests for the inﬂuence of repeated
measures were performed (see Analysis: Comparison of
LED and HPS industrial-scale lighting). Suitable nights
were considered to be nights with a forecast air
temperature at 21:00 of at least 158C. Actual air
temperature at 22:00 that was recorded at the study site
at an elevation of 10 m was used for analyses (see
Analysis: Comparison of LED and HPS industrial-scale
lighting). All LED and HPS comparisons were conduct-
ed at the PanPac wood processing facility, Hawkes Bay,
New Zealand. Pairs of lights were established on the
edge of industrial buildings at the site. The site is
bordered by an extensive Pinus radiata plantation forest
to the west and by coastal grassland to the east with the
ocean located ,1 km to the east. Insects are known to
disperse into the site from the forest as they are attracted
by the bright site lighting.
Data collection: Comparison of LED color temperature
The attraction of ﬂying invertebrates to white LEDs
with six different color temperatures (see Appendix,
FIG. 1. Relative spectral emission of (a) high-pressure
sodium (HPS) lamp (Sunlux ACE, NH-360 FLX; EYE
Lighting, Wacol, Queensland, Australia) and (b) light-emitting
diode (LED) 4000 K color temperature high bay lamp
(LUXEON M, LXR7-SW40; Koninklijke Philips, Amsterdam,
The Netherlands). Emission spectra are normalized to the
spectrum with the maximum intensity and were kindly provided
by each manufacturer.
S. M. PAWSON AND M. K.-F. BADER1562 Ecological Applications
Vol. 24, No. 7
Table 1, Fig. 2) were compared to a control treatment
(18-ohm resister that matched the power consumption
and heat output of the tested LEDs) in a completely
randomized block design with three replicates (see Plate
1). Each of the three blocks consisted of a linear transect
with seven sampling points located at 20-m intervals.
Within each block, the six color treatments (and control)
were initially assigned at random to one of the seven
sampling points. On subsequent nights the individual
color treatments were then rotated sequentially to
account for any potentially confounding spatial effects
that may have occurred due to the location of the
sampling point along the transect within the block.
LEDs were powered by 12-V DC batteries and the
experiment was conducted over four hours each night
(20:00 to 00:00) on seven non-consecutive days between
30 January and the 26 February 2013. Delays in
sampling were due to periods of unsuitable weather
when invertebrate ﬂight activity was minimal.
LEDs were mounted in a heat sink housing (Makers-
LED, Aimes, Iowa, USA) and the attraction of ﬂying
invertebrates to lights was quantiﬁed using A3 Perspex
catching panes installed 30 cm in front of each LED.
Tanglefoot-coated Mylar sheets were attached to the
Perspex to sample ﬂying invertebrates attracted to the
light. Unimpeded light was visible to the sides of the
Tanglefoot-coated Mylar sheets, however a portion of
light emitted during the study did have to pass through
the Mylar sheet, potentially altering the spectral compo-
sition. To test for this the spectral emission of LEDs with
and without a Tanglefoot-coated Mylar sheet were
compared (Fig. 2). This showed that there was some
absorption of light in the 600þnm wavelengths (red and
infrared), but this change should not have biased our
results, as most insects are not sensitive to changes in the
far red portion of the spectra (Briscoe and Chittka 2001).
In the highly attractive blue-green portion of the spectra,
the Tanglefoot coated Mylar sheet did not impact the
relative proportion of light emitted.
The input current of individual LEDs was adjusted
(via a potentiometer attached to a LuxDrive BuckPuck
Driver [LED Dynamics, Randolph, Vermont, USA]) to
produce a uniform power output of 12 61 mW (mean
6SD) for each color temperature (see Appendix, Table
1). To measure power output for calibration the
emission from individual LEDs was collimated using a
25.4 mm focal length 1-inch diameter (1 inch ¼2.54 cm)
lens that was focused on a Coherent J-50MB-HE
thermopile sensor (Coherent, Santa Clara, California,
USA). Power output was averaged over 10-s intervals,
using a Coherent FieldMaxII-TO.
All LED color temperature trials were conducted on the
boundary of the Synlait facility, Rakaia, Christchurch,
New Zealand. Each of the three independent blocks was
established on areas of long grass (intermittently mown)
or gravel. The area is surrounded by introduced exotic
pastoral grass with shelter belts of Pinus radiata. There are
no substantial areas of non-productive ecosystems in the
immediate vicinity of the site.
Analysis: Comparison of LED and HPS industrial-scale
Total number of ﬂying invertebrates (pooling panes
that were ‘‘facing’’ and ‘‘away’’) were standardized by
the nightly trapping duration to account for the
variation in trapping times ranging from 3.7 to 4.2
hours. This standardization procedure changed the
scale of the response variables from a discrete to a
continuous scale and thus allowed the use of linear
mixed effects models (R package nlme [Pinheiro et al.
2014]). The ﬁxed term of the model contained light
type, air temperature, wind speed, and their interac-
tions as explanatory variables. Trapping date nested in
light type and sampling location were modeled as
random terms to account for the hierarchical design
and repeated measures. Plots of the standardized
residuals vs. ﬁtted values and for each of the
explanatory variables were used for graphical model
validation. The validation plots indicated heterosce-
dasticity, which was modeled using a power variance
structure that incorporated the ﬁtted values as a
variance covariate and light type as grouping variable
(i.e., allowing for stratiﬁed variance modelling). The
signiﬁcance of the ﬁxed model terms was assessed via
backward selection using likelihood ratio tests (Zuur et
al. 2009). The ﬁnal models showed high correlation
between the intercept and the slopes of the ﬁxed
factors. To overcome this issue the models were
reﬁtted using centered air temperature and wind speed
Analysis: Comparison of LED color temperature
We applied generalized linear mixed models (GLMM)
with Poisson errors and log link ﬁt by Laplace
approximation (R package lme4 [Bates et al. 2014]) to
analyze the trap catch data. The total number of ﬂying
invertebrates caught per trap was compared against
color temperature, location (sampling point within
transect), and their interaction as ﬁxed terms within
the model. Block and trapping date were modeled as
random effects. Overdispersion was detected (ratio of
residual deviance to residual degrees of freedom .1)
and accounted for by incorporating a per-observation-
TABLE 1. Results of the optimal linear mixed-effects model
results for standardized invertebrates catches at lights
equipped with light-emitting diode (LED) or high-pressure
sodium (HPS) lamps.
Parameter Estimate SE df tP
Intercept 21.65 2.77 50 7.81 ,0.001
Light 8.55 2.55 42 3.35 0.002
Temp 5.61 0.99 42 5.67 ,0.001
Light 3temp 2.53 1.10 42 2.30 0.027
Notes: Sample size n¼5 blocks. Abbreviations are light, light
type; temp, air temperature; and wind, wind speed.
October 2014 1563ECOLOGICAL EFFECTS OF LED LIGHTING
level random effect. The GLMM was followed by a
multiple comparison test using Tukey contrasts to allow
pairwise comparisons between color temperatures (R
package multcomp [Hothorn et al. 2008]). Plots of the
Pearson residuals vs. ﬁtted values and against the
response variable(s) were applied for graphical model
validation. The signiﬁcance of the ﬁxed model terms was
assessed via backward selection using Akaike’s infor-
mation criterion (AIC; Zuur et al. 2009). The AIC was
favored over the likelihood ratio test, as used for the
comparison between LED and HPS lamps (see Analysis:
Comparison of LED and HPS industrial-scale lighting).
Our rationale for using the AIC is that it includes a
penalty for the number of parameters, which discour-
ages over-ﬁtting of the model (in this case 36 parameters
were associated with the color temperature 3location
interaction term). When DAIC 2, we considered the
FIG. 2. Relative spectral emission of ‘‘white’’ LED of differing color temperatures. Emission spectra were measured using an
OceanOptics USB2000 ﬁber-coupled spectrometer (Ocean Optics, Dunedin, Florida, USA) with an integration time of 100 ms.
Emission spectra are normalized to the maximum spectral intensity in the 400–500 nm range, as insects are most attracted to this
portion of the visible spectrum. The shape and form of the curve in the blue-green region is almost identical between naked LEDs
and operational LEDs where measurements were taken behind the Perspex and Tanglefoot-coated Mylar.
FIG. 3. Number of ﬂying invertebrates (standardized by the
3.7–4.2 h trapping duration) caught in light traps equipped with
LED or HPS light. Values are means 6SE, n¼5 replicates.
S. M. PAWSON AND M. K.-F. BADER1564 Ecological Applications
Vol. 24, No. 7
competitive models to provide similar goodness of ﬁts
and opted for the model with fewer parameters.
Standardization of trapping times was not required for
the comparison of color temperatures as the sample
duration varied by less than 10 minutes between
treatments. All analyses were conducted in R version
2.15.3 (R Development Core Team 2013).
Comparison of LED and HPS industrial-scale lighting
In total 7300 invertebrates were caught including,
3811 Diptera, 1376 Trichoptera, 994 Coleoptera, 409
Hymenoptera, 308 Hemiptera, 173 Ephemeroptera,
111 Psocidae, and less than 100 Lepidoptera, Neurop-
tera, Thysanoptera, Araneae, Plecoptera, Isoptera,
Orthoptera, and Blattodea. Sampling panes equipped
with LED lamps attracted 48%more ﬂying inverte-
brates on average than HPS lamps (Fig. 3, Table 1).
Insect attraction to light was signiﬁcantly affected by
air temperature (Table 1), we observed a precipitous
decline in catch numbers during a single night when
air temperatures were favorable for ﬂight (;208C).
This coincided with a strong easterly wind blowing
from the ocean. The prevailing wind directions at the
study site were from west and southwest suggesting
that ﬂying invertebrates from nearby forested land
would have to ﬂy into a headwind to reach the
Comparison of LED color temperatures
In total 12 860 invertebrates were caught, including
8879 Diptera, 1674 Lepidoptera, 1089 Thysanoptera,
450 Coleoptera, 379 Hymenoptera, 144 Neuroptera,
and ,100 Hemiptera, Psocoptera, Trichoptera, Ara-
neae, Ephemeroptera, Collembola, and Mantodea (in
decreasing order of abundance). When considering the
pooled catch of all ﬂying invertebrate taxa (removing
Araneae, Collembola, and Acarina) LED lamps at-
tracted signiﬁcantly more ﬂying invertebrates than
control traps irrespective of color temperature (Fig. 4,
Table 2). However, the difference between control and
LED lamps was taxon dependent, strong effects were
observed in Diptera and Lepidoptera and no effect
observed for Thysanoptera, Hymenoptera, and Cole-
optera (Fig. 4). Irrespective of taxa, the LED color
temperature had no signiﬁcant effect on invertebrate
attraction (Fig. 4).
Light pollution is recognized as a global threat to the
conservation of biological diversity that could drive
reductions in the quality of provisioning, regulating and
PLATE 1. Image showing one experimental block with ﬁve of the six different color temperature LEDs. Note the dark spot
between the 4th and 5th light as the location of the just visible control treatment. The 6th color temperature is not visible and is to
the left of the image. Individual lights were placed 20 m apart with different blocks placed 200 m apart. Photo credit: S. M. Pawson,
October 2014 1565ECOLOGICAL EFFECTS OF LED LIGHTING
cultural ecosystem services (Ho
¨lker et al. 2010b). There
is increasing evidence that existing light pollution has
signiﬁcant ecological effects (Rich and Longcore 2005,
Gaston et al. 2012, 2013, Perkin et al. 2014). However,
the spatial extent, density, and spectral composition of
light pollution is predicted to change (Gaston et al.
2012). Continued urban expansion will result in a
concomitant encroachment of light pollution into areas
that are currently naturally lit, however, spectral
changes may alter the type of impacts that these new
areas experience, e.g., changed species interactions
(Davies et al. 2013).
FIG. 4. Effect of LED color temperature on ﬂying invertebrates trap catch over a 4-h sampling period; C is the control trap,
values below the bars indicate the color temperature in Kelvin (n¼3 blocks, values are means 6SE). Different lower case letters
indicate statistically signiﬁcant differences at a¼0.05 (multiple comparison procedure using Tukey contrasts).
TABLE 2. Results of a backward selection applied to the
generalized linear mixed effects model for LED color
temperature and trap position.
Dropped term AIC
Color temperature 3location 595.11
Color temperature 658.01
Notes: AIC stands for Akaike’s information criterion.
The most parsimonious model, which only contained LED
color temperature as explanatory variable (i.e., the model with
the interaction term and the factor location dropped).
S. M. PAWSON AND M. K.-F. BADER1566 Ecological Applications
Vol. 24, No. 7
White LED lighting for both municipal and indus-
trial applications is predicted to increase dramatically
in the next decade (Anonymous 2012). Gaston et al.
(2012) have referred to this anticipated shift from high-
pressure sodium (HPS) lights to LEDs, as the
formation of a ‘‘white-light night.’’ Our results suggest
that a white-light night shift could signiﬁcantly increase
the ecological impacts of light pollution as white LEDs
attracted 48%more ﬂying invertebrates than existing
HPS lamps. Our study only accounts for the absolute
loss of individuals from the population of nocturnal
ﬂying invertebrates attracted to LED lights. However,
the true extent of white LED light pollution will require
an assessment of ecological effects across multiple
ecological levels, e.g., species, populations, and com-
munities to address varying complexity in the potential
interactions (as suggested by Fox ). In addition
further research is required to understand the land-
scape-scale inﬂuence of LED lights at broader spatial
scales. As discussed by Davies et al. (2013), light
pollution may affect visually guided behaviors of both
individuals and of interactions between species or
ecological guilds, e.g., predator avoidance and/or prey
detection, navigation, pollination, and foraging, may
be inﬂuenced by light pollution. However, the magni-
tude of such potential effects may prove to be
dependent on both habitat structure (e.g., forest
canopy vs. open grassland) and the spatial arrangement
Manipulating LED color temperature is one poten-
tial intervention that could minimize the ecological
impacts of white LEDs as it reduces the intensity of
blue spectral emissions that are attractive to inverte-
brates. However, our results show that the attraction of
nocturnal ﬂying invertebrates to currently available
phosphor coated white LEDs does not vary with LED
color temperature. This effect was strongly observed in
Diptera and Lepidoptera as they were the most
numerous taxa attracted to the LEDs (Fig. 4).
However, there was no observed effect of white LED
lights for Hymenoptera, Thysanoptera, and Coleop-
tera. Given the low sample size for these three
taxonomic groups it is difﬁcult to draw absolute
conclusions as these individuals may represent acci-
dental by-catch in the hour before dusk that it took to
install the sticky sheets, as opposed to actual nocturnal
Our general ﬁnding for all taxa (Fig. 4) is contrary to
our initial hypotheses, and implies that careful
selection of currently available off-the-shelf color
temperatures is unlikely to mitigate the potential
ecological impacts of a broad-scale shift to white
LED lighting for municipal and industrial applica-
tions. One potential explanation for this is that current
low color temperature white LEDs still emit a
proportion of blue-green spectra (Fig. 2). It may be
possible to overcome this issue using longpass optical
ﬁlters, or alternatively by selecting speciﬁc monochro-
matic LEDs (with narrow spectral wavelengths) that
avoid the highly attractive blue-green spectra, however
this has yet to be tested.
In addition to their direct ecological impacts light
pollution from white LEDs is likely to exacerbate
existing domestic, e.g., midge swarms and industrial
infestations of sanitary and phytosanitary pests that are
known to be highly attractive to white lighting (Pawson
et al. 2009, Goretti et al. 2011). The potential nuisance
impact of such unwanted domestic pest species is an
additional factor that should be considered in the
selection of municipal lighting. However, more impor-
tant is the potential for white LED lighting to increase
phytosanitary and biosecurity risks that could lead to
additional indirect ecological impacts. For example,
white light is more attractive than light emitted from
HPS lamps to gypsy moth (Lymantria dispar); an
invasive, polyphagous, forest pest (Walliner et al.
1995). The potential ecological impacts from the
establishment of gypsy moth in new regions are severe,
e.g., defoliation affecting productivity (Sharov et al.
2002) and local extinction of other Lepidoptera
(Wagner and Van Driesche 2010), and ships infested
with egg masses are a known pathway that is actively
monitored by a number of countries, including
Australia and New Zealand (MacLellan 2011). Thus a
transition to white LEDs at, or near, ports may elevate
the risk of egg masses moving on a transoceanic
pathway, which potentially increases the risk of
establishment in new regions.
Although we have shown that the color temperature
of existing yellow phosphor white LEDs cannot be used
to reduce their ecological impact on ﬂying inverte-
brates, other options may reduce the effects of white
LEDs in the future. Gaston et al. (2012) highlight the
potential of white LEDs derived from a combination of
monochromatic LED light sources, e.g., red, blue, and
green, that together would form a white light. This may
provide greater ability to avoid certain spectral
emissions to reduce the effects of light pollution.
However, before multiple primary LEDs (e.g., RGB
emitters) can become a reality for large-scale illumina-
tion there are signiﬁcant technological breakthroughs
required for both green and red LEDs (Krames et al.
2007). Alternatively, longpass ﬁlters could be used to
remove speciﬁc spectral emissions as was previously
suggested to reduce the attractiveness of mercury vapor
and high-pressure sodium lamps to particular pest
species (e.g., gypsy moth; Walliner et al. 1995).
Practically such ﬁlters may have limitations as they
would signiﬁcantly alter color rendering and may
increase per unit costs and energy consumption per
unit of light emitted.
Phosphor-coated white LED lamps have the potential
to increase the impacts of light pollution dramatically.
Given the strong impetus for their adoption in municipal
and industrial applications, it is imperative to fully
understand the potential long-term impacts of white
LED lights on ecological communities, populations, and
species. A comprehensive assessment of overall impacts
and knowledge about the inﬂuence of each region of the
visible spectrum will allow technologists to work with
October 2014 1567ECOLOGICAL EFFECTS OF LED LIGHTING
ecologists to focus future developments in lighting
technology that balance the needs of illumination with
reduced ecological impact.
The authors acknowledge Paul Fielder and Zachary Treamer
from LED Dynamics for advice on LED binning and the
supply of speciﬁc LED components. Rob Eagle of PanPac and
Simon Causer of Synlait milk products for access to their
respective facilities to conduct experiments. Sebastian Horvath
for the analysis of LED spectral frequencies and the calibration
of power output from different color temperature LEDs. Jess
Kerr, Brooke O’Connor, Liam Wright, Tia Uaea, Thornton
Campbell, Krystal Jansen, and Arild Roberts for the establish-
ment of ﬁeld trial sites and Sarah Cross for assistance in
identifying and counting insect samples.
Anonymous. 2012. Lighting the way: perspectives on the global
lighting market. Second edition. McKinsey and Company,
New York, New York, USA.
Bates, D., M. Maechler, B. Bolker, and S. Walker. 2014. lme4:
linear mixed-effects models using Eigen and S4. R package
version 1.1-7. http://CRAN.R-project.org/package¼lme4
Bogard, P. 2013. The end of night: searching for natural
darkness in an age of artiﬁcial light. Little, Brown and
Company, New York, New York, USA.
Briscoe, A. D., and L. Chittka. 2001. The evolution of color
vision in insects. Annual Review of Entomology 46:471–510.
Davies, T. W., J. Bennie, and K. J. Gaston. 2012. Street lighting
changes the composition of invertebrate communities.
Biology Letters 8:764–767.
Davies, T. W., J. Bennie, R. Inger, N. H. de Ibarra, and K. J.
Gaston. 2013. Artiﬁcial light pollution: are shifting spectral
signatures changing the balance of species interactions?
Global Change Biology 19:1417–1423.
Fox, R. 2013. The decline of moths in Great Britain: a review of
possible causes. Insect Conservation and Diversity 6:5–19.
Gaston, K. J., J. Bennie, T. W. Davies, and J. Hopkins. 2013.
The ecological impacts of nighttime light pollution: a
mechanistic appraisal. Biological Reviews 88(4):912–927.
Gaston, K. J., T. W. Davies, J. Bennie, and J. Hopkins. 2012.
Review: reducing the ecological consequences of night-time
light pollution: options and developments. Journal of
Applied Ecology 49:1256–1266.
Goretti, E., A. Coletti, A. Di Veroli, A. M. Di Giulio, and E.
Gaino. 2011. Artiﬁcial light device for attracting pestiferous
chironomids (Diptera): a case study at Lake Trasimeno
(Central Italy). Italian Journal of Zoology 78:336–342.
¨lker, F., et al. 2010a. The dark side of light: a transdisci-
plinary research agenda for light pollution policy. Ecology
and Society 15(4):13.
¨lker, F., C. Wolter, E. K. Perkin, and K. Tockner. 2010b.
Light pollution as a biodiversity threat. Trends in Ecology
and Evolution 25:681–682.
Hothorn, T., F. Bretz, and P. Westfall. 2008. Simultaneous
inference in general parametric models. Biometrical Journal
Krames, M. R., O. B. Shchekin, R. Mueller-Mach, G. O.
Mueller, L. Zhou, G. Harbers, and M. G. Craford. 2007.
Status and future of high-power light-emitting diodes for
solid-state lighting. IEEE/OSA Journal of Display Technol-
Le Tallec, T., M. Perret, and M. The
´ry. 2013. Light pollution
modiﬁes the expression of daily rhythms and behavior
patterns in a nocturnal primate. PLoS ONE 8:e79250.
Longcore, T., and C. Rich. 2004. Ecological light pollution.
Frontiers in Ecology and the Environment 2:191–198.
MacLellan, R. 2011. Gypsy moth surveillance in New Zealand.
Pawson, S. M., M. S. Watt, and E. G. Brockerhoff. 2009. Using
differential responses to light spectra as a monitoring and
control tool for Arhopalus ferus (Coleoptera: Cerambycidae)
and other exotic wood boring pests. Journal of Economic
Perkin, E. K., F. Ho
¨lker, and K. Tockner. 2014. The effects of
artiﬁcial lighting on adult aquatic and terrestrial insects.
Freshwater Biology 59:368–377.
Pinheiro, J., D. Bates, S. DebRoy, D. Sarkar, and R Core
Team. 2014. nlme: linear and nonlinear mixed effects models.
R package version 3.1-117. http://CRAN.R-project.org/
R Development Core Team. 2013. R: a language and
environment for statistical computing. R Foundation for
Statistical Computing, Vienna, Austria. http://www.
Rich, C., and T. Longcore, editors. 2005. Ecological conse-
quences of artiﬁcial night lighting. Island Press, Washington,
Schubert, E. F., and J. K. Kim. 2005. Solid-state light sources
getting smart. Science 308:1274–1278.
Sharov, A. A., D. Leonard, A. M. Liebhold, E. A. Roberts, and
W. Dickerson. 2002. ‘‘Slow the Spread’’: a national program
to contain the gypsy moth. Journal of Forestry 100:30–35.
Stone, E. L., G. Jones, and S. Harris. 2012. Conserving energy
at a cost to biodiversity? Impacts of LED lighting on bats.
Global Change Biology 18:2458–2465.
van Langevelde, F., J. A. Ettema, M. Donners, M. F. Wallis-
DeVries, and D. Groenendijk. 2011. Effect of spectral
composition of artiﬁcial light on the attraction of moths.
Biological Conservation 144:2274–2281.
Wagner, D. L., and R. G. Van Driesche. 2010. Threats posed to
rare or endangered insects by invasions of nonnative species.
Annual Review of Entomology 55:547–568.
Baranchikov, and R. T. Carde. 1995. Response of adult
lymantriid moths to illumination devices in the Russian far
east. Journal of Economic Entomology 88:337–342.
Zuur, A. F., E. N. Leno, N. J. Walker, A. A. Saveliev, and
G. M. Smith. 2009. Mixed effects models and extensions in
ecology with R. Springer, New York, New York, USA.
Photos of experimental setup and speciﬁcations of light-emitting diode (LED) and high-pressure sodium vapor (HPS) lamps
used for all studies (Ecological Archives A024-191-A1).
Raw invertebrate data of LED color comparison and LED vs. HPS comparison (Ecological Archives A024-191-S1).
S. M. PAWSON AND M. K.-F. BADER1568 Ecological Applications
Vol. 24, No. 7