ArticlePDF AvailableLiterature Review

Abstract

In recent decades, advances in lighting technology have precipitated exponential increases in night sky brightness worldwide, raising concerns in the scientific community about the impact of artificial light at night (ALAN) on crepuscular and nocturnal biodiversity. Long‐term records show that insect abundance has declined significantly over this time, with worrying implications for terrestrial ecosystems. The majority of investigations into the vulnerability of nocturnal insects to artificial light have focused on the flight‐to‐light behavior exhibited by select insect families. However, ALAN can affect insects in other ways as well. This review proposes five categories of ALAN impact on nocturnal insects, highlighting past research and identifying key knowledge gaps. We conclude with a summary of relevant literature on bioluminescent fireflies, which emphasizes the unique vulnerability of terrestrial light‐based communication systems to artificial illumination. Comprehensive understanding of the ecological impacts of ALAN on diverse nocturnal insect taxa will enable researchers to seek out methods whereby fireflies, moths, and other essential members of the nocturnal ecosystem can coexist with humans on an increasingly urbanized planet.
Ecology and Evolution . 2018 ;1–2 2 .    
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www.ecolevol.org
1 | INTRODUCTION
Localized illumination of nocturnal landscapes by anthropogenic
sources of light such as street lamps, path lights, and vehicle head
lights, hereafter referred to collectively as artificial light at night
(ALAN), is likely to disrupt populations of crepuscular and nocturnal
animal species present in affected habitats (Davies & Smyth, 2018;
Gaston & Holt, 2018; Navara & Nelson, 2007; Rich & Longcore,
2006). Recent global surveys of night sky brightness have concluded
that 23% of land surfaces between 75°N and 60°S, including 88%
of Europe and 47% of the United States, experience nightly “light
pollution” in the form of an increase in night sky brightness at least
8% above the natural level (Falchi et al., 2016) and that this artifi‐
cial illumination is gradually invading biodiversity hot spots (Guetté,
Godet, Juigner, & Robin, 2018). A meta‐analysis of night sky bright‐
ness over time has found regional increases ranging from 0% to 20%
per year, averaging 6% (Hölker, Moss, et al., 2010). Such increases
are expected to match the pace of urban expansion (Elvidge et al.,
2014). In the past two decades, however, some regions have experi‐
enced net decreases in night sky brightness, which may be due to en‐
vironmental and economic incentives to reduce ALAN, as well as to
technological innovations such as shielded lights that mitigate light
trespass by restricting artificial illumination to desired areas (Bennie,
Davies, Duffy, Inger, & Gaston, 2014, but see Kyba et al., 2017).
Received:17June2018 
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  Accepted :14July2018
DOI: 10.1002/ece3.4557
REVIEW
The impact of artificial light at night on nocturnal insects: A
review and synthesis
Avalon C. S. Owens | Sara M. Lewis
This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium,
provided the original work is properly cited.
© 2018 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
Department of Biology, Tufts University,
Medford, Massachusetts
Correspondence
Avalon C. S. Owens, Department of Biology,
Tufts University, Medford, MA.
Email: avalon.owens@tufts.edu
Funding information
Tufts University; Zoological Lighting
Institute
Abstract
In recent decades, advances in lighting technology have precipitated exponential in‐
creases in night sky brightness worldwide, raising concerns in the scientific commu‐
nity about the impact of artificial light at night (ALAN) on crepuscular and nocturnal
biodiversity. Long‐term records show that insect abundance has declined signifi‐
cantly over this time, with worrying implications for terrestrial ecosystems. The ma‐
jority of investigations into the vulnerability of nocturnal insects to artificial light
have focused on the flight‐to‐light behavior exhibited by select insect families.
However, ALAN can affect insects in other ways as well. This review proposes five
categories of ALAN impact on nocturnal insects, highlighting past research and iden‐
tifying key knowledge gaps. We conclude with a summary of relevant literature on
bioluminescent fireflies, which emphasizes the unique vulnerability of terrestrial
light‐based communication systems to artificial illumination. Comprehensive under‐
standing of the ecological impacts of ALAN on diverse nocturnal insect taxa will en‐
able researchers to seek out methods whereby fireflies, moths, and other essential
members of the nocturnal ecosystem can coexist with humans on an increasingly
urbanized planet.
KEYWORDS
artificial light at night, bioluminescence, fireflies, light pollution, nocturnal insects, visual
ecology
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The ecological effects of ALAN depend on both its intensity and
its spectral composition (as well as its flicker rate; see Inger, Bennie,
Davies, & Gaston, 2014 for a review). Historically, ALAN sources have
mainly comprised low‐ and high‐pressure sodium lamps, which emit
characteristic emission spectra concentrated in the yellow‐to‐orange
region of the visible spectrum, in addition to “whiter” broad‐spectrum
mercury vapor lamps, which also emit a large amount of UV radiation
(Figure 1). Most recently, environmental and economic concerns have
galvanized movements to replace these traditional, energy‐inten
sive light sources with energy‐efficient alternatives, primarily LEDs
(Davies, Bennie, Inger, & Gaston, 2013; Gaston, Davies, Bennie, &
Hopkins, 2012). LEDs can emit monochromatic light of any desired
wavelength within or adjacent to the visible spectrum; white light of a
given color temperature is produced by coating blue LEDs in a phos
phor material that absorbs a percentage of this light and re‐emits it in
longer wavelengths (Krames et al., 2007). As a result of this process,
most commercial white LED street lamps emit more of their light in
the blue region of the visible spectrum than do other ALAN types
(Figure 1). Exposure to blue light at night is known to cause insomnia
and increased disease risk in humans (American Medical Association,
2016). However, the effects of this widespread spectral shift on other
organisms have only recently attracted the attention of researchers
(Davies et al., 2017; Donners et al., 2018; Gaston, Visser, & Hölker,
2015; Justice & Justice, 2016; Lewanzik & Voigt, 2017; Longcore et
al., 2015, 2018 ; Pawson & Bader, 2014; Plummer, Hale, O’Callaghan,
Sadler, & Siriwardena, 2016; Somers‐Yeates, Hodgson, McGregor,
Spalding, & Ffrench‐Constant, 2013; Spoelstra et al., 2015; van
Grunsven et al., 2014; van Langevelde, Grunsven, Veenendaal, &
Fijen, 2017; Wakefield, Broyles, Stone, Jones, & Harris, 2016).
Worldwide, around 30% of vertebrates and more than 60% of in
vertebrates are nocturnal (Hölker, Wolter, Perkin, & Tockner, 2010).
These species are undoubtedly most vulnerable to artificial illumina
tion; they also tend to be underrepresented in the scientific literature.
To bring greater attention to the ecological consequences of human
activity in an often ignored temporal niche, herein we review known
individual‐level impacts of ALAN on nocturnal animals, organized
into the following five categories: temporal and spatial disorientation,
attraction, desensitization, and recognition. Temporal disorientation
covers both alterations in circadian clock s and photoperiodism as well
as the partitioning of activity between day and night (sensu Gaston,
Bennie, Davies, & Hopkins, 2013). To distinguish between discrete
effects of ALAN on nocturnal insect taxa, visual perception (sensu
Gaston et al., 2013) has been subdivided into the categories of recog
nition and desensitization; similarly, spatial orientation (sensu Gaston
et al., 2013) has been subdivided into spatial disorientation and at
traction. As the behavioral responses of individuals determine how
ALAN will affect populations, and then entire ecological communi
ties, we believe that this level of assessment is illuminating.
Recent studies have suggested that insect diversity and abundance
are both undergoing a rapid decline (Hallmann et al., 2017). Insects are
essenti al components of al l terrestrial fo od webs, and any loss es in insect
biomass are likely to have widespread ecological ramifications. ALAN
FIGURE 1 Spectral emission of different ALAN types. ALAN sources, such as incandescent and halogen bulbs, and mercury vapor lamps
emit large amounts of energy as infrared radiation (heat); mercury vapor and metal halide lamps also emit a non‐negligible amount of UV
radiation. Low‐pressure sodium lamps and LEDs are comparatively efficient ALAN sources, both capable of emitting nearly monochromatic
visible light (see text). Neutral (red) and cool (blue) temperature white LEDs have been plotted on the same graph for comparison. Modified
from Elvidge, Keith, Tuttle, and Baugh (2010) with permission
    
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affects insects in unique ways related to their body size and visual sys
tem. We have therefore chosen to focus our review around nocturnal
and crepuscular insects, although our classification may be applicable to
other organisms as well. Previous reviews cover the effects of ALAN on
plants (Bennie, Davies, Cruse, & Gaston, 2016), butterflies (Seymoure,
2018), and stream and riparian ecosystems (Perkin et al., 2011), as
well as the effects of artificial light regimes on pest insects (Johansen,
Vänninen, Pinto, Nissinen, & Shipp, 2011) and poultry (Van Nuffel, Bujis,
& Delezie, 2015); for general reviews of the impacts of AL AN on all spe
cies, see Rich and Longcore (2006) and Gaston et al. (2013).
Fireflies (Coleoptera: Lampyridae), click beetles (Coleoptera:
Elateridae), and glowworms (Diptera: Keroplatidae) are among the
most charismatic of all nocturnal insects, and their unique light‐based
communication system may make them especially vulnerable to arti
ficial illumination. We therefore conclude with a review and synthesis
of existing studies concerning the impact of ALAN on bioluminescent
insects, followed by recommendations for future research.
2 | INSECT VISION
As background for understanding the ecological effects of ALAN,
some familiarity with how light is measured, and how nocturnal
insects perceive light, is necessary. These topics are briefly intro
duced below; for a more in‐depth primer, see Cronin et al. (2014)
and Honkanen, Immonen, Salmela, Heimonen, and Weckström
(2017).
Light behaves both as a wave and as a particle (or photon).
Wavelength is measured in nanometers (nm), with wavelengths
between 400 and 700 nm corresponding to colors visible to hu
mans. Intensity (particle density) is measured in units of counts per
unit area (photons cm−2 s−1), with more photons corresponding to
greater intensity. Brightness, a subjective perception of intensity,
is influenced by individual spectral sensitivity, which is generally
viewed as fixed for a given species. In trichromatic humans, who
have blue‐, green‐, and red‐sensitive photoreceptors, luminance
sensitivity peaks around 555 nm under photopic (well‐lit) condi
tions (Figure 2a,b). Photon counts can be weighted by this func
tion to obtain measurements of brightness as perceived by the
human eye, given in units of lumens (lm). Light emitted from a point
source can be standardized to candelas, or lumens per solid angle
(cd = lm/sr), and light incident on a flat surface is converted to lux,
or lumens per area (lux = lm/m2). Although lux is widely used by
engineers and policymakers, and is easiest to measure, it poorly
approximates brightness as it is perceived by non‐human animals
(Longcore & Rich, 2004).
FIGURE 2 Spectral sensitivity in humans and insects. (a) Spectral sensitivities of blue (S, short‐wavelength), green (M, mid‐wavelength),
and red (L, long‐wavelength) sensitive photoreceptors in humans presented as 10° cone fundamentals, calculated from Stiles and Burch
(1959) color matching functions (Stockman & Sharpe, 2000; www.cvrl.org/). Humans are insensitive to UV wavelengths (<390 nm),
perceive visible light wavelengths (390–700 nm) as color, and perceive infrared wavelengths (>700 nm) as heat. (b) Human luminosity
function, used to predict relative “brightness” as perceived by humans. (c) Spectral sensitivity of the photoreceptors of honeybee workers,
thought to have retained the trichromatic color vision of ancestral insects (Briscoe & Chittka, 2001; modified from Peitsch et al., 1992). (d)
Electroretinography of a male Big Dipper firefly Photinus pyralis suggests that this species may have lost its short‐wavelength sensitive opsin
(modified from Lall, Chapman, Ovid Trouth, & Holloway, 1980)
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Ancestral insects likely possessed three types of photorecep
tors or opsins (Briscoe & Chittka, 2001; Figure 2c): one sensitive to
ultraviolet wavelengths (UV, 300–400 nm), another to short wave‐
lengths (blue, 400–480 nm), and a third to long wavelengths (green
to amber, 480–600 nm). Subsequent evolution into different eco‐
logical niches and light environments then selected for variation
in spectral sensitivity (Endler, 1993; Lall, Seliger, Biggley, & Lloyd,
1980; Théry, Pincebourde, & Feer, 2008). In butterflies, dragon‐
flies, and other diurnal insects, more visual opsins improve color
discrimination (Briscoe, 2008; Feuda, Marlétaz, Bentley, & Holland,
2016; Futahashi et al., 2015; Lunau, 2014; Sharkey et al., 2017). In
contrast, species occupying nocturnal and subterranean (aphotic)
habitats often lose one or more opsins, reducing their capacity for
color vision (reviewed in Feuda et al., 2016, Tierney et al., 2017;
but see Kelber, Balkenius, & Warrant, 2003; Meyer‐Rochow, 2007;
Somanathan, Borges, Warrant, & Kelber, 2008; White, Xu, Münch,
Bennett, & Grable, 2008).
In addition to reduced spectral sensitivity, nocturnal insects
often sacrifice spatial and temporal resolution in order to optimize
total visual sensitivity under low‐light conditions (but see Kelber et
al., 2011). Moths and some nocturnal beetles have evolved super‐
position eyes with rhabdoms that collect light from multiple facets,
sacrificing resolution to gain up to 1,000× more sensitivity than
apposition eyes of similar dimension (Cronin et al., 2014; Horridge,
1969). Some crepuscular bees, ants, and dung beetles have more re
cently transitioned to a nocturnal niche, and possess apposition eyes
with larger lenses and wider rhabdoms that sum photons over time
and/or space to increase sensitivity (Baird, Fernandez, Wcislo, &
Warrant, 2015; Warrant & Dacke, 2011). Among dung beetles, larger
eye size, smoother facets, and absence of screening pigment cor‐
relate to nocturnal activity (McIntyre & Caveney, 1998). Nocturnal
bees, wasps, and ants have dorsal ocelli—simple eyes used to detect
ambient light, movement, and/or orientation—that are significantly
larger than those of their diurnal relatives (Narendra & Ribi, 2017;
Somanathan, Kelber, Borges, Wallén, & Warrant, 2009; Warrant,
Kelber, Wallén, & Wcislo, 2006). The ocelli of nocturnal bees and
cockroaches are sensitive, but poor at perceiving fast movements;
they also lack UV‐sensitive opsins, perhaps as an adaptation to their
UV‐poor light environments (Berry, Wcislo, & Warrant, 2011); simi‐
larly, the ocelli of nocturnal ants may have lost polarization sensitiv‐
ity in response to the loss of polarized skylight signals (Narendra &
Ribi, 2017).
3 | EFFECTS OF ALAN ON NOCTURNAL
INSECTS
When considering the environmental impact of ALAN, it is vital
to distinguish between astronomical and ecological light pollution
(Longcore & Rich, 2004). The former, often referred to as “skyglow,”
is the result of upwelling illumination of the night sky. Skyglow may
spread to cover an area far beyond its origin and is not blocked
by local terrain (Gaston et al., 2015). In contrast, ecological light
pollution refers to the infiltration of point sources of light into habi‐
tats on t he ground, which c an affect loc al species with out necessaril y
influencing night sky brightness. Although shielded light technology
may ameliorate skyglow, this light may still impact local biodiversity.
Similarly, important is the position of insects relative to light sources;
downwelling light may affect species on the ground more than those
in the air, while upwelling light (path lights, tree lights, etc.) is more
likely to affect insects in flight.
To provide a comprehensive framework for understanding the
ecological impacts of artificial light on nocturnal insects, we have or‐
ganized the relevant literature into five categories, described below.
4 | TEMPORAL DISORIENTATION
ALAN may cause temporal disorientation, desynchronization of or
ganisms from their typical biorhythms. The vast majority of ter‐
restrial species have circadian, circamensual, and/or circannual
patterns of activity (foraging, reproduction, migration, etc.) that are
synchronized to daily, monthly, and yearly light cycles, respectively.
Specialized photoperiodic photoreceptors use external light signals
(Zeitgebers or “time‐givers”) to entrain the internal clock to the out‐
side environment (Numata, Miyazaki, & Ikeno, 2015; Numata, Shiga,
& Morita, 1997). In nocturnal insects, daily emergence time and
duration of feeding and courtship activity are dictated by internal
clocks entrained by ambient light or temperature (Saunders, 2009;
Tataroglu & Emery, 2014), while monthly or yearly cycles of eclo‐
sion, mating, and oviposition can be entrained by moonlight or day
length cues (for a review of lunar entrainment, see Kronfeld‐Schor
et al., 2013). If AL AN is sufficiently intense and/or sustained in time,
and of a specific spectral composition, it can desynchronize the in‐
ternal clock (reviewed by Saunders, 2012). For example, the pink
bollworm Pectinophora gossypiella can be artificially entrained by
monochromatic yellow light emitted by low‐pressure sodium (LPS)
lamps (Pittendrigh & Minis, 1971), and diurnal Bombus terrestris bum
blebees by UV illumination (Chittka , Stelzer, & Stanewsky, 2013).
Desynchronization reduces reproductive fitness of Drosophila mela
nogaster fruit flies (Xu, DiAngelo, Hughes, Hogenesch, & Sehgal,
2011, see also McLay, Nagarajan‐Radha, Green, & Jones, 2018) and
has the potential to disrupt vital biological processes in other taxa
(Dominoni, Borniger, & Nelson, 2016; Gaston et al., 2013; Saunders,
2012).
Exposure to artificial light in the laboratory inhibits a variety
of biological processes in nocturnal insects (e.g., Botha, Jones, &
Hopkins, 2017), possibly due to a buildup of melatonin (Honnen,
Johnston, & Monaghan, 2016; Jones, Durrant, Michaelides, & Green,
2015). In moths, constant light can inhibit female sex pheromone
release (Fatzinger, 1973; Geffen, Groot, et al., 2015; Sower, Shorey,
& Gaston, 1970), reduce male attraction (Geffen, Eck, et al., 2015),
induce male sterility (Bebas, Cymborowski, & Giebultowicz, 2001;
Giebultowicz, 2001; Riemann, Johnson, & Thorson, 1981), and dis
rupt female oviposition (Yamaoka & Hirao, 1981). Because ALAN
rarely completely conceals natural day–night cycles (Longcore &
    
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Rich, 2004; but see Honnen et al., 2016), it is unclear how these
findings apply to natural populations. If skyglow is sufficiently con‐
stant and intense, some animals may respond by adopting around‐
the‐clock activity patterns, as polar animals do during seasons with
continuous light (Bloch, Barnes, Gerkema, & Helm, 2013). Even low‐
level ALAN increases perceived day length (Kyba, Ruhtz, Fischer, &
Hölker, 2011a), and could thereby induce seasonal polyphenism in
stink bugs (Niva & Takeda, 2003) and aphids (Hardie, 2010; Sanders
et al., 2015), or alter the calling songs of katydids (Whitesell &
Walker, 1978). Low‐level ALAN is known to accelerate development
in a range of insect taxa (Kehoe, Cruse, Sanders, Gaston, & Veen,
2018, van Geffen, Grunsven, Ruijven, Berendse, & Veenendaal,
2014, but see Durrant, Botha, Green, & Jones, 2018), with varying
effects on fitness.
ALAN has been shown to conceal monthly and seasonal regimes
of lunar sk y brightness in urban areas (Davies, Bennie, Inger, Ibarra, &
Gaston, 2013; Kyba, Ruhtz, Fischer, & Hölker, 2011b), and this could
affect circamensual and circatidal rhythms of certain insect species
(but see Satoh, Yoshioka, & Numata, 2008). Some populations of the
mayfly Povilla adusta eclose 2 days after the full moon, which pro‐
vides increased visibility in which to court, copulate, and oviposit
(Corbet, Sellick, & Willoughby, 1974). Intertidal midges Clunio mari
nus (Neumann, 1989) and Pontomyia oceana (Soong, Lee, & Chang,
2011) use moonlight cues to synchronize activities (e.g., eclosion,
mating, and oviposition) during intertidal periods. The corn earworm
Helicoverpa zea will not mate unless its eyes are dark‐adapted to
ambient light below 0.05 lux, similar to that produced by a quarter
moon (Agee, 1969). Artificial light also suppresses H. zea oviposition,
perhaps because this behavior is synchronized to the lunar cycle
(Nemfc, 1971). Masking of lunar cycles by ALAN may disrupt these
vital circamensual rhythms.
In addition to disrupting natural rhythms, ALAN can desynchro‐
nize ecological interactions of mutualist species. Different species
have evolved to use either day length and temperature as Zeitgebers
(Bale et al., 2002; Jayatilaka, Narendra, Reid, Cooper, & Zeil, 2011).
Because ALAN alters day length but not temperature, it can lead to
ecological mismatches. For example, the nightly foraging activity of
the nocturnal carpenter bee Xylocopa tranquebarica is triggered by
twilight and coincides with the opening of night‐blooming flowers
(Somanathan et al., 2008). By delaying the onset of foraging in this
bee (a “phase shift”; Pittendrigh, 1993), ALAN may disrupt this vital
pollination mutualism, especially if flower opening is entrained by
ambient temperature instead of light (van Doorn & van Meeteren,
2003, see also Seymoure, 2018).
ALAN may also prolong the foraging activity of diurnal and/
or crepuscular competitors, which generally become more ac
tive with increasing illumination (Kempinger, Dittmann, Rieger,
& Helfrich‐Forster, 2009; Kronfeld‐Schor et al., 2013). Initiation
of foraging activity in moths (Dreisig, 1980; Eaton, Tignor, &
Holtzman, 1983; Riley, Reynolds, & Farmery, 1983) and crickets
(Campbell, 1976) is triggered when ambient light intensity declines
to a species‐specific level. Foraging activity of nocturnal and cre
puscular bees is both initiated and inhibited by specific light levels
(Dyer, 1985; Kelber et al., 2006). In dung beetle communities, tem
poral partitioning helps reduce competition. Each species is phys
iologically adapted to a particular temporal niche: Nocturnal dung
beetles have larger eyes and greater body size, which reduces
radiant heat loss on cold nights (Caveney, Scholtz, & McIntyre,
1995). Crepuscular insects that phase shift into nocturnal niches
may experience cold stress and could be slow to adapt (Urbanski
et al., 2012). In this way, the disconnect of temperature and light
environment may further disrupt ecological interactions by shift
ing species into temporal niches in which they experience greater
competition and/or lowered fitness.
5 | SPATIAL DISORIENTATION
ALAN may also result in spatial disorientation, disrupting an organ‐
ism’s ability to navigate in three‐dimensional space. In nocturnal
landscapes, the lack of visual information makes navigation difficult.
The most consistently visible landmarks are the moon and stars, fol
lowed by the dim pattern of polarized moonlight produced by at‐
mospheric filtering (Dacke, Byrne, Baird, Scholtz, & Warrant, 2011).
These orientation cues vary predictably throughout the night and
seasonally, and nocturnal insects often use the moon or stars to cal‐
culate navigational bearings (e.g., black carpenter ants, Klotz & Reid,
1993; earwigs, Ugolini & Chiussi, 1996; heart‐and‐dart moths, Baker,
1987; and harvester termites, Leuthold, Bruinsma, & Huis, 1976).
Sand hoppers use the moon to orient along beaches and are
known to orient to artificial fiber optic moons in the laboratory
(Ugolini, Boddi, Mercatelli, & Castellini, 2005). Other animals can
navigate by stars alone (reviewed by Foster, Smolka, Nilsson, &
Dacke, 2018). On moonless nights, Noctua pronuba yellow under
wing moths orient with respect to the north star (Sotthibandhu &
Baker, 1979), and Scarabaeus satyrus dung beetles use the Milky Way
as a cue to guide themselves away from dung piles in a maximally ef‐
ficient straight line (Dacke, Baird, Byrne, Scholtz, & Warrant, 2013).
On moonlit nights, S. satyrus and its relative Scarabaeus zambesianus
(Dacke, Byrne, Scholtz, & Warrant, 2004; Dacke, Nilsson, Scholtz,
Byrne, & Warrant, 20 03) navigate using polarized moonlight; other
nocturnal insects, including wasps (Warrant et al., 20 06), bees
(Greiner, Cronin, Ribi, Wcislo, & Warrant, 2007; Warrant & Dacke,
2011), and crickets (Herzmann & Labhart, 1989), may do so as well.
The bull ant Myrmecia pyriformis uses polarized moonlight in addition
to visual landmarks in the surrounding terrain to navigate to and from
its nest during nightly foraging trips (Narendra, Reid, & Raderschall,
2013; Reid, Narendra, Hemmi, & Zeil, 2011).
ALAN has potential to interfere with all these forms of noctur‐
nal navigation in two ways: Ecological light pollution introduces new
sources of light into the nocturnal landscape, which could be con
fused for the moon or stars (see Attraction for one example), while
atmospheric light pollution reduces the visibility of existing cues.
Skyglow dramatically reduces star visibility in urban areas (Falchi et
al., 2016), and artificial lighting has been shown to disrupt polariza
tion signals as well (Kyba et al., 2011b).
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Some nocturnal insects possess highly sensitive eyes capable of
color discrimination under minimal illumination, including the hawk
moth Deilephila elpenor (Kelber et al., 2003) and the nocturnal bee
X. tranquebarica (Somanathan et al., 2008). Both X. tranquebarica and
the nocturnal sweat bee M. genalis use optic flow and visual land‐
marks such as trees and flowers to navigate to and from their nests
(Baird, Kreiss, Wcislo, Warrant, & Dacke, 2011; Warrant et al., 2004).
Several nocturnal ants (Hölldobler & Taylor, 1983; Kaul & Kopteva,
1982; Klotz & Reid, 1993), bees (Warrant et al., 2006), and the shield
bug Parastrachia japonensis (Hironaka, Inadomi, Nomakuchi, Filippi,
& Hariyama, 2008) have been shown to navigate by canopy pat‐
terns, and the site‐specific shape of leaves and branches silhouetted
against the bright night sky overhead. Urban skyglow associated
with ALAN could increase this contrast, while artificial illumination
beneath the canopy is likely to erase it entirely.
Nocturnal insects such as wasps and bees use their enlarged
ocelli for navigation (Berry et al., 2011; Goodman, 1965), and direct
illumination may cause navigation problems for these and other spe‐
cies. Because these insects adjust their wing angle so that the lighter
half of their visual field is always overhead, upward‐directed illumi‐
nation could cause a maladaptive response. Indeed, light‐reflecting
mulching films are used in open crop fields to suppress the arrival
of alate aphids, thrips, and whiteflies (Shimoda & Honda, 2013).
Disoriented insects might wander into unsuitable habitat, and those
without the ability to “beeline” back to their nests at the end of each
night may fall prey to overheating, desiccation, or predation the next
day.
6 | ATTRACTION
Through positive phototaxis, many flying insects are attracted to
ALAN (Verheijen, 1960). Some exhibit characteristic spiraling flight
patterns, while others approach the light directly. Some orbit the
light source, frequently changing their angular velocity and direc
tion to remain within its vicinity (Muirhead‐Thompson, 1991), while
others perch on or under the light, apparently stunned. Physiological
and behavioral explanations of this phenomenon abound (see
Nowinszky, 2004), and their explanatory power varies with species.
The light compass theory (Baker & Sadovy, 1978; Sotthibandhu &
Baker, 1979) suggests insects that orient themselves by maintaining
a constant angle to light rays, historically emitted only by the moon
or stars, will spiral into artificial light sources. Other theories involve
the illusion of open sky (Goldsmith, 1990) or dark “Mach bands” at
light–dark borders (Hsiao, 1973), or disorientation due to “dazzling”
(Robinson, 1952, Verheijen, 1960, Hamdorf & Höglund, 1981; see
Desensitization below).
Historically, light traps have been used by scientists to survey
community composition, monitor beneficial insects (Nabli, Bailey,
& Necibi, 1999), and control insect pest populations (e.g., Goretti,
Coletti, Veroli, Giulio, & Gaino, 2011; Pawson, Watt, & Brockerhoff,
2009; Wallner & Baranchikov, 1992). The most common insect or‐
ders attracted to and captured in light traps are Diptera, Coleoptera,
and Lepidoptera (Mikkola, 1972; van Grunsven et al., 2014;
Wakefield et al., 2016). Light‐trapping equipment can differ from
ALAN in important ways: Experimental light traps usually emit more
short wavelengths, are often without glass shields (which filter UV),
and are placed near the ground (Degen et al., 2016). However, exper‐
iments that vary the intensity and spectral composition of light traps
can still offer insight into the potential effects of ALAN on positively
phototactic insects. The impact of ALAN on negatively phototactic
insects such as cockroaches and earwigs has not yet been well‐ex‐
plored (Bruce‐White & Shardlow, 2011, but see Farnworth, Innes,
Kelly, Littler, & Waas, 2018), despite a clear potential for adverse
effects (see Siderhurst, James, & Bjostad, 2006).
Among the common positively phototactic insects, moths (Frank,
1988, 2006 ; MacGregor, Pocock, Fox, & Evans, 2015) and aquatic
insects (Perkin, Hölker, & Tockner, 2014; Yoon, Kim, Kim, Jo, & Bae,
2010) are best studied. Comparative surveys have shown that, rel‐
ative to their calculated visibility, short wavelengths are dispropor‐
tionately attractive to many insects (Barghini & de Medeiros, 2012;
Mikkola, 1972; see also Wakefield et al., 2016 for a discussion of
infrared wavelengths). Although most insects can perceive short
wavelengths (Briscoe & Chittka, 2001; Kelber & Roth, 2006), cer‐
tain families of moths are more attracted to them than others (van
Langevelde, Ettema, Donners, WallisDeVries, & Groenendijk, 2011;
Somers‐Yeates et al., 2013, see also Wölfling, Becker, Uhl, Traub, &
Fiedler, 2016). LPS lamps rarely attract moths (Plummer et al., 2016;
Robinson, 1952; Rydell, 1992), even though most species can detect
the yellow wavelengths they emit (Briscoe & Chittka, 2001; Mikkola,
1972). Some nocturnal insects are disproportionately attracted to
polarized light sources as well (Danthanarayana & Dashper, 1986;
see Recognition below).
About 30%–40% of insects that approach street lamps die
soon thereafter (Eisenbeis, 2006), as a result of collision, overheat‐
ing, dehydration, or predation (Minnaar, Boyles, Minnaar, Sole, &
McKechnie, 2015; Yoon et al., 2010). The presence of foraging bats
does not repel moths from ALAN sources (Acharya & Fenton, 1999),
and under mercury vapor light, Operophtera brumata and O. fagata
moths lacked their normal evasive responses to simulated ultrasonic
bat signals (Svensson & Rydell, 1998). Depending on its placement,
ALAN may also impede the movement of insects among habitat
patches, lure them into bodies of water, or divert them into traffic
(Frank, 2006). Insects not killed immediately may become trapped in
a “light sink,” unable to forage (Langevelde, Grunsven, et al., 2017),
search for mates, or reproduce—especially when different sexes are
disproportionately attracted to ALAN, as is the case for many moth
species (Altermatt, Baumeyer, & Ebert, 2009; Altermatt & Ebert,
2016; Degen et al. , 2016; Frank, 1988; Garris & Snyd er, 2010; see also
Farnworth et al., 2018). Ecological traps that result in mortality or re‐
productive failure are predicted to lead to rapid population decline
and ultimately extinction (Kokko & Sutherland, 2001; Robertson,
Rehage, & Sih, 2013). Long‐term records confirm that positively
phototactic macro‐moths (Langevelde, Braamburg‐Annegarn, et al.,
2017) in lit habitats (Wilson et al., 2018) have undergone dispropor‐
tionate declines in abundance over the past 50 years.
    
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 7
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Perhaps due to selection, when newly eclosed moths from urban
populations are tested under standardized conditions, they are
less attracted to ALAN (Altermatt & Ebert, 2016). ALAN in urban
settings may also be generally less attractive due to a reduction in
background contrast (Frank, 2006), although one study comparing
declines in macro‐moth abundance at light‐trap sites with and with‐
out artificial night sky brightness did not support this suggestion
(Conrad, Warren, Fox, Parsons, & Woiwod, 2006, see also White,
2018).
7 | DESENSITIZATION
The highly sensitive visual systems of nocturnal insects may not
always function well in illuminated environments. Crepuscular and
nocturnal Myrmecia ants are capable of flexible, rapid light adaptation
(Narendra, Greiner, Ribi, & Zeil, 2016), and some even forage more
effectively under illumination (Narendra et al., 2013). However, the
photoreceptors of other insects, such as nocturnal flies, bees, and
cockroaches, are saturated at modest light levels (see Honkanen et
al., 2017). When exposed to too many photons at once, some insects
may be temporarily dazzled or even permanently blinded (Stark,
Walker, & Eidel, 1985). In Gryllus bimaculatus crickets, the photo
receptors of nocturnal adults show structural degeneration after
exposure to bright UV light, while those of diurnal nymphs are less
affected (Meyer‐Rochow, Kashiwagi, & Eguchi, 2002). If not bright
enough to cause permanent damage, a discrete ALAN source may
still temporarily blind a nocturnal insect to other orientation cues by
inducing a rapid process of light adaptation (see Laughlin & Hardie,
1978), perhaps causing it to fly directly into said light (McGeachie,
1988; Robinson, 1952; Verheijen, 1960). In the hawkmoth D. elpenor,
an 8‐s exposure to bright blue light can reduce visual sensitivity by
two to three orders of magnitude in minutes (Hamdorf & Höglund,
1981); even 0.125 s of white light exposure can effectively blind
cockroach ocelli for 15–20 s (Ruck, 1958). Should an affected insect
escape into darkness, it may take hours for it to completely recover
its original visual sensitivity (Bernhard & Ottoson, 1960). Light adap‐
tation can also have behavioral consequences: When moths encoun‐
ter yellow or green light above a certain brightness, light adaptation
of their eyes suppresses nocturnal behaviors, including flying, forag‐
ing, and mating, a response that has been successfully used in pest
control (Shimoda & Honda, 2013).
8 | RECOGNITION
The effects of environmental illumination on the ability of noc
turnal insects to recognize objects in their environment (con
specifics, predators, food plants, etc.) will depend on both the
wavelength and intensity of the ALAN source under considera
tion. Nocturnal insects may be able to detect nearby objects more
easily if the source emits (at sufficient intensity) wavelengths to
which their visual systems are sensitive. However, some taxa may
gain or lose their ability to discriminate colors, depending on the
range of wavelengths emitted (Davies, Bennie, Inger, Ibarra, et al.,
2013, but see Johnsen et al., 2006). As a result, ALAN has the
potential to impede visual signaling and/or undermine camouflage
(reviewed by Delhey & Peters, 2017). The body color of dusk‐
active beetles is most apparent in a purplish light environment
(Endler, 1993; Théry et al., 2008) and may become less visible to
conspecifics under broad‐spectrum ALAN illumination. Similarly,
the aposematic coloration of Heliconius butterfly wings, which is
especially apparent in their typical light environments, may be ob
scured by AL AN (Seymoure, 2016). The UV emissions of mercury
vapor lamps accentuate UV‐reflective markings on flowers and
wings, which may benefit bees, moths, and other nocturnal in
sects sensitive to these signals (Kevan, Chittka, & Dyer, 2001). In
contrast, illumination by LPS lamps could obscure these markers
(Frank, 2006). Many aquatic insects use polarized light to locate
suitable oviposition sites (reviewed by Horváth, Kriska, Malik, &
Robertson, 2009, Perkin et al., 2014, Villalobos Jiménez, 2017).
Artificial illumination of smooth dark surfaces such as asphalt
simulates the polarization of light reflected off bodies of water,
causing some aquatic insects to maladaptively oviposit on bridges
or cars (Egri et al., 2017; Szaz et al., 2015), or to congregate on
windows (Horváth et al., 2009; Kriska, Malik, Szivák, & Horváth,
2008).
Increased visibility is thought to benefit predators over prey
(Kronfel d‐S chor et al., 2013; Youthed & Moran, 1969), but evidence of
this phenomenon in arthropod systems is mixed (see Grenis, Tjossem ,
& Murphy, 2015; Skutelsky, 1996; Tigar & Osborne, 1999). By simu
lating bright moonlit nights every night of the month (Davies, Bennie,
Inger, & Gaston, 2013), ALAN may reduce foraging time and increase
the starvation risk of nocturnal prey insects (Schmitz, Beckerman, &
O’Brien, 1997). Conversely, it may prolong the foraging activity of
diurnal and crepuscular insects such as the sweat bee Lasioglossum
texanum (Kerfoot, 1967), the desert ant Veromessor pergandei (Hunt,
1977), and the hemipteran Nilaparvata lugens (Riley, Reynolds, &
Farrow, 1987), all of which engage in nocturnal foraging during the
full moon.
9 | COMMUNITY‐LEVEL IMPACTS
By altering the light environment experienced by nocturnal in‐
sects, ALAN can disrupt their temporal patterns, interfere with
their spatial orientation, act as a fatal attraction, reduce their vis‐
ual sensitivity, and alter foraging activity and species interactions.
When populations of such abundant and ubiquitous organisms are
disrupted (Gaston & Bennie, 2014; Kurvers & Hölker, 2015), entire
communities will be affected (Davies et al., 2017; Davies, Bennie,
Inger, Ibarra, et al., 2013; Sanders & Gaston, 2018). The population‐
and community‐level effects of ALAN have been relatively under‐
studied, despite their great potential for use in predicting the future
composition of artificially illuminated habitats; we summarize exist‐
ing research below.
8 
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   OWENS aNd LEWIS
Insects that are attracted to ALAN sources are readily ex
ploited by predators: Orb‐weaving spiders prefer artificially lit
web sites (Enders, 1977), which may net them more prey (Heiling,
1999, but see Yuen & Bonebrake, 2017). Bats ( Jung & Kalko, 2010;
Minnaar et al., 2015; Rydell, 2006), birds (Robertson, Kriska,
Horvath, & Horvath, 2010), and invasive cane toads (González‐
Bernal, Greenlees, Brown, & Shine, 2016) congregate around
streetlights and lit buildings for a similar reason. Usually, diurnal
anole lizards and jumping spiders have been observed hunting for
insects at night in artificially illuminated locations (Frank, 2009;
Garber, 1978; Wolff, 1982). Moths frozen under illumination may
provide stable search images that enable birds to recognize the
camouflaged wing patterns of these species in other contexts
(Frank, 2006). Decreases in moth abundance have a negative im
pact on nocturnal pollen transport (Fox, 2013; Frank, 1988; Knop
et al., 2017; Macgregor et al., 2015; Macgregor, Evans, Fox, &
Pocock, 2017), with cascading effects on populations of plants
and insect herbivores.
Artificial illumination in urban areas may reduce the population
persistence of nocturnal species by preventing movement between
habitat patches (Farnworth et al., 2018; Gaston & Bennie, 2014;
Guetté et al., 2018). For example, moths attempting to cross road
networks were impeded by a line of closely spaced street lights
(Degen et al., 2016). In a riparian ecosystem, emerging aquatic in
sects were drawn to artificially illuminated patches rather than the
surrounding habitat, potentially reducing nutrient exchange and
species dispersal (Manfrin et al., 2017). At the same time, illumi
nation of the water may increase predation risk for invertivorous
fish, reducing predation on aquatic insects and leading to locally
increased insect abundance (Manfrin et al., 2017). Riparian preda
tors respond to increased prey availability by congregating around
light sources (Meyer & Sullivan, 2013; Perkin et al., 2011).
Several studies have noted an influx of predatory and scaveng
ing arthropods into lit areas (Davies, Bennie, & Gaston, 2012; Šustek,
1999), though this response appears to be taxon‐specific (Eccard,
Scheffler, Franke, & Hoffmann, 2018; Manfrin et al., 2017; Meyer &
Sullivan, 2013; van Grunsven, Jähnichen, Grubisic, & Hölker, 2018).
Broad‐spectrum LED lights in combination with urban heat reduced
pea aphid populations by increasing visibility and lengthening the
activity period of their visually oriented coccinellid predators (Miller
et al., 2017); however, in similar experiments, bright illumination
decreased or did not affect rates of parasitism by parasitoid wasps
(Kehoe et al., 2018; Sanders et al., 2015, see Sanders, Kehoe, Cruse,
Veen, & Gaston, 2018). The introduction of artificial light and noise
causes parasitic frog‐biting midges to be unable to locate and feed
from their túngara frog hosts (McMahon, Rohr, & Bernal, 2017).
Herbivorous insects may additionally be affected by changes in their
food plants (reviewed by Vänninen, Pinto, Nissinen, Johansen, &
Shipp, 2010). For example, long‐wavelength ALAN reduced the pop
ulation sizes of pea aphids by inhibiting flowering in their host plant
via the phy tochrome pathway (Bennie, Davies, Cruse, Inger, & Gaston,
2015), and light from high‐pressure sodium (HPS) lamps increased
plant toughness and decreased the mass of cutworm larvae (Grenis
& Murphy, 2018). The resultant absence of midges, pea aphids, and
other prey insects in urban areas is likely to have widespread effects
on populations of their host plants, pollinated plants, and predators.
10 | EFFECTS OF ALAN ON
BIOLUMINESCENT INSECTS
Firefly beetles (Coleoptera: Lampyridae) comprise the most wide
spread and diverse group of bioluminescent species on land. The
approximately 2000 lampyrid species, all of which glow aposemati‐
cally as larvae (Branham & Wenzel, 2003), enjoy a worldwide dis‐
tribution. In many adult fireflies, either one or both sexes employ
bioluminescence as a courtship signal (Lloyd, 2008). Other biolumi‐
nescent insect taxa, primarily click beetles (Coleoptera: Elateridae),
railroad worms (Coleoptera: Phengodidae), and fungus gnats
(Diptera: Keroplatidae), employ bioluminescence as an aposematic
signal or predatory lure (Meyer‐Rochow, 2007, Redford, 1982). All
of these essential signals may be masked by ALAN, which has been
suggested as one of several factors contributing to a worldwide de‐
cline in firefly populations (Khoo, 2014; Lewis, 2016; Lloyd, 2006).
Recent studies on bioluminescent ostracods (Gerrish, Morin, Rivers,
& Patrawala, 2009) and fungus gnats (Merritt & Clarke, 2013; Mills,
Popple, Veidt, & Merritt, 2016) have shown inhibitory effects of ar‐
tificial light on signaling activity, raising similar concerns. In this sec‐
tion, we briefly discuss firefly visual ecology then use our framework
to examine how ALAN may affect the courtship and reproductive
success of these and other bioluminescent insects.
11 | FIREFLY VISION AND
BIOLUMINESCENCE
Bioluminescent fireflies employ a diverse range of courtship sign
aling systems as part of their sexual communication (Lewis, 2009;
Takatsu, Minami, Tainaka, & Yoshimura, 2012). In some taxa, sed‐
entary females produce long‐lasting glows that attract flying males,
the latter sometimes incapable of producing light. In other taxa,
including most North American species, both sexes use flash sig‐
nals—discrete bursts of light—to communicate with potential mates.
North American Photinus fireflies engage in courtship dialogs that
involve precisely timed flash signals encoding species identity and
sex. Typically, sedentary females respond to advertisement flashes
emitted by flying males. In congregating South‐East Asian species,
however, clusters of stationary males emit synchronous flashes to
attract flying females.
Bioluminescence can be a highly efficient visual signal: against
a black background, its contrast is effectively infinite, and the dis‐
tances across which it can be perceived limited only by habitat struc‐
ture and the visual sensitivity of the receiver (Cronin et al., 2014).
However, fireflies do not always signal against a black background.
While nocturnal species initiate courtship flashing long after night
fall, crepuscular species become active shortly after sunset (Lloyd,
    
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 9
OWENS aNd LE WIS
1966). Some species flash in shady patches during the daytime
(Vencl, Luan, Fu, & Maroja, 2017). The color of firefly biolumines‐
cence varies as follows: Among crepuscular species, flashes are
generally yellower compared to the greener flashes emitted by noc
turnal fireflies (Seliger, Lall, Lloyd, & Biggley, 1982); this is thought
to maximize signal contrast against the green foliage dominating the
background at dusk. However, signal color also shows intraspecific
variation, perhaps related to differences in habitat typ e (Hall, Sander,
Pallansch, & Stanger‐Hall, 2016).
Due to their temporally restricted courtship activity periods,
fireflies are highly sensitive to ambient light cues indicating time
of day. When ambient light intensity descends to a certain species‐
specific threshold, courtship signaling begins (Buck, 1937a; Table 1).
Intracerebral ocelli described from Japanese Luciola cruciata and
TABLE 1 Effect of ambient light on flash activity of various firefly speciesa
Species Flash (lux) No flash (lux) Location References
Aspisoma lineatum 0.85 Southeast Region, Brazil Hagen and Viviani (2009)
>0.05 Hagen et al. (2015)
Apisoma physonotum <0.2 Hagen and Viviani (2009)
>0.05 Hagen et al. (2015)
Api som a sp2 <0.2 Hagen and Viviani (2009)
Apisoma sp4 <0.2 Hagen and Viviani (2009)
Bicellonychia lividpennis 4.5 Hagen and Viviani (20 09)
Bicellonychia ornaticollis <0.2 Hagen and Viviani (2009)
Cratomorphus concolor <0.2 Hagen and Viviani (2009)
Cratomorphus sp4 <0.2 Hagen and Viviani (2009)
Lampyris noctiluca ()1.3 10 Ebeltoft, Denmark Dreisig (1975)
0.28 Dreisig (1975)
Lampyris noctiluca (larva) 6.85 Dreisig (1975)
Luciola italica 0.25 0.47 ± 0.26 Turin, Italy Picchi, Avolio, Azzani,
Brombin and Camerini
(2013)
Photinus interdius (diurnal) 339.82 ± 88 >1,000 Darien Province, Panama Vencl, Luan, Fu, and
Maroja (2017)
Photinus pyralis 210–320 Laboratory Buck (1937a)
301.24 ± 89.07 Clarke County, Virginia, USA Firebaugh and Haynes
(2016)
Photinus sp1 <0.2 >0.234 Southeast Region, Brazil Hagen et al. (2015)
>1. 5 Hagen and Viviani (2009)
Photinus spp. 1.2 Piedmont Region, Maryland,
USA
Costin and Boulton (2016)
Photinus umbratus 1Highlands County, Florida, USA Dreisig (1975)
4Dreisig (1975)
Photuris "A" 0.38 Dreisig (1975)
Photuris congener 0.25 2.5 Dreisig (1975)
Photuris pennsylvanica 160,000 Laboratory Harvey (1926)
Photuris missouriensis 3,800 Laboratory Case and Trinkle (1968)
Photuris versicolor ()Highlands County, Florida, USA Dreisig (1975)
301.24 ± 89.07 Clarke County, Virginia, USA Firebaugh and Haynes
(2016)
Pteroptyx maipo 0.2–0.3 Tin Shui Wai, Hong Kong Yiu (2012)
Pteroptyx valida 7–14 Samut Prakan Province, Thailand Prasertkul (2018)
Pyrogaster moestus >0.05 Southeast Region, Brazil Hagen et al. (2015)
Pyrogaster sp1 <0.2 Hagen and Viviani (2009)
a“Flash” column gives ambient light levels shown to be dim enough to induce bioluminescence for each species; “No flash” gives ambient light levels
shown to inhibit firefly flash activity.
10 
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   OWENS aNd LEWIS
Luciola lateralis fireflies may assist in entraining this circadian be
havior (Hariyama, 2000). Lampyris noctiluca females usually begin
emitting courtship signals when ambient light levels fall below 1.3
lux and are never active above 15 lux; for Photuris congener males,
these values are 0.25 and 2.5 lux, respectively, reflecting the
shorter duration of twilight in their habitat (Dreisig, 1975). Ambient
light intensity also affects other aspects of courtship behavior. In
crepuscular Photinus species, males fly higher as light levels decline,
and when passing under the shade of trees (Lewis & Wang, 1991;
Lloyd, 2000).
The superposition compound eyes of fireflies are finely attuned
to conspecific signals: They are highly sensitive, but only to a narrow
range of wavelengths. To date, only UV‐ and long‐wavelength‐sen‐
sitive (UVS and LWS) opsin genes have been isolated from fireflies
(Martin, Lord, Branham, & Bybee, 2015; Sander & Hall, 2015), al‐
though blue sensitivity has been observed in both North American
and Japanese species using electroretinography (ERG) recordings
(Eguchi, Nemoto, Meyer‐Rochow, & Ohba, 1984; Lall, Lord, & Ovid
Trouth, 1982; Lall, Strother, Cronin, & Seliger, 1988). The peak sen‐
sitivity of the LWS photoreceptor generally corresponds to the peak
waveleng th of signals produ ced by that species (Sa nder & Hall, 2015).
Signal reception is further tuned using filter pigments: In species
that emit yellow bioluminescence, reddish filter pigments narrow
visual sensitivity to the yellow region of the spectrum by blocking
out green wavelengths (Booth, Stewart, & Osorio, 2004; Cronin,
Järvilehto, Weckström, & Lall, 2000). This screening could allow
some lampyrids to communicate visually even within light‐polluted
habitats. However, current evidence suggests that AL AN does have
a demonstrable impact on firefly signaling (Table 2).
Studies in urban and rural regions in São Paulo, Brazil, have
found that several lampyrid species are limited to areas with am
bient light levels below 0.2 lux, approximately equivalent to that of
the night sky during a full moon (Hagen & Viviani, 2009; Viviani,
Rocha, & Hagen, 2010). However, the crepuscular species Aspisoma
lineatum and Bicellonychia lividipennis were found signaling near so
dium vapor lamps, in areas with illumination measuring 0.85 and
4.5 lux, respectively; note that B. lividipennis emerges early in the
evening when ambient light levels (without ALAN) reach about 4.5
lux. Similarly, surveys of Luciola italica in Turin, Italy, have found
populations concentrated in dimly illuminated areas, with a negative
correlation between firefly abundance and certain indices of urban
ization (Picchi et al., 2013). In descriptive studies such as these, the
effects of ALAN are confounded by many other variables associated
with urbanization. It is also unclear whether population loss occurs
due to movement away from illuminated habitats or to reduced re
productive success within urban populations.
Below, we summarize all available evidence concerning the
impact of ALAN on fireflies, organized according to the five cate
gories of impact described earlier. Many of these findings may be
applicable to all bioluminescent insects, although data on other
taxa are sparse. Should these insects be unable to cope with the
following challenges, their survivorship, foraging success, and
mating success—and therefore population persistence—will be
reduced.
TABLE 2 Effect size of experimental studies examining impact of ALAN on firefly courtship
Species
Treatment (ALAN
type) Metric Intensity (lux)
Effect size
(G's Δ)Study
Lampyris noctiluca High‐pressure
sodium street lamps
Green LED lure (# of trapped
males)
L1: 46–64 L1:−0.74 Ineichen and
Rüttimann (2012)
L2: 0.1–0.4 L2:−0.97
Lampyris noctiluca Incandescent
flashlight
Green LED lure (# of trapped
males)
L1: 0.3 L1:−0.37 Bird and Parker (2014)
L2: 0.18 L2:−0. 37
L3: 0.09 L3:−0. 23
L4: 0.07 L4:−0.02
Photinus sp1 Multi‐metal vapor
floodlights
Transect count (# of flashing
individuals)
T1: 4.45 T1:−0. 25 Hagen et al. (2015)
T2: 1.5 T2:−0.22
T3: 0.05 T3:−0.17
Photuris versicolor White LED
floodlights
Flash count (flashes/min) 301 at plot
center
−0. 459 Firebaugh and Haynes
(2016)
Photinus pyralis Flash count (flashes/min) 301 at plot
center
NS
Flash count, tethered females
(15 min total)
167.21 on
average
−0 .617
Photinus spp. Mercury vapor bulb Flash count (flashes/min) 1.2 at plot edge −0.653 Costin and Boulton
(2016)
Note. Because all studies involve comparison of groups of the same size (e.g., firefly populations before and after ALAN exposure), Glass’s Δ is an ap‐
propriate estimate of effect size, as it uses only the standard deviation of the control group (Kline, 2013). NS indicates that no significant ef fect on
firefly courtship activity was observed.
    
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 11
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11.1 | Temporal disorientation
ALAN has the potential to disrupt both larval and adult activity
cycles, but little is known about the magnitude of this impact.
Pyractomena borealis larvae kept indoors under continuous light
pupated several months earlier than anticipated (Lloyd, 2006),
but whether this occurs in natural populations remains unclear.
Although adult fireflies rarely live long enough to express circa
mensual rhythms of activity, L. noctiluca larvae in the field have
been shown to hide during the full moon (Gunn & Gunn, 2012),
and may reduce their foraging activity if night sky brightness is
increased by skyglow.
Nightly onset of courtship activity by adult fireflies is deter
mined by ambient light intensity. P. pyralis adults begin flashing
earlier in the evening on cloudy days (Buck, 1937a), although the
opposite is predicted for urban areas, where reflection of AL AN
makes the sky brighter on cloudy days than on clear ones (Kyba
et al., 2011a). In light‐polluted habitats, the flight periods of cre
puscular species might be extended into those normally occupied
by nocturnal species. If males or females are responsive to het
erospecific signals, this could impact their reproductive success
by reducing dialog efficiency and/or increasing the frequency of
heterospecific matings. Nocturnal Photuris fireflies are a major
predator of other firefly species (Eisner, Goetz, Hill, Smedley, &
Meinwald, 1997; Eisner, Wiemer, Haynes, & Meinwald, 1978;
Lloyd, 1997). If ALAN causes crepuscular species to extend their
courtship into the nocturnal niche occupied by Photuris, they may
experience higher predation rates. It has been posited that, over
evolutionary time, predation by nocturnal Photuris fireflies drove
some fireflies into crepuscular and diurnal niches (Deyrup et al.,
2017; Gronquist et al., 2006).
11. 2| Spatial disorientation
ALAN may disorient fireflies that navigate with respect to the sun
or moon, such as the lar vae of P. borealis, which choose an aerial pu‐
pation site located on the southern side of trees to maximize their
exposure to direct sunlight (Gentry, 2003). However, little is known
about the balance of positive and negative phototaxis in fireflies. In
Photinus fireflies, males produce courtship signals while searching
within their habitat, and immediately orient toward a female when
they detect her response flash. Because these fireflies do not appear
capable of distinguishing between small signals that are nearby and
large signals farther away (Cratsley & Lewis, 2003), they could pos‐
sibly mistake artificial lights with certain emission spectra for recep
tive conspecifics (see below).
11.3| Attraction
In the European glowworm L. noctiluca, flying non‐bioluminescent
males are attracted to larviform females that glow steadily from
perches on raised display sites. Numerous experimental studies
conducted with this species have employed glowing light lures to
elucidate their signaling system (e.g., Bird & Parker, 2014; Mikkola,
1972; Schwalb, 1961). By measuring the attraction of males to dif‐
ferent LEDs or chemiluminescent lures, these studies have described
male preferences for the color and spatial pattern of female glow
signals, and have demonstrated male attraction to light traps that
emit yellow‐green light (reviewed by De Cock, 2009).
Although L. noctiluca is geographically widespread in Europe,
its populations have been declining for some time (Gardiner,
2011). One potential contributing factor could be that males are
more attracted to certain point sources of ALAN than they are
to conspecific female glow signals. In North Wales, Bek (2015)
reported significant attraction of male L. noctiluca to LPS street
lamps, compared to HPS, LED, or unlit lamps: The vast majority of
males (556 of 564) were found under LPS lamps, while none were
found below LED lamps or those that had been switched off. A
notable difference between LPS and LED bulbs is that the latter
emit a large percentage of their light in blue wavelengths; binary
choice experiments on L. noctiluca suggest that the addition of
blue wavelengths significantly decreases male attraction to light
lures (Booth et al., 2004).
Though L. noctiluca is not representative of all firefly spe‐
cies, other glowworms may also be attracted to artificial lights. In
Uganda, Bowden and Church (1973) found multiple Lamprigera
specimens caught in a Robinson trap illuminated by a mercury vapor
bulb. In Rwanda, Diaphanes males were collected from a light trap
emitting red light (Pacheco, Martin, & Bybee, 2016). Pleotomus and
Pleotomodes glowworms are also attracted to light traps, including
those exclusively emitting UV wavelengths (Faust, 2017; Lloyd,
2006).
Beyond glowworms, other firefly species could also be attracted
to artificial light sources. Male Pteroptyx fireflies in South‐East Asia
congregate in particular display trees and flash together en masse, and
may be attracted to flashing string lights imitating these mating con
gregations (Cratsley, Prasertkul, & Thancharoen, 2012; Thancharoen,
Srinual, & Laksanawimol, 2017). If these light sources draw individuals
away from traditional display sites or differentially attract males and
females, they could disrupt Pteroptyx courtship and reduce mating
success.
11.4 | Desensitization
Due to their highly sensitive visual systems (Horridge, 1969),
fireflies may be vulnerable to blinding by bright ALAN sources.
Numerous previous studies have used ERG recordings to meas
ure firefly spectral sensitivity, by observing the level of electrical
activity in the eye triggered by exposure to small point sources
of light (e.g., Lall, 1981; Lall et al., 1982, 1988; Lall, Chapman, et
al., 1980; Lall & Lloyd, 1989). These studies show that the com
pound eyes of Photinus fireflies can take several hours to become
fully dark‐adapted (Lall, 1993); intermittent light exposures may
decrease resulting gains in visual sensitivity, but the duration and
intensity of exposure sufficient to cause a complete reversal to
the light‐adapted state, let alone to cause dazzling and/or blinding,
12 
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   OWENS aNd LEWIS
have yet to be investigated (A. Lall, pers. comm.). Nonetheless,
it appears likely that bright sources of ALAN such as LED street
lamps could at minimum slow the dark adaptation process. This
may disproportionately diminish long‐wavelength sensitivity in
female fireflies (Oba & Kainuma, 2009), reducing their ability to
recognize potential mates.
11. 5| Recognition
While some background illumination is often present in their sign‐
aling milieu, especially for crepuscular firefly species, artificial il
lumination could reduce courtship success by interfering with the
perception of male signals by receptive females, or vice‐versa. In
addition, females might be less responsive to male signals that they
perceive as dimmer under ALAN. Lloyd (2006) suggested that mon‐
ochromatic yellow light from LPS lamps might disproportionately
impact the signal exchange of yellow‐flashing (crepuscular) species.
Because broad‐spectrum illumination from white LED street lamps
includes many wavelengths, these ALAN sources may obscure the
bioluminescent signals of green‐flashing (nocturnal) firefly species
as well.
Several recent field studies have examined how ALAN affects
firefly flash activity and courtship behavior (Table 2). Ineichen and
Rüttimann (2012) assessed the impact of HPS street lighting on
L. noctiluca males in suburban Switzerland. They compared male
attraction to LED lures simulating the glows of conspecific females,
placed either under (46–64 lux) or between (0.1–0.4 lux) HPS street
lamps. Significantly, more males were attracted to lures in darker
locations between the HPS street lamps. Perhaps males were un
able to detect the lures in the highly illuminated surroundings, or
perhaps males prefer signals with greater contrast against the back
ground (Hopkins, Baudry, Candolin, & Kaitala, 2015). In either case,
these results suggest HPS street lighting interferes with L. noctiluca
courtship communication. Ineichen and Rüttimann (2012) noted
that female display sites appeared to be uniformly distributed with
respect to street lighting. L. noctiluca females select their display
sites during daytime as larvae and rarely relocate in their flightless
adult form (Tyler, 2013). Thus, females located underneath ALAN
sources are likely to experience lower mating success.
In another observational study on a university campus in
Sorocaba, Brazil, Hagen, Santos, Schlindwein, and Viviani (2015)
found that courtship flashing by several firefly species was affected
by multi‐metal vapor spotlights illuminating an outdoor sport court.
On nights when the spotlights were turned on, flash activity by
the most common species, Photinus sp1, was significantly reduced.
ALAN reduced the number of flashing individuals encountered along
transects that were both directly and indirectly illuminated by the
spotlights; Photinus sp1 flashed only below measured light intensi‐
ties of 0.23 lux. While this and other studies clearly demonstrate
that ALAN reduces flash activity (Table 1), its impact on firefly mat‐
ing success remains unknown.
Experimental field studies of ALAN are useful for removing
confounding variables associated with urbanization. Working in an
undisturbed British chalk grassland, Bird and Parker (2014) intro
duced a point source of light and then measured attraction of L. noc‐
tiluca males to glowing LED lures. Lures were place d 0.5, 1.0, 1.5, and
2 m from an upward‐directed filament bulb flashlight (its emission
spectra resembling that of a mercury vapor lamp) with measured
intensities of 0.3–0.07 lux. Compared to a non‐illuminated control,
levels of ALAN as low as 0.09 lux significantly reduced the number
of males attracted to simulated female glows. When ambient illumi‐
nation was brighter than 0.18 lux, none of the males approached the
simulated females.
Although both studies showed that ALAN can interfere with the
ability of male L. noctiluca to locate females, Bird and Parker (2014)
found effects at much lower ALAN levels compared to Ineichen and
Rüttimann (2012). Several explanations are possible. Bird and Parker
attributed this difference to the orientation of the ALAN source,
suggesting that upward‐directed light is more likely to dazzle males
or reduce trap visibility than downward‐directed street lamps. The
studies also differed in ALAN type as well as design of the female
lures. Furthermore, urban and suburban firefly populations may
have adapted to cope with higher levels of ALAN. Male choice is
another potential factor: In both studies, males chose the compara
tively brightest lures of those presented.
Firebaugh and Haynes (2016) conducted an experimental study
of the impact of ALAN on the flash activity of both male and female
fireflies at a rural site in Virginia, USA. They established four pairs
of 20‐m‐diameter plots, placing a downward‐facing LED floodlight
(301 lux at plot center) at the center of each plot. The flash activ‐
ity of Photuris versicolor, a nocturnal species, declined significantly
in the illuminated plots compared to non‐illuminated control plots.
However, no difference in flash activity was detected for males of
P. pyralis, a crepuscular species. In addition, AL AN did not alter the
timing of the nightly onset of P. pyralis flash activity.
In a separate experiment, the authors tethered P. pyralis females
to display platforms and observed the impact of LED floodlights (ap‐
proximately 167 lux) on male approach frequency and flash count,
as well as on the frequency of female response flashes. Although
ALAN again had no significant effect on the flash activity of P. pyra‐
lis males, it significantly reduced the female response rate. This dif‐
ferential effect on male and female flash behavior could be due to
the downward directionality of the ALAN source. Flying males may
be able to detect a female flashing below them, even against more
brightly illuminated vegetation. However, females perch close to the
ground and look upward to detect males, typically against a dark sky.
Therefore, the females may have been unable to detect male sig‐
nals viewed against the downward‐facing ALAN. Alternatively, since
P. pyralis females are known to prefer brighter simulated flashes
(Vencl & Carlson, 1998), they may be less responsive to male signals
that, when set against an illuminated background, appear less bright.
In an experimental field study in rural Maryland, USA, Costin and
Boulton (2016) introduced ALAN using a mercury vapor bulb (1.2
lux at plot edge). At each of the six sites starting 30 min after sun‐
set, they counted firefly flashes for 30 min one night, then added
ALAN, and repeated the count the next night. On nights when
    
|
 13
OWENS aNd LE WIS
ALAN was introduced, the authors found a significant decline in
flash rates. Multiple species were present at each site, with the cre‐
puscular Photinus marginellus most frequently observed. In contrast,
Firebaugh and Haynes (2016) found no effect of ALAN on flash ac‐
tivity of P. pyralis. The discrepancy between these studies could re
flect differing emission spectra of their artificial light sources, and/
or differences between firefly species. Although both are crepuscu
lar, P. marginellus emerges slightly later in the evening (Lloyd, 1966)
and might be more sensitive to ALAN than P. pyralis. Because control
counts were always made on the night preceding ALAN treatment
at each site, the results may also reflect temporal changes in firefly
abundance.
Yiu (2012) performed a similar experiment on synchronously
flashing males of a Hong Kong firefly Pteroptyx maipo, introducing a
compact fluorescent lamp for several minute intervals and counting
flashes before and during exposure. AL AN illumination (0.21–2 .0 lux)
significantly decreased the average flash frequency of this species,
and these decreases were reversed when the lamp was switched off.
However, during a survey of Pteroptyx malaccae and Pteroptyx valida
in Thailand, Prasertkul (2018) found robust congregations of both
species flashing within close proximity to white fluorescent street
lamps (up to 7–14 lux), suggesting that ALAN does not prevent ag‐
gregation or courtship flashing in these species.
In contrast to the complex light environments and shifting con‐
ditions typical of field experiments, laboratory experiments allow
for more precise manipulation of ambient light levels. Early light
exposure experiments conducted in the laboratory demonstrated
that sufficiently bright light (from 50 to 1,000 lux) completely in‐
hibits bioluminescent signaling in several firefly species (Table 1).
More recently, Thancharoen (2007) exposed pairs of the Thai firefly
Sclerotia (formerly Luciola) aquatilis to different intensities of fluores‐
cent lighting (0.05, 0.1, 0.2, and 0.3 lux) and observed their mating
behavior. Artificial illumination prolonged courtship, mounting, and
mating duration in this species. Though all pairs ultimately mated
successfully in the laboratory, ALAN could reduce mating success
under field conditions by increasing the difficulty of locating a mate.
Using monochromatic LEDs, Owens, Meyer‐Rochow and Yang
(2018) investigated the impact of ambient light on the alarm flash
behavior of an aquatic Taiwanese firefly, Aquatica ficta. Short‐wave
length light (444–533 nm) caused males to flash more brightly, but
less frequently, while long‐wavelength light (597–663 nm) had no
significant effect on alarm flash behavior. These results indicate that
fireflies can respond to increased background illumination by produc
ing light signals of greater intensity. These results also suggest that,
at least for this species, long‐wavelength artificial light (>597 nm,
amber to red) is less disruptive than short‐wavelength (444–533 nm,
violet to green) ALAN (but see Buck, 1937b, Pacheco et al., 2016).
12 | CONCLUSIONS
Widespread nocturnal artificial illumination radically disrupts the
habitats of night‐active species. Nocturnal and crepuscular insects
are abundant and important components of these ecosystems. Thus,
the impact of ALAN on insect fitness and abundance can provide a
useful metric of overall ecosystem disturbance. The potential effects
of ALAN on insects can be categorized as temporal disorientation,
spatial disorientation, attraction, desensitization, and recognition.
The severity of impact will depend on the degree of overlap be‐
tween the spectral sensitivity of the insect in question and spectral
emission (and intensity) of the particular ALAN source (Gaston et al.,
2015). Recently, many urban areas have begun to phase out mono‐
chromatic long‐wavelength LPS lamps in favor of broad‐spectrum
white LED lighting. This spectral shift represents an ecological ex‐
periment on a global scale, with potentially devastating results.
One recommendation is of paramount importance: future stud
ies concerning the impact of ALAN on nocturnal organisms should
employ more objective, less anthropocentric methods for measuring
light. Light meter measurements in lux are adjusted for human lumi‐
nous sensitivity and can be greatly affected by differences in dis‐
tance from and angle to a light source. Insects often occupy discrete
microhabitats within larger light environments. Therefore, research
ers should strive to objectively measure how light varies within, as
well as among, nocturnal habitats.
Among night‐active insects, bioluminescent fireflies are con
spicuous and charismatic flagship species that can attract support
for conservation efforts aimed at minimizing excess nocturnal il‐
lumination in urban areas. As a result, recent studies conducted in
several geographic locations have focused on the impact of ALAN
on fireflies. Laboratory and field studies reviewed here demonstrate
that ambient light can inhibit the courtship flashing of several firefly
species. ALAN reduced courtship signaling by both sexes in a North
American firefly and reduced the ability of male European glow‐
worms to locate females. Laboratory studies testing the effects of
monochromatic light found that an Asian firefly is capable of increas‐
ing its flash intensit y in response to ALAN and that long wavelengths
(amber to red) were least disruptive.
This review also identifies several key gaps in our knowledge
concerning the impacts of AL AN on fireflies, and highlights several
important directions for future research. We currently know little
about temporal disorientation or desensitization by AL AN, and these
effects deserve further study. For example, comparative studies of
firefly phenology and nightly emergence times could reveal the de
gree to which ALAN delays and/or shortens the temporal scope of
courtship activity. Furthermore, despite evidence from several fire
fly species that ALAN interferes with mate location, it remains to
be seen whether this disruption has consequences for mating and
reproductive success. We also need information on how these be‐
havioral impacts translate into longer‐term effects on population
size and persistence.
Fireflies may be able to successfully cope with ALAN in various
ways, including increased dispersal or evolutionary adaptation. Can
fireflies move away from artificial lights toward darker habitat? We
currently know little about dispersal abilities in either the adult or
larval stages for any firefly species. While evidence suggests that
the colors of firefly bioluminescence are under selection to maximize
14 
|
   OWENS aNd LEWIS
signal contrast in different natural light environments (Hall et al.,
2016; Lall, Seliger, et al., 1980), whether ALAN selects for genetic
changes in firefly signals remains unknown. Comparison of urban
and rural populations may provide insight into adaptations that allow
fireflies to cope with high levels of ALAN.
It will also be important to investigate how different intensities
and colors of ALAN affect the courtship signaling of different firefly
species. While physiological studies (e.g., ERGs) have revealed varia‐
tion among species in their spectral sensitivity, the impact of differ‐
ent wavelengths on courtship success has yet to be determined. One
particularly important aspect of this issue is understanding how the
blue wavelengths emitted by commercial LED street lamps influence
courtship signals and mate location across different firefly species.
This research can help guide development of new lighting tech
nology that balances the need for public safety, energy efficiency,
and conservation, and inform policy recommendations for firefly‐
friendly ALAN sources that can be deployed on public, commercial,
and private lands in or near firefly habitat.
ACKNOWLEDGMENTS
We thank Tufts University and the Zoological Lighting Institute for
supporting this work.
CONFLICT OF INTEREST
None declared.
AUTHORS CONTRIBUTION
ACSO conceived and drafted the first version of this manuscript
and performed all data analyses. Both authors contributed signifi‐
cantly to revisions and have given final approval of the version to
be published.
DATA ACCESSIBILITY
Effect size calculations: Dryad https://doi.org/10.5061/dryad.jm8ps6b.
ORCID
Avalon C. S. Owens http://orcid.org/0000‐0001‐72122275
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How to cite this article: Owens ACS, Lewis SM. The impact
of artificial light at night on nocturnal insects: A review and
synthesis. Ecol Evol. 2018;00:1–22. https://doi.org/10.1002/
ece3.4557