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BAAE-51130; No.


Basic and Applied Ecology xxx (2017) xxx–xxx

Light pollution may create demographic traps for nocturnal

Ariel Firebaugha , Kyle J. Haynesb,∗
Department of Environmental Sciences, University of Virginia, Charlottesville, VA, 22904-4132, USA
Blandy Experimental Farm, University of Virginia, 400 Blandy Farm Lane, Boyce, VA, 22620, USA

Received 7 March 2018; received in revised form 6 July 2018; accepted 8 July 2018

Light pollution impacts both intra- and inter-specific interactions, such as interactions between mates and predator–prey
interactions. In mobile organisms attracted to artificial lights, the effect of light pollution on these interactions may be intensified.
If organisms are repelled by artificial lights, effects of light pollution on intra- and inter-specific interactions may be diminished
as organisms move away. However, organisms repelled by artificial lights would likely lose suitable habitat as light pollution
expands. Thus, we investigated how light pollution affects both net attraction or repulsion of organisms and effects on intra- and
inter-specific interactions. In manipulative field studies using fireflies, we found that Photuris versicolor and Photinus pyralis
fireflies were lured to artificial (LED) light at night and that both species were less likely to engage in courtship dialogues
(bioluminescent flashing) in light-polluted field plots. Light pollution also lowered the mating success of P. pyralis. P. versicolor
is known to prey upon P. pyralis by mimicking the flash patterns of P. pyralis, but we did not find an effect of light pollution
on Photuris–Photinus predator–prey interactions. Our study suggests, that for some nocturnal insects, light-polluted areas may
act as demographic traps, i.e., areas where immigration exceeds emigration and inhibition of courtship dialogues and mating
reduces reproduction. Examining multiple factors affecting population growth in concert is needed to understand and mitigate
impacts of light pollution on wildlife.
© 2018 Gesellschaft für Ökologie. Published by Elsevier GmbH. All rights reserved.

Keywords: Artificial light; Courtship; Population sink; Predator–prey

Introduction acoustic, tactile, and olfactory regimes (i.e., sensory pollu-

tion; Halfwerk & Slabbekoorn 2015). Because organisms use
Changes in human land use (e.g., urbanization and agricul- sensory information to interpret and interact with their sur-
tural intensification) affect large swaths of the globe (Venter roundings, sensory pollution is increasingly recognized as a
et al. 2016), and are considered major drivers of biodiversity conservation concern, especially in cases where the organ-
loss (Foley et al. 2005; Newbold et al. 2015). One component ism’s response decreases fitness (Delhey & Peters 2017).
of land-use change is alteration of sensory inputs: new photic, Light pollution — defined here as artificial illuminance —
alters visual information for crepuscular and nocturnal organ-
isms (Davies, Bennie, Inger, & Gaston 2013). Light pollution
∗ Corresponding
already covers over 22.5% of the globe’s terrestrial area
E-mail address: (K.J. Haynes).
(Falchi et al. 2016), and Cinzano, Falchi, and Elvidge (2001)
1439-1791/© 2018 Gesellschaft für Ökologie. Published by Elsevier GmbH. All rights reserved.

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showed that light pollution is increasing spatially at a rate that used non-visual cues to find prey. While some predator
of 6% per year. As light pollution spreads, understanding its species may exploit lights to their advantage, other predator
ecological consequences is increasingly critical. species may be negatively impacted.
Impacts of light pollution on intra-specific interactions Fireflies are ideal for studying effects of light pollution on
are largely unexplored (Longcore & Rich 2004), but one intra- and inter-specific interactions due to their unique light-
area where they do seem to be important is sexual behav- based communication system used for both courtship and
ior (Delhey & Peters 2017). For example, misleading light predation (many predaceous species lure in prey species by
cues could disrupt the release of reproductive hormones mimicking their flash patterns; Lloyd 1975). Fireflies are bee-
(Russ et al. 2015), pheromone signaling (van Geffen et al. tles in the family Lampyridae. There are about 2000 species
2015), or courtship behaviors prior to mating (Endler 1992; described worldwide, and they occur on almost every con-
Endler & Thery 1996; Heindl & Winkler 2003). Botha, Jones, tinent (Lloyd 2004). Many firefly species are nocturnal or
& Hopkins (2017) show light-pollution effects on sexual crepuscular and communicate using bioluminescent flashes.
behavior are complex; lifetime exposure to elevated-light Laboratory experiments have shown artificial light can stop
conditions in the lab increased the mating probability of male or delay the timing of firefly flashing (Buck 1937; Merritt,
crickets, decreased precopulatory behaviors of female crick- Rodgers, Amir, & Clarke 2012). Observational field studies
ets associated with mating efficiency, and had no effect on the have reported reduced nocturnal firefly flashing in light-
number or structure of acoustic courtship calls. Light pollu- polluted areas (Hagen & Viviani 2009; Viviani, Rocha, &
tion could also impact reproduction indirectly by affecting Hagen 2010; Picchi, Avolio, Azzani, Brombin, & Camerini
movement behaviors that determine the spatial aggregation 2013; Hagen, Santos, Schlindwein, & Viviani 2015). Addi-
of conspecific individuals. There is evidence that attraction tionally, manipulative experiments showed light pollution
to artificial lights is sex dependent in some moth species reduced firefly flashing activities in the field. Costin and
(Altermatt, Ebert, & Altermatt 2016; Degen et al. 2016), but Boulton (2016) found that firefly flashes per minute decreased
the effects on mating of attraction to artificial lights remain by around 50% on nights when artificial light was introduced.
unknown. Firebaugh and Haynes (2016) found that Photinus pyralis
Light pollution may also impact inter-specific interac- females were three times more likely to flash in response
tions (Longcore & Rich 2004). For example, attraction to to males during courtship in control plots where no artifi-
or repulsion from artificial lights may alter encounter rates cial light was added than in plots with added artificial light.
between predators and prey. Minnaar, Boyles, Minnaar, Sole, The effects of light pollution on predator–prey interactions
& McKechnie (2015) found moth consumption by bats between firefly species have not been evaluated. In addition,
increased near lights despite lower moth abundances in lit the effects of light pollution on firefly movement (attraction
areas. Rydell (1991) showed predators adjust the areas over or repulsion from artificial lights) is also poorly understood
which they forage in response to light-mediated changes (Firebaugh & Haynes 2016).
in prey availability. Conversely, prey behavior may change We conducted a field experiment to investigate whether
if predation risk in artificially lit areas is heightened or is Photuris versicolor (hereafter, the predator species) and P.
perceived to be heighted. Yorzinski et al. (2015) observed pyralis (hereafter, the prey species) fireflies displayed net
peahens increased anti-predator vigilance behaviors under movement towards (attraction) or away (repulsion) from arti-
artificial lights, and Rotics, Dayan, & Kronfeld-Schor (2011) ficial lights at night. We also conducted a series of field
found one mouse species spent less time performing con- experiments to examine effects of light pollution on intra-
spicuous activities under lights. Additionally, light from the specific (courtship) interactions of the predator species and
full moon is known to increase prey anti-predator or vigi- inter-specific (predator–prey) interactions between the two
lance behaviors (Beauchamp & McNeil 2003; Biebouw & firefly species. Previously, females of the prey species were
Blumstein 2003; Kotler, Brown, Mukherjee, Berger-tal, & shown to flash three times less frequently in the presence
Bouskila 2010). By giving advantage to a predator or its prey of artificial lights (Firebaugh & Haynes 2016), but effects
(or vice versa), light pollution could potentially destabilize on courtship in the predator species were not studied. To
predator–prey interactions. assess how light pollution impacts the predator species’ mat-
Few studies have examined effects of light pollution on ing behavior, we observed courtship dialogues in the field
predator–prey interactions in invertebrates. Bennie et al. with and without artificial lighting. To explore interactions
(2015) found no interactive effect of light pollution and preda- between light pollution, predator–prey interactions, and prey
tor presence on aphid abundance in experimental mesocosms. mating behaviors, we conducted a manipulative experiment
Underwood, Davies, & Queir (2017) showed dogwhelk mol- where the prey species was exposed to light pollution or
lusks acclimated to light pollution were more likely to engage natural-light conditions with and without the predator species
in conspicuous foraging activities in the presence of olfactory present. We predicted light pollution would decrease flash-
predator cues. Miller et al. (2017) showed light pollution ing by the predator species based on a previous finding of
decreased aphid consumption by one nocturnal ladybug reduced flashing activity in experimentally lit portions of a
species that used visual cues to find prey, but had no effect field (Firebaugh & Haynes 2016). We also predicted the pres-
on aphid consumption by another nocturnal ladybug species ence of the predator species would reduce flashing by the prey

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species as courtship flashing may make the prey species more lights is typically higher before midnight than between mid-
vulnerable to predators. night and dawn (Wakefield, Broyles, Stone, Jones, & Harris
2016). Effects of light pollution on the abundances of the
prey and predator species on the traps was analyzed using
Materials and methods a linear-mixed-effects (LME) model with light pollution as
the fixed effect and observation round (sunset, 1 h after sun-
Study system set, 2 h after sunset, or midnight) as both a fixed and random
effect (Bondell, Krishna, & Ghosh 2011). Observation round
Firefly flashes enable species identification during is included as both a fixed and a random effect because we
courtship. Male fireflies flash with a species-specific flash were interested in changes in firefly captures throughout the
pattern, and conspecific females respond by flashing back evening, but observation round cannot be randomly assigned.
(Lloyd 1971). Flashing begins at or soon after sunset when The abundances of the prey and predator species on the traps
ambient light levels are low (Lall, Seliger, Biggley, & Lloyd were log transformed to improve the normality of the LME
1980). In addition to their role in courtship, firefly flashes model residuals.
play a role in predator–prey interactions between predatory
fireflies in the genus Photuris and Photinus (prey) fireflies.
Photuris males and females mimic the flash patterns of Phot- Courtship of the predator species
inus prey to lure in Photinus prey (Lloyd 1981). Predation
of Photinus prey allows Photuris individuals to accumulate To assess how light pollution affects the predator species’
lucibufagin compounds, which make them distasteful to gen- mating behavior, we observed courtship dialogues between
eralist predators (Eisner, Goetz, Hill, Smedley, & Meinwald free-flying males and stationary females in the field with and
1997). without artificial lighting. We placed females (11 per light
This study was conducted during the summer of 2016 at pollution treatment) in 10 × 10 × 10 cm nylon-mesh contain-
Blandy Experimental Farm (BEF), a University of Virginia ers (1 female per container). The bottom of the container was
research station (39◦ 03 50.43 N,78◦ 03 47.20 W). The prey open so the female could sit on vegetation (warm-season
species (P. pyralis) and the predator species (P. versicolor) grasses), as they normally do during courtship activities
dominate the firefly community at BEF. The prey species is (Lloyd 2004). Females were given 10 min to acclimate to the
common throughout eastern North America (Lloyd 2004). cage before we began recording behavior. We observed each
At BEF, the prey species’ courtship displays usually occur female for 15 min, and recorded the number of flashing males
within 90 min of sunset at BEF, whereas the predator species within 1 m of the female, and the number of flashes from
is most active in courtship displays 1–3 h after sunset. The the caged female. Plots were paired with a distance of 20 m
predator species’ flash patterns are more rapid than those of between plots within a pair. The light-pollution treatment,
the prey species (Lloyd 1990), making it easy to differentiate consisting of 2 downward-pointing white LED floodlights
between the two species. (Ultra-Tow 9-32 Volt LED Floodlight) mounted at a height
of 3 m, was randomly assigned to one plot within each pair.
Mean illuminance at chest height directly under the light
Firefly attraction to artificial lights fixture at plots receiving the light-pollution treatment was
57.4 ± 0.89 lx, comparable to illuminance on a bright urban
We conducted an experiment to test whether fireflies are street (Gaston, Duffy, Gaston, Bennie, & Davies 2014). Mean
attracted to, or repulsed from, artificial lights at night. We illuminance at unlit plots was 0.03 ± 0.01 lx. Trials took place
counted firefly individuals on paired lit and unlit sheet traps from 9:30 to 11:30 PM over the course of 10 nights in June.
hung in lawn areas at BEF. Each sheet trap consisted of an For both sexes of the predator species, lack of variability in the
opaque white plastic sheet (177 w × 182 cm) hung between rate of flashing in the light-pollution treatment (see Section
two vertical objects (e.g., trees, fence posts, etc.) at a height “Results”) prevented statistical analysis.
of 1.5 m so that the bottom edge of the shower curtain just
touched the ground. Traps were lit with an LED flood-
light (Ultra-Tow 9-32 Volt LED Floodlight, Northern Tool Predator–prey interactions and prey mating
& Equipment Company, Inc., Burnsville, Minnesota, USA) behaviors
positioned 0.3 m above the ground and 1.5 m away from
the sheet, with the light pointed at the center of the sheet. We conducted a manipulative field experiment to exam-
The paired lit and unlit traps were located 10 m apart. We ine effects of light pollution on predation and to assess
counted fireflies on the traps for 15 pairs of traps over the whether the prey species altered its rate of courtship flashing
course of 14 nights at sunset, one hour after sunset, two hours in response to presence of the predator species under dark
after sunset, and midnight. We stopped the trials at midnight or artificially lit conditions. The experiment was conducted
because firefly flashing activities at BEF are highest between in 3 × 3 × 3 m floorless, mesh-sided tents (Fig. 1; Caravan
sunset and midnight, and because insect capture using LED Canopy, Caravan Global US, La Mirada CA) placed over

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rounds (one hour after sunset and two hours after sunset) were
analyzed using a generalized-linear-mixed-effects (GLME)
model, specifying a binomial distribution (female did or did
not flash) and the logit link function. Light pollution was
a fixed effect and observation round and trial were treated
as random effects. Predator-species flashes at sunset were
excluded from the model as there was little to no flashing
at this time. The number of prey species flashes per minute
was log transformed to improve the normality of the LME
model residuals. Effects of light pollution and predator pres-
ence on whether or not the females of the prey species mated
were tested using a GLME model, specifying a binomial dis-
tribution and the logit link function, with light pollution and
predator presence as fixed effects, and date as a random effect.
Fig. 1. Experimental mesocosms used to assess predator–prey inter- All statistical analyses were conducted in R (R Core Team
actions and prey mating behaviors in the presence and absence of 2015). The LME and GLME models were performed using
artificial illumination. the R package ‘lme4’ (Bates, Mächler, Bolker, & Walker
mowed vegetation in an old field meadow. We stocked all
mesocosms with 3 males and 1 female of the prey species.
Light-pollution (present or unlit control) and predator- Results
density treatments (1 female predator or no predators) were
applied to the mesocosms according to a factorial design Firefly attraction to artificial lights
with 1 replicate of each treatment completed during each
of 14 trials. Light pollution was added by mounting 1 LED On average, 189% more prey-species individuals were
light (Ultra-Tow 9-32 Volt LED Floodlight) in the center of found on the lit sheet traps than the unlit sheet traps by
a mesocosm at a height of 2.5 m. Light intensity at chest the end of each collection night (i.e., midnight) (Fig. 2A;
height in mesocosms in which artificial light was added mea- t1,14 = −9.76, p ≤ 0.001). The lit traps captured, on aver-
sured 174.53 ± 1.08 lx. Light intensity in unlit cages was age, 0.21 ± 0.52 individuals of the predator species, while
0.03 ± 0.02 lx. Each trial took place over approximately 7 h none were found on the unlit traps (Fig. 2B; t4,113 = −3.27,
(5 PM–12 AM). Densities (no. per m−2 ) of the predator and p = 0.001). For both species, the number found on the
prey species in the experiment were approximately one third traps varied significantly among rounds for both the prey
and one fifth, respectively, the peak densities observed at BEF species (t1,14 = −4.23, p = 0.048) and the predator species
in 2015 (Firebaugh & Haynes 2016). (t1,14 = 2.15, p = 0.034), with the number of individuals
Fireflies were marked with fluorescent powder to facilitate increasing over the course of the evening (Fig. 2).
recapture and were put in the mesocosms at 5:00 PM before
each trail. For each of three observation rounds (sunset, one
Courtship in the predator species
hour after sunset, and two hours after sunset), we counted
the number of flashes during three 1-min periods by the
Females tethered under artificial lights never flashed,
prey-species (3 males and 1 female) and the predator-species
whereas all 11 females tethered in unlit plots flashed at least
female in each tent. Each trial lasted 1 night. After the final
once (Fig. 3A). On average, there were 7.27 ± 9.08 flashes
observation round, we recovered all fireflies and recorded the
from males within 1 m of each female in unlit plots, but males
number living and dead. A UV flashlight was used to recover
never flashed near females in lit plots (Fig. 3B).
the fireflies (the fluorescent powder glowed under UV light).
We assessed whether females of the prey species had mated
by checking their genitalia for the presence of the fluorescent Predator–prey interactions and prey mating
color used to mark males of the prey species. The presence behaviors
of the powder color used to mark the males on a female’s
genitalia suggested that mating was likely. In the predator–prey experiment, light pollution decreased
Effects of light pollution and predator presence on the flashes by males of the prey species by 25% (Fig. 4A;
prey species’ flashing activity were analyzed using a LME t5,485 = 2.93, p = 0.004). There was no significant effect of
model, with light pollution, predator presence, and light pol- the presence of a predator (t5,485 = 0.410, p = 0.681), and no
lution × predator presence as fixed effects, and observation significant interactive effect of predator presence and light
round (sunset, 1 h after sunset, and 2 h after sunset) and trial pollution (t5,485 = −1.57, p = 0.117) on the number of flashes
as random effects. Effects of light pollution on the preda- by males of the prey species. Light pollution significantly
tor species’ flashing activity during the final two observation decreased the number of predator species flashes (Fig. 4B;

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Fig. 2. Abundances of the prey species (Photinus pyralis) (A) and the predator species (Photuris versicolor) (B) observed on lit and unlit
sheet traps at sunset, 1 h after sunset, 2 h after sunset, and midnight (n = 15 pairs). Error bars represent ±1 standard error of the mean.

Fig. 3. Photuris versicolor flashes in (A) females and (B) males observed with and without artificial illumination (n = 11 pairs).

z = 5.56, p ≤ 0.001). None of the prey-species individuals this study, we found that two firefly species were attracted
were eaten during this experiment; all marked individuals to ‘white’ LED lighting (Fig. 2), which is rapidly becom-
were recovered alive. The proportion of females of the prey ing the dominant outdoor lighting technology (Baumgartner
species that mated was twice as high in lit than unlit meso- et al. 2012; Stanley et al. 2015). Our results also show
cosms (Fig. 5), however, this difference was not significant that ‘white’ LED lighting at night dramatically reduced the
(z = 1.58, p = 0.114). The presence or absence of a predator courtship behavior (flashing) and likely decreased mating
had no significant effect on the portion of females of the prey of the prey species (as evidenced by observations of males
species that mated (z = 0.453, p = 0.651). and females in coitus or transfer of fluorescent powder from
males to females), consistent with the findings of Firebaugh
and Haynes (2016), while also strongly reducing flashing by
Discussion the predator species. We were not able to assess effects of
light pollution on the predator species’ mating success, but
Gaston and Bennie (2014) proposed that light-polluted the lack of flashing activity under light-polluted conditions
areas may act as population sinks (immigration > emigration suggests mating would also be reduced in this species. For
and mortality > births). However, few studies have assessed fireflies, the combined effects of nighttime LED lighting –
effects of light pollution on multiple demographic processes attraction, strongly reduced flashing, and likely decreased
in any single species (Gaston & Bennie 2014, but see Mclay, mating success – suggest areas where outdoor LED lighting
Green, & Jones 2017; McMahon, Rohr, & Bernal 2017). In is installed may become demographic traps. The rapid accu-

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Fig. 4. Number of prey species (Photinus pyralis) flashes (A) and predator species (Photuris versicolor) flashes (B) in experimental mesocosms
with and without artificial illumination (n = 56). Error bars represent ±1 standard error of the mean.

higher if Photuris consumed Photinus to meet nutritional

Light pollution has the potential to intensify (Rydell
2006; Miller et al. 2017) or weaken (McMahon et al. 2017)
intra- and inter-specific interactions through its effects on
movement and behavior. Though our results suggest the
predator–prey interaction between the two species we studied
is weak (the predator species never consumed the prey species
in our mesocosm experiment), light pollution may further
weaken this interaction because light pollution strongly
Fig. 5. Proportion of prey species (Photinus pyralis) females that inhibited flashing behaviors (Fig. 4B), which the predator
mated in mesocosms that were or were not artificially lit, in the species use to lure the prey species. Though we found that
presence or absence of the predator species (Photuris versicolor;
both species aggregate near artificial lights, which could
n = 56).
intensify interactions by increasing encounters between indi-
viduals, it seems likely light pollution weakens both the
intra-specific interactions (mating) and inter-specific inter-
actions (predation) of our study organisms given that these
mulation of fireflies on sheet traps lit by LED lighting but not interactions are mediated by flashing dialogues.
on unlit control sheet traps indicates this LED lighting caused
immigration to exceed emigration. An important question for
future research is whether the attraction of nocturnal insects
to light-polluted areas, coupled with decreases in courtship Acknowledgements
and mating, leads to the creation of population sinks.
We found no evidence for lethal or sub-lethal effects of We thank two anonymous reviewers for providing helpful
predator presence on the prey species. The prey species was comments on a previous draft of this manuscript. This work
not eaten during the mesocosm experiment, and the prey was supported by funding from Blandy Experimental Farm
species’ flashing activity was the same in the presence or and the University of Virginia Department of Environmental
absence of the predator species (Fig. 4A). It is possible the Sciences. Assistance with the fieldwork was provided by B.
mesocosms altered activity levels or flight behavior of the Cook, M. Hey, and O. Lewis.
predator species, but it appeared to exhibit typical flying
and flashing behaviors in the mesocosms. The lack of preda-
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