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REVIEWS REVIEWS REVIEWS The nature, extent, and ecological 347 implications of marine

Thomas W Davies*, James P Duffy, Jon Bennie, and Kevin J Gaston

Despite centuries of use, artificial light at night has only recently been recognized as a cause for environmental concern. Its global extent and ongoing encroachment into naturally lit has sparked scientific interest into the many ways in which it may negatively affect human health, societal attitudes, scientific endeavors, and biological processes. Yet, perhaps because sources of artificial light are largely land based, the potential for artifi- cial light pollution to interfere with the biology of the ocean has not been explored in any detail. There is little information on how light pollution affects those species, behaviors, and interactions that are informed by the intensity, spectra, and periodicity of natural nighttime light in marine ecosystems. Here, we provide an overview of the extent of marine light pollution, discuss how it changes the physical environment, and explore its poten- tial role in shaping marine ecosystems.

Front Ecol Environ 2014; 12(6): 347–355, doi:10.1890/130281

rtificial light at night is globally widespread (Cinzano systems, document its current spatial extent, and high- A et al. 2001), rapidly expanding in spatial extent light a wide array of known and potential ecological (Hölker et al. 2010), and shifting in its spectral character- impacts. istics (Davies et al. 2013a). Increasing concern over these trends has recently fuelled a surge in research toward n Nature and extent understanding the ecological impacts of artificial light pollution (eg Davies et al. 2012, 2013a; Stone et al. 2012; Satellite images of the Earth’s nighttime lights are a testi- Dominoni et al. 2013). While the impacts of light pollu- mony to humankind’s widespread colonization and subse- tion are increasingly well documented in terrestrial quent influence on the planet’s ecosystems (Figure 1). As ecosystems, marine have received comparatively an initial estimate, in 2010, 354 760 km (22.2%) of the little attention (Depledge et al. 2010). Indeed, in a recent world’s coastlines (excluding Antarctica) were exposed to survey in which 592 coastal scientists were asked to prior- nightly artificial light pollution (see Table 1 for details). itize global coastal research questions, artificial light was While it might be expected that the coastlines of the not included (Rudd and Lawton 2013). Yet, worldwide, developed world are most affected, this is not so: Asia and more than a billion people (~23% of the global popula- Africa, for example, have the second and third highest tion) live within 100 km of a coastline (Small and percentage of coastline influenced by light pollution, Nicholls 2003), and many coastal marine ecosystems are respectively, with Europe having the highest (Figure 1; exposed to artificial light at night. Here, we draw atten- Table 1). The amount of artificial light on land is contin- tion to artificial light as an environmental issue in marine uing to increase at a rate of 6% per year globally (Hölker et al. 2010). Given the rapid growth of many developing world economies, future increases are expected to be In a nutshell: greater in these regions as compared with the developed • Artificial light pollution is globally widespread in marine world over the coming decades, with unknown conse- environments, altering the natural colors, cycles, and intensi- quences for some of the planet’s most biodiverse marine ties of nighttime light, each of which guide a variety of bio- ecosystems (Aubrecht et al. 2008). logical processes • Known and potential impacts include those on navigation, Sources of artificial light in the marine environment reproduction and , predator–prey interactions, vary, with shipping and light fisheries contributing as and communication in a myriad of marine species and ecosys- temporary sources in nearshore and offshore waters tems, including some of the world’s most biologically diverse (Figure 2, a and f). Offshore oil platforms and land-based and functionally important developments such as towns, cities, and their harbors pro- • Research into these impacts is needed to inform conservation strategies and policy decisions relevant to the holistic man- vide more permanent sources that can increase nighttime agement of marine ecosystems experiencing an increasingly light intensities across large geographical areas such as diverse array of anthropogenic stressors estuaries, bays, and continental shelf seas (Figure 2, b and e). Scattering of upwardly emitted or reflected artificial light in the atmosphere and reflection by clouds (artificial Environment and Sustainability Institute, University of Exeter, skyglow) further extends the spatial influence of land- Cornwall, UK *([email protected]) based light sources (Kyba et al. 2011) into offshore waters,

© The Ecological Society of America www.frontiersinecology.org Marine light pollution TW Davies et al.

348 lightscapes at night (Figure 2c), which were previously lit only by the comparatively constant broad spec- trum of moonlight (Figure 2d). This is particularly pertinent to depth- dependent biological processes that can be guided by the spectrum of light as it changes through the water column. Blue artificial light will pen- etrate deeper in the open ocean due to the faster attenuation of red light, increasing its potential scale of influ- ence. Higher concentrations of bio- logical and nonbiological particulates alter this attenuation so that, for instance, green artificial lights would penetrate deepest in temperate regions where phytoplankton absorb other wavelengths. Given the increasing proportion of coastlines that are artificially illumi- nated and the variety of ways this alters the natural light regimes with which marine organisms have Figure 1. The global extent of marine light pollution. Images are derived from the evolved, it is essential that scientists Defense Meteorological Satellite Program Operational Line Scan nighttime lights image gain a better understanding of the for 2010 (Behrmann equal-area projected; pixels aggregated to 8-km resolution). Graph known and potential ecological con- inserts in (a) represent the distribution of artificial light intensity across continental sequences of artificial light in marine coastlines, calculated from raw data at the original ~800-m resolution. The x axes ecosystems. represent artificial light intensity (0–63 Digital Number [DN]), where the brightest pixels are assigned the highest values); the y axes represent log km of coastline affected by that n Ecological implications intensity of light. Coastlines (highlighted) were defined as anywhere up to 10-km inland. The vast majority of species have and masks natural rhythms of lunar light intensity evolved under natural and predictable regimes of moon- (Davies et al. 2013b) that provide information for regulat- light, sunlight, and starlight. These regimes define ing a number of biological processes in the ocean (Naylor species’ activity times (eg nocturnal, crepuscular, diur- 1999). Ongoing shifts in prevailing technologies are nal), offer a useful navigational aid, help to regulate and resulting in an increasing use of “white” or broader spec- coordinate maturation and reproductive events, and pro- trum lights (Gaston et al. 2012), and a greater variety of vide a relatively constant irradiance spectrum that can spectrally distinct lighting types is creating complex spa- regulate physiology and inform visually guided behaviors tial patterns of color and brightness across marine such as and communication (Gaston et al. 2013). Many of these processes are affected by artificial Table 1. The spatial extent of coastal light pollution nighttime light in terrestrial ecosystems: for example, in Kilometers of Percent of , sexual maturation is advanced (Dominoni et al. Region coastline affected coastline affected 2013), effort is intensified (Titulaer et al. 2012), Europe 115 383 54.3 and the timing of dawn song is extended into the night Asia (excluding Russia) 113 166 34.2 (Nordt and Klenke 2013). Some species are attracted to Africa 18 589 22.1 artificially lit areas where they may experience increased South America 24 197 15.5 predation (Rydell 1992), while others avoid artificially lit North America 64 356 11.8 areas (Stone et al. 2012), and so are displaced from habi- Oceania 11 692 7.9 tats that would otherwise be suitably dark in the absence Russia 7377 6.1 of artificial lighting. Many of the same types of light cues Total 354 760 22.2 are just as intrinsic to the of marine species as Notes: Values are derived from a Behrmann equal-area projected Defense they are for terrestrial species. Artificial nighttime light is Meteorological Satellite Program Operational Line Scan nighttime lights image from 2010 (see Figure 1). Light-polluted areas of coastline were defined as those therefore likely to affect a diverse array of ecological where the pixel intensity was greater than five on an uncalibrated scale between processes in the marine environment (Figure 3). Here, we zero and 63. Antarctica was omitted when calculating the total % of coastline. highlight some of the major impacts, providing examples

www.frontiersinecology.org © The Ecological Society of America TW Davies et al. Marine light pollution from taxa known to be affected 349 as well as functionally impor- tant taxa that are likely to be affected.

Orientation

Perhaps the best-known impact of artificial light is the disorien- tation experienced by species (a) (b) that use natural light cues to navigate, most notably birds and sea turtles (Tuxbury and 2005; Merkel 2010) in marine systems. strikes, involving a variety of species, on artificially lit vessels at sea are common at night (Merkel 2010), whereas coastal lighting disorientates turtle hatchlings and prevents or delays them (c) (d) from locating the sea (Tuxbury and Salmon 2005), ultimately reducing the number of turtle nesting sites in artificially lit locations (Mazor et al. 2013). In the intertidal environment, moonlight provides a compass for navigation by the inverte- brate sandhopper Talitrus saltator (Ugolini et al. 2005), raising the question of whether artifi- (e) (f) cial lighting is disrupting lunar- Figure 2. Sources of artificial light in marine ecosystems. (a) Artificial lights from ships that orientated tidal migrations. anchor in Falmouth Bay, UK. (b) The lights of Plymouth, UK, illuminating the Tamar This behavior is also seen in Estuary and its creeks and tributaries. (c) Decorative, harbor, and municipal lighting spilling terrestrial species, such as dung onto the Tamar, viewed from Cremyll, UK. (d) Moonlight cast onto the shores of Cremyll in beetles (Dacke et al. 2003), and the Tamar Estuary, UK. (e) Oil rig lighting across the North Sea captured by satellite. (f) is likely to be widespread across Light fisheries target squid spawning grounds in the Tsushima Strait. Satellite images courtesy of both terrestrial and marine NASA/NOAA Suomi NPP satellite 2012 (www.earthobservatory.nasa.gov/NaturalHazards/ arthropods. Behavioral respon- view.php?id=79765). ses to artificial light have also been demonstrated in various fish species (Marchesan et likely to influence the behavior and survivorship of many al. 2005). For instance, sea cages that are artificially lit nontarget as well as target species over large spatial scales. affect depth selection in Atlantic salmon (Salmo salar; Light intensity and spectra are especially useful cues by Oppedal et al. 2011), while coastal lighting around estu- which organisms regulate their depth in the pelagic envi- aries can aggregate fish in artificially lit habitats (Becker ronment where landmarks are absent. While some et al. 2012). Indeed, the attraction of the larval stages of species have evolved the ability to navigate horizontally many fish species to artificial lights has led to the devel- via detection of the Earth’s geomagnetic field, the visible opment of light-based trapping methods that are analo- portion of the electromagnetic spectrum is one of the gous to those used for trapping nocturnal most important means of vertical navigation. Con- (Doherty 1987). Artificial lights are commonly used by sequently, light intensity informs the vertical movement the fishing industry to attract and catch several squid of zooplankton species that migrate to surface waters at species. Those lights are so powerful that nighttime satel- night to graze while avoiding predation (Cohen and lite images can be used to quantify fishing pressure, Forward 2009). Given that artificial skyglow is more spawning grounds, and migration routes (Figure 2f; widespread than direct artificial light (Cinzano et al. Kiyofuji and Saitoh 2004). The geographically wide- 2001) and frequently occurs at intensities greater than spread use of powerful artificial lights in these fisheries is (and thus directly interferes with) natural lunar sky

© The Ecological Society of America www.frontiersinecology.org Marine light pollution TW Davies et al.

350 light regimes to regulate rates of repro- ductive maturation and synchronize (b) broadcast spawning events (the release (a) of male and female gametes into the surrounding environment). In the (b) aquaculture industry, these photoperi- (c) odic responses are manipulated with artificial lighting to control sexual mat- uration (Oppedal et al. 2011). A classic example of apparent lunar periodicity (h) (h) in broadcast spawning is the palolo worm (Eunice viridis), which releases (f) gametes over a few days of the year dur- (d) ing the third-quarter moon in October (d) (Naylor 1999). Other marine poly- (g) chaetes apparently also synchronize (f) reproductive events to lunar cycles (Bentley et al. 1999), as do some corals (e) (Harrison 2011) and echinoderms (Lessios 1991). The degree to which lunar light intensity alone regulates such events, as compared to the role of Figure 3. Known and potential impacts of artificial light pollution on marine variables such as day length, tempera- ecosystems. (a) Suppression of zooplankton by artificial ture, and tidal conditions, is unknown skyglow. (b) Bird strikes on lit ships at night. (c) Extension of visual foraging behavior for many taxa. However, it seems likely in coastal wading birds into the night. (d) Disruption of settlement site selection in that in numerous cases a combination sessile invertebrate larvae. (e) Aggregation of fish under pier lights leads to intensified of these cues is used to regulate gamete predation. (f) De-synchronization of broadcast spawning from lunar phase (corals development, with lunar light intensity releasing gametes). (g) Displacement of nesting sea turtles from artificially lit nesting providing the final trigger for spawning areas. (h) Disorientation of seaward migration in hatchlings by street lights. (Harrison 2011). In addition, it has been postulated that the spectral con- brightness (Davies et al. 2013b), it seems likely that tem- trasts between moonlight and daylight vary more pre- poral patterns of zooplankton migration will be affected dictably than intensity alone (Sweeney et al. 2011), in artificially lit waters. Indeed, artificial light reflected implying that changes in artificial light spectra, and con- back to Earth from urban areas (a primary source of artifi- sequently the colors of skyglow, may also affect broadcast cial skyglow) limits the vertical migration of Daphnia spp spawning synchronicity. The advantage of synchronizing in freshwater lakes (Moore et al. 2000). spawning events is unclear, although it is probably linked The nightly vertical movement of zooplankton toward to maximizing reproductive contact between conspecifics the ocean surface is arguably the largest daily migration of (Harrison 2011). Lunar light intensity and day length on the planet (Hays 2003). The consumption of offer two cues that vary predictably and independently of near-surface phytoplankton by migrating zooplankton other environmental variables that are influenced by during the night and the subsequent defecation of fecal more stochastic Earth-bound processes. This includes pellets at depth during the day (Hays 2003; Cohen and tidal height and current velocity, which – in contrast to Forward 2009) constitutes a major pathway in the carbon the single detectable peak of lunar brightness every 29.5 cycle, and the adaptive behavior of predators to the verti- days – have multiple peaks within this time frame that are cal movements of their prey results in the daily migration more spatially variable in magnitude and timing. The of entire food webs (Hays 2003). Artificial light pollution “lunar clock” can, however, be masked by artificial sky- has the potential to disrupt this vertical migration pat- glow (Davies et al. 2013b), which may interfere with the tern, and with it the and cycling of carbon synchronization of spawning events, resulting in and nutrients in marine ecosystems. decreased cross fertilization and ultimately a decline in recruitment among broadcast-spawning species. Reproduction and recruitment It is well known that the intensity and spectral charac- teristics of light inform the orientation and settlement of The movements of the celestial bodies create predictable the larvae of a broad array of sessile invertebrates regimes in the intensity and duration of light that organ- (Thorson 1964), including corals, arthropods, poly- isms use as a clock to synchronize activity (Naylor 1999). chaetes, echinoderms, tunicates, and bryozoans. Altering In the marine environment, many species use natural the intensity or spectral signature of light could interfere

www.frontiersinecology.org © The Ecological Society of America TW Davies et al. Marine light pollution with both the pelagic and pre-settlement stages of marine 351 invertebrate life cycles, with major consequences for ses- sile invertebrate ecosystems. For example, as in many shallow seabed communities, coral reef assemblages are structured vertically with increasing depth (Vermeij and Bak 2002). Coral larvae identify their optimum settle- ment zones using the intensity and spectral characteris- tics of light (Mundy and Babcock 1998), which vary with depth due to the differential absorption of different wave-

lengths of light in water. In a similar fashion, light inten- Semenov A sities allow the larvae of sessile invertebrates in shallow Figure 4. The bioluminescent comb jelly, Beroe cucumis, subtidal environments to avoid settling in areas that are occurs at depths that can be penetrated by artificial light. sufficiently lit to facilitate the growth of competitive algal species (Glasby 1999), giving rise to horizontal structur- (Rydell 1992). In the marine environment, direct artifi- ing of ecological communities. Artificial lights alter the cial light aggregates both small prey fish and larger preda- spectrum and intensity of light that larvae experience, tory species, increasing predation pressure in a similar resulting in suboptimal selection of sites to settle and fashion (Becker et al. 2012). In another example, turning metamorphose into adult form. For instance, barnacle off the lighting on bridges spanning the Puntledge River larvae can choose settlement zones under white light in British Columbia, Canada, reduced the intensity of pre- intensities of 1 × 10–5 lumen m–2 (lux) (Crisp and Ritz dation by harbor seals (Phoca vitulina) on migrating juve- 1973), which is four orders of magnitude less than that nile Pacific salmon (Oncorhynchus spp; Yurk and Trites emitted by urban skyglow (Gaston et al. 2013). In sessile 2000). Similarly, artificially lit intertidal mud flats allow invertebrates, settlement site selection ultimately decides coastal waders to employ more profitable visually guided survival and reproductive success; hence changing nat- foraging behaviors later into the night (Dwyer et al. 2013). ural light regimes could plausibly affect the composition Where the distribution of artificial light is patchy, this can and functioning of these communities, the members of result in for optimal foraging zones based on many of which (eg reefs) are engineers. artificial light intensity (Dwyer et al. 2013) and intensified predation within more brightly lit patches. Predation Many marine taxa emit light generated from biochemi- cal reactions (; Figure 4) to avoid preda- A variety of cryptic anti-predator defenses have evolved tion or to lure prey. Although popular perceptions of biolu- in the marine environment, of which and minescence are often associated with the deep sea, it is nocturnality are two of the most widespread. The ability evolutionarily and geographically more widespread to locate prey or avoid a predator depends largely on the throughout the oceans, occurring in fish, tunicates, echin- predator’s ability to recognize its quarry against complex oderms, arthropods, annelids, ribbon worms (Nemertea), backgrounds of shape, color, and pattern, and the prey’s cnidarians, mollusks, dinoflagellates, bacteria, and ability to disguise itself to avoid detection (Troscianko et chaetognaths in both oceanic and nearshore environments al. 2009). Many species have evolved color patterns to (Haddock et al. 2010). Anti-predator strategies include blend in with their environment and nocturnal behaviors using light displays to confuse predators, sacrificing light- to avoid predator contact, while polymorphic color vari- emitting body parts to distract predators, attracting the eties allow some species populations to rapidly adapt to predator’s own predators, and mimicking ambient light different environments and to predators with contrasting from above to avoid silhouette detection by predators from methods of prey search behavior. The increased intensity below (Haddock et al. 2010). The effects of artificial light- of artificial light as compared to moonlight, and the ing on these types of interactions are thus far largely unex- broadening of artificial light spectra, provides greater plored. The narrow spectrum of 590-nanometer (nm) visi- opportunities for predatory species to recognize their prey, ble light emitted by widely used 20th-century low-pressure potentially allowing diurnal and crepuscular foraging sodium lighting technologies and the short-wave spectral behaviors to encroach further into the night, displacing peak of ~470 nm emitted by most bioluminescent marine nocturnal prey or predator species from habitats that were taxa (Haddock et al. 2010) suggest that in the past, in previously exposed only to natural regimes of darkness. nearshore environments at least, land-based municipal Prey species may experience intensified predation pres- lighting may have had limited impacts on interactions gov- sure, resulting in localized population declines or shifts in erned by bioluminescent communication. However, the the frequency of particular polymorphs toward types that increasing use of broader spectrum lighting technologies are less conspicuous under artificial night lighting. (eg light-emitting diodes [LEDs]) over the coming decades Several studies have shown that the attraction of noc- will result in an increasing amount of land-based artificial turnal Lepidoptera to street lights at night results in aggre- light illuminating the 470-nm communication channel gations upon which opportunistic species often prey (Davies et al. 2013a) in inshore waters.

© The Ecological Society of America www.frontiersinecology.org Marine light pollution TW Davies et al.

352 Communication Quantifying threat extent

Complex eyes have evolved independently in several Quantifying the extent of the threat posed by light pollu- marine lineages, including fish, cephalopods, and arthro- tion to the ecology of marine systems requires research on pods, and just as in many terrestrial taxa, the spectra, pat- the number and phylogenetic breadth of species affected, terns, and angles of light reflected from individuals are use- the spatial influence of artificial light in marine ecosys- ful elements in inter- and intraspecies communication. As tems, and the range of biological complexity across which with butterflies and birds, the boldness of colorful markings impacts occur. communicates competitive to potential mates in In order to guide this process it is useful to know which many marine arthropods (Detto 2007) and fish (Siebeck et species are likely to be vulnerable and whether they are al. 2010). Cephalopods use adaptive displays of color and currently or are likely to be exposed to artificial light pol- pattern, created by a combination of reflecting and color- lution. Ecologists can identify vulnerable species both by changing cells to communicate (Mäthger et al. 2009); the looking for marine analogs of biological responses in ter- iridescent markings on squid, for example, vary in appear- restrial ecosystems and by extrapolating from studies that ance depending on the position of the onlooker. This document the role of natural light in marine ecosystems. could facilitate communication of shoaling behavior Screening for responses across many taxa can assist in the between conspecifics, a type of visual communication that development of comprehensive lists of vulnerable species may also be exploited by fish for the same purpose or behaviors, but such an approach can also be costly and (Mäthger et al. 2009). time-consuming. Targeting research toward species and The current introduction of artificial lights that emit ecosystems that have obvious ramifications for human broader spectra means that visually guided behaviors well-being (eg biomass production, coastal protection, such as mate selection are more likely to be influenced ecosystem stability, biogeochemical processes) will pro- by light pollution, since many of the physical features vide clear economic incentives for developing a thorough that are used to communicate fitness will be more recog- understanding of the problem, while remote-sensing tools nizable under whiter lighting (Davies et al. 2013a). This can be used to identify regions already exposed to and includes the use of lights that emit (UV), a likely to become exposed to light pollution in the future communication channel that some species exploit as priority research areas. almost exclusively to avoid predation or to distinguish Global satellite images of nighttime lighting have proven between closely related species (Siebeck et al. 2010). useful for identifying areas of coral reefs that are exposed to The Ambon damselfish (Pomacentrus amboinensis), for light pollution (Aubrecht et al. 2008), and this approach instance, uses UV light reflected in species-specific pat- could be extended to map artificial light in marine pro- terns to discriminate between conspecifics and other tected areas, as well as other regions containing species species of damselfish that appear identical at visible known to be sensitive to artificial lighting. However, satel- wavelengths (Siebeck et al. 2010). lite images are limited to measuring light emitted upward, Bioluminescence signals are also used in sexual communi- under clear-sky conditions while organisms are exposed to cation by marine species in a way that is analogous to fire- direct and scattered lighting and light reflected back from flies (Haddock et al. 2010). Examples include ostracods that the atmosphere, the amount of which can vary with local attract mates using bioluminescent signals and ponyfishes meteorological conditions in space and time. A large-scale (Leiognathidae) that use dimorphic bioluminescent dis- spatial and temporal analysis of light pollution trends and plays to differentiate between sexes (Haddock et al. 2010). long-term biological effects therefore requires a suitable The potential for artificial lights to disrupt mating behav- array of ground-based sensors with which to augment and iors that rely on bioluminescent cues has been raised with groundtruth satellite data so that accurate predictive mod- (Longcore and Rich 2004); artificial light could also els can be developed. Sky Quality Meters (SQMs; possibly disrupt such interactions in marine taxa. Unihedron, Grimsby, Canada) have proven useful for quan- tifying nighttime sky brightness at ground level in terrestrial n Moving forward systems (Kyba et al. 2011; Davies et al. 2013b); however, only a few SQMs have been deployed in coastal areas and We have drawn attention to artificial light as a threat to none so far in the open ocean. Offshore infrastructure marine and highlighted some of the diverse including oil platforms, “ships of opportunity” (ships willing array of species, behaviors, interactions, and ecosystems to participate in long-term monitoring programs), and the that are likely to be affected. The challenge now is to Global Ocean Observing System (sea surface buoys) all quantify the extent of the threat posed in regions where hold potential for establishing such a network of devices, such species and ecosystems exist, to develop a sound providing the logistical challenges associated with offshore knowledge base from which to design protective mea- deployment can be overcome. sures, and to identify how such measures can best be Measuring and monitoring marine light pollution at implemented. In this section, we highlight some of the large spatial scales offers the possibility of identifying key components of such work. threatened regions, quantifying trends, and formulating

www.frontiersinecology.org © The Ecological Society of America TW Davies et al. Marine light pollution predictive models of where future impacts are most likely on associated species, trophic structure, and ecosystem 353 to occur. Understanding where sensitive species and functioning, thereby compromising the delivery of ecosys- ecosystems are exposed to artificial light also requires that tem services. Various long-term ecological field manipula- mapping and modeling take account of vertical as well as tions of light pollution have recently been established in horizontal variability in artificial light, so that factors terrestrial systems. By artificially illuminating previously such as turbidity, which determines how spectrum and dark habitats at night, these experiments are elucidating depth interact to affect intensity, can be quantified. the impact of light pollution at the and ecosys- tem levels. The intertidal and shallow sublittoral marine Developing a sound knowledge base environments offer several tractable systems where similar experiments could be performed. As with many emerging ecological issues, research into marine light pollution will initially focus on documenting Reducing environmental impacts the range of species, behaviors, and places that are affected. It is also imperative that investigators explore the Initiatives to preserve naturally lit landscapes exist context of these effects to identify ecologically sound mit- through national and international programs that recog- igation measures that could provide alternatives to the nize the value of dark skies (eg www.darkskyparks.org). complete elimination of artificial lighting (as this will not The International Dark-Sky Association’s Dark Sky Parks always be possible). For example, many biological program, The Starlight Foundation’s Starlight Reserves, responses – including those associated with communica- and the UK’s Science and Technology Facilities Council’s tion, larval recruitment, or phototactic reactions in Dark Sky Discovery Sites are all examples of initiatives mobile fauna – are likely to be spectrally dependent. that award dark sky status to areas where light pollution is Narrow-band optical filters and LEDs offer the potential minimized. By participating in such programs, local gov- for controlling the intensity and spectrum of artificial erning authorities are incentivized to adopt measures that light independently, enabling investigators to identify preserve natural darkness and the cultural, ecological, aes- those regions of the light spectrum that have minimal bio- thetic, and scientific benefits this provides. Although logical impacts. Experiments of this nature can be used as coastal regions have been awarded status under some of a basis for the development of mitigation strategies, while the national schemes, expanding these initiatives more those that document the impacts of artificial lighting in widely to include “marine dark sky parks” would be a posi- the environment provide the impetus for doing so. tive step toward protecting currently dark regions of the Long-term databases offer opportunities to quantify marine environment against encroaching light pollution, the role of light pollution in driving population fluctua- while capitalizing on benefits to society, the environment, tions, and to monitor the success of preventative and and economic gains through nature tourism. remedial measures. An example of one such monitoring Artificial light pollution has only recently become scheme is the British Antarctic Survey’s bird strike log, widely recognized as an environmental issue. Statutory where bird strikes on ships are digitally recorded. tools for mitigating against its ecological impacts are Rolling out such a program across shipping globally therefore largely lacking in marine environments. At pre- would help to identify “hotspots” of human–wildlife sent, the International Convention for the Prevention of conflict, where operational procedures could be put in Pollution from Ships (MARPOL) does not recognize arti- place to minimize ecological impact. The data gener- ficial light as a pollutant. Some examples of voluntary mit- ated also offer the capability to identify seasonal and cli- igation measures have, however, been adopted by individ- matic influences on bird strike occurrences, and to eval- ual nations in sensitive areas such as the Antarctic, uate the success of mitigation measures such as although these guidelines have yet to be adopted formally switching off deck lighting during non-operational by all Antarctic Treaty Committee members (Secretariat hours and installing lights with narrow emission spectra of the Antarctic Treaty 2010). Indeed, in many cases it is that reduce bird collision incidents (Secretariat of the difficult for policy makers to introduce statutory mitiga- Antarctic Treaty 2010; Merkel 2010). tion measures where these conflict with local economic To appreciate the consequences of marine light pollution gains or operational safety issues and there is a limited for the goods and services provided to humanity by marine understanding of the severity of the problem. This is high- ecosystems, investigators can focus on whether direct lighted in the European Commission Marine Strategy impacts on one species can result in cascading impacts Framework Directive (Commission decision 2010/ throughout ecological networks. For instance, while there 477/EU; MFSD 2010). Despite being recognized under is evidence that artificial light facilitates nocturnal visual Descriptor 11, the Commission did not specify any formal foraging in wading birds (Dwyer et al. 2013), it is not criteria for light, stating that “Additional scientific and known how this affects the macro-infaunal communities technical progress is still required” (MFSD 2010). As a on which the birds feed or the ecosystem processes these result, standards defining environmentally “acceptable” communities perform. Many of the ecological responses to levels of light pollution in marine waters are unlikely to artificial light discussed here could lead to secondary effects feature in the corresponding legislative tools drawn up by

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354 the member nations of the European Union (EU). A con- Cohen JH and Forward RB. 2009. Zooplankton diel vertical migra- certed research effort is clearly needed to document and tion – a review of proximate control. Oceanogr Mar Biol 47: understand the environmental impacts of light pollution 77–110. in the marine environment so that effective statutory pro- Crisp DJ and Ritz DA. 1973. Responses of cirripede larvae to light. I. Experiments with white light. Mar Biol 23: 327–35. tective measures can be developed. Dacke M, Nilsson D-E, Scholtz CH, et al. 2003. Animal behaviour: insect orientation to polarized moonlight. Nature 424: 33. n Conclusion Davies TW, Bennie J, and Gaston KJ. 2012. Street lighting changes the composition of invertebrate communities. Biol Lett 8: Artificial light pollution is a global environmental issue, 764–67. the ecological impacts of which are only now beginning Davies TW, Bennie J, Inger R, et al. 2013a. 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U.S. Environmental Protection Agency Office of Research and Development National Exposure Research Laboratory HIGH-LEVEL CAREER OPPORTUNITY Computational Exposure Scientist Announcement Number: RTP-ORD-42-2014-0004

EPA’s Office of Research and Development (ORD), National Exposure Research Laboratory (NERL), is seeking a highly recognized scientific leader to direct and conduct a cutting edge research program in the area of computational exposure science. The position will be located in Research Triangle Park, North Carolina. NERL is leading the development of approaches to exposure science that take advantage of new advances in information and monitoring tech- nology and large new sources of data to improve the rate and scope of exposure assessments. This new approach, “computational exposure science,” integrates advances in informatics, chemistry, computer sciences, mathematics, statistics and social and behavioral sciences with new and numerically efficient models to more effectively predict real-world exposures.

More information on ORD’s NERL can be found at: www.epa.gov/nerl

The incumbent will develop and implement complex computational solutions that facilitate the use of “big data” to develop an understanding of the factors that lead to exposure and to develop rapid exposure and dose exposure for chemicals. In doing so, the incumbent will utilize knowledge of advanced informatics methods, tools and resources in chemistry, environmental science, ex- posure science, and behavioral science to interpret and understand the importance of various data sets relative to characterizing human and ecological exposure. Salary and Benefits: This is a Title 42 authority appointment. The position is a full-time, renewable, 5-year term appointment. Determination of salary is dependent upon qualifications, experience, and other factors (e.g., current salary, market salary rates). The selected applicant will be eligible for full benefits including relocation expenses, health and life insurance, retirement, and vacation and sick leave. Applicants must be U.S. citizens or permanent residents. EPA is an Equal Opportunity Employer, and selection for this position will be based solely on merit without regard to race, color, religion, age, gender, national origin, political affiliation, disability, sexual orientation, marital or family status or any other non-merit factors. Specific information on the position and instructions for applying can be found at: www.usajobs.gov/ Applications must be received by 11:59 p.m. ET on August 15, 2014, in order to be considered. Applications will not be considered if submitted via USAJOBS.

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