Splitting light pollution: Wavelength effects on the activity of two sandy beach species.

Artificial Light at Night (ALAN) threatens to disrupt most natural habitats and species, including those in coastal settings, where a growing number of studies have identified ALAN impacts. A careful examination of the light properties behind those impacts is important to better understand and manage the effects of this stressor. This study focused on ALAN monochromatic wavelengths and examined which types of light spectra altered the natural activity of two prominent coastal species from the Pacific southeast: the talitroid amphipod Orchestoidea tuberculata and the oniscoid isopod Tylos spinulosus. We compared the natural daylight/night activity of these organisms with the one they exhibit when exposed to five different ALAN wavelengths: lights in the violet, blue, green, amber, and red spectra. Our working hypothesis was that ALAN alters these species' activity at night, but the magnitude of such impact differs depending on light wavelengths. Measurements of activity over 24 h cycles for five consecutive days and in three separate experiments confirmed a natural circadian activity pattern in both species, with strong activity at night (∼90% of probability) and barely any activity during daylight. However, when exposed to ALAN, activity declined significantly in both species under all light wavelengths. Interestingly, amphipods exhibited moderate activity (∼40% of probability) when exposed to red lights at night, whereas isopods shifted some of their activity to daylight hours in two of the experiments when exposed to blue or amber lights, suggesting a possible alteration in this species circadian rhythm. Altogether, our results were consistent with our working hypothesis, and suggest that ALAN reduces night activity, and some wavelengths have differential effects on each species. Differences between amphipods and isopods are likely related to their distinct adaptations to natural low-light habitat conditions, and therefore distinct sensitivity to ALAN.

Artificial light at night alters the feeding activity and two molecular indicators in the plumose sea anemone Metridium senile (L.).

Artificial light at night (ALAN) is becoming a widespread stressor in coastal ecosystems, affecting species that rely on natural day/night cycles. Yet, studies examining ALAN effects remain limited, particularly in the case of sessile species. This study assessed the effects of ALAN upon the feeding activity and two molecular indicators in the widespread plumose sea anemone Metridium senile. Anemones were exposed to either natural day/night or ALAN conditions to monitor feeding activity, and tissue samples were collected to quantify proteins and superoxide dismutase (SOD) enzyme concentrations. In day/night conditions, sea anemones showed a circadian rhythm of activity in which feeding occurs primarily at night. This rhythm was altered by ALAN, which turned it into a reduced and more uniform pattern of feeding. Consistently, proteins and SOD concentrations were significantly lower in anemones exposed to ALAN, suggesting that ALAN can be harmful to sea anemones and potentially other marine sessile species.

Cross-examining the influence of upwelling and seaweed quality on herbivores' feeding behavior and growth.

At the regional scale, upwelling conditions are known to influence ecosystems and communities and their primary and secondary productivity. However, the influence of upwelling on local herbivore-algae interactions is less well understood. We address this question by cross-examining herbivores and seaweeds from sites associated with upwelling and downwelling conditions along the Humboldt Current System. Specifically, we quantified the feeding and benefits attained by the black sea urchin (Tetrapygus niger) and the black sea snail (Tegula atra) while consuming a widespread kelp species (Lessonia spicata). We hypothesized that food quality drives herbivores' preference, consumption, and growth rates, regardless of the origin or "prior" conditions of the consumers. Laboratory trials measured algal consumption rates with (preference) and without a choice, and consumer's growth rates, to assess the influence of food quality (algae from upwelling vs downwelling sites) and the site of origin of the consumers. Our results showed that algal quality was a prevailing factor for both herbivores: they chose, consumed more, and grew faster on high quality (upwelling) algae. By comparison, the origin of the consumer was only significant for sea snails: those coming from an upwelling site, consumed significantly more and grew faster than those from downwelling. The bulk of our results provided strong support to our hypothesis and suggest that the high nutritional quality of algae associated with upwelling centers has a strong influence on consumers' preferences, consumption, and performance (growth). The fact that origin was found to be relevant for one of the herbivores suggests that the conditions in which species grow may dictate some of their efficiency as consumers.

Artificial Light at Night (ALAN) causes size-dependent effects on intertidal fish decision-making.

Artificial Light at Night (ALAN) alters cycles of day and night, potentially modifying species' behavior. We assessed whether exposure to ALAN influences decision-making (directional swimming) in an intertidal rockfish (Girella laevisifrons) from the Southeastern Pacific. Using a Y-maze, we examined if exposure to ALAN or natural day/night conditions for one week affected the number of visits and time spent in three Y-maze compartments: dark and lit arms ("safe" and "risky" conditions, respectively) and a neutral "non-decision" area. The results showed that fish maintained in natural day/night conditions visited and spent more time in the dark arm, regardless of size. Instead, fish exposed to ALAN visited and spent more time in the non-decision area and their response was size-dependent. Hence, prior ALAN exposure seemed to disorient or reduce the ability of rock fish to choose dark conditions, deemed the safest for small fish facing predators or other potential threats.

Field experimental evidence of sandy beach community changes in response to artificial light at night (ALAN).

Artificial light at night (ALAN) is a pervasive but still under-recognized driver of global change. In coastal settings, a large majority of the studies assessing ALAN impacts has focused on individual species, even though it is unclear whether results gathered from single species can be used to predict community-wide responses. Similarly, these studies often treat species as single life-stage entities, ignoring the variation associated with distinct life stages. This study addresses both limitations by focusing on the effects of ALAN on a sandy beach community consisting of species with distinct early- and late-life stages. Our hypothesis was that ALAN alters community structure and these changes are mediated by individual species and also by their ontogenetic stages. A field experiment was conducted in a sandy beach of north-central Chile using an artificial LED system. Samples were collected at different night hours (8-levels in total) across the intertidal (9-levels) over several days in November and January (austral spring and summer seasons). The abundance of adults of all species was significantly lower in ALAN treatments. Early stages of isopods showed the same pattern, but the opposite was observed for the early stages of the other two species. Clear differences were detected in the zonation of these species during natural darkness versus those exposed to ALAN, with some adult-juvenile differences in this response. These results support our hypothesis and document a series of changes affecting differentially both early and late life stages of these species, and ultimately, the structure of the entire community. Although the effects described correspond to short-term responses, more persistent effects are likely to occur if ALAN sources become established as permanent features in sandy beaches. The worldwide growth of ALAN suggests that the scope of its effect will continue to grow and represents a concern for sandy beach systems.

Hemocyanin as a biological indicator of artificial light at night stress in sandy beach amphipods.

The influence of artificial light at night (ALAN) is becoming evident in marine sandy beaches. These habitats are dominated by species reliant on natural daylight/night regimes, making the identification of biological indicators a priority. We assessed the applicability of hemocyanin, an oxygen-transport protein in the hemolymph of many invertebrates, as an indicator of ALAN-related stress. Unlike total proteins, hemocyanins signal metabolic function and stress, so we expected them to increase in response to ALAN. We adapted spectrophotometry protocols to describe spatial variation in hemocyanins and total proteins in four populations of the talitroid amphipod Americorchestia longicornis. Then, a two-week experiment tested for changes in response to ALAN. Hemocyanin levels increased by 17 % and 40 % with respect to experimental controls after 7 and 14 d, respectively, and were higher than any measurements conducted in the field. These results suggest good prospects for hemocyanin as an indicator of ALAN effects.

Artificial light at night (ALAN) causes variable dose-responses in a sandy beach isopod.

Artificial Light at Night (ALAN) is expanding worldwide, and the study of its influence remains limited mainly to documenting impacts, overlooking the variation in key characteristics of the artificial light such as its intensity. The potential dose-response of fitness-related traits to different light intensities has not been assessed in sandy beach organisms. Hence, this study explored dose-responses to ALAN by exposing the intertidal sandy beach isopod Tylos spinulosus to a range of light intensities at night: 0 (control), 20, 40, 60, 80 and 100 lx. We quantified the response of this species at the molecular (RNA:DNA ratios), physiological (absorption efficiency) and organismal (growth rate) levels. Linear and non-linear regressions were used to explore the relationship between light intensity and the isopod response. The regressions showed that increasing light intensity caused an overall ~ threefold decline in RNA:DNA ratios and a ~ threefold increase in absorption efficiency, with strong dose-dependent effects. For both response variables, non-linear regressions also identified likely thresholds at 80 lx (RNA:DNA) and 40 lx (absorption efficiency). By contrast, isopod growth rates were unrelated (unaltered) by the increase in light intensity at night. We suggest that ALAN is detrimental for the condition of the isopods, likely by reducing the activity and feeding of these nocturnal organisms, and that the isopods compensate this by absorbing nutrients more efficiently in order to maintain growth levels.

Artificial Light at Night (ALAN) negatively affects the settlement success of two prominent intertidal barnacles in the southeast Pacific.

Many coastal processes are regulated by day/night cycles and are expected to be altered by Artificial Light at Night (ALAN). The goal of this study was to assess the influence of ALAN on the settlement rates of intertidal barnacles. A newly designed settlement plate equipped with a small central LED light source was used to quantify settlement rates in presence/absence of ALAN conditions. "ALAN plates" as well as regular settlement plates were deployed in the mid rocky intertidal zone. Both ALAN and control plates collected early and late settlers of the barnacles Notochthamalus scabrosus and Jehlius cirratus. Early settlers (pre-metamorphosis cyprids) were not affected by ALAN. By contrast, the density of late settlers (post-metamorphosis spats) was significantly lower in ALAN than in control plates for both species, suggesting detrimental ALAN impacts on the settlement process. The new ALAN plates represent an attractive and alternative methodology to study ALAN effects.

Artificial light at night alters the activity and feeding behaviour of sandy beach amphipods and pose a threat to their ecological role in Atlantic Canada.

Artificial light at night (ALAN) is a growing source of stress for organisms and communities worldwide. These include species associated with sandy beaches, which consume and process stranded seaweeds (wrack) in these ecosystems. This study assessed the influence of ALAN on the activity and feeding behaviour of Americorchestia longicornis, a prominent talitrid amphipod living in sandy beaches of Prince Edward Island, Atlantic Canada. First, two parallel field surveys were conducted to document the natural daily cycle of activity of this species. Then, three related hypotheses were used to assess whether ALAN disrupts its locomotor activity, whether that disruption lasts over time, and whether it affects the feeding behaviour and growth of the amphipods. Tanks equipped with actographs recorded amphipod locomotor activity for ~7 days and then its potential recovery (after ALAN removal) for additional ~3 days. Separate tanks were used to compare amphipod food consumptions rates, absorption efficiency and growth rates under natural daylight / night (control) and altered conditions (ALAN). The results of these manipulations provide support to two of the three hypotheses proposed and indicate that ALAN was temporarily detrimental for (i.e. significantly reduced) the surface activity, consumption rates and absorption efficiency of the amphipods, whereas growth rates remained unaffected. The results also rejected the remaining hypothesis and suggest that the plasticity exhibited by these amphipods confer them the capacity to recover their natural rhythm of activity shortly after ALAN was removed from the system. Combined, these results suggest that ALAN has a strong, albeit temporary, influence upon the abundant populations of A. longicornis. Such influence has implications for the ecosystem role played by these amphipods as consumers and processors of the subsidy of stranded seaweeds entering these ecosystems.

Effects of artificial light at night and predator cues on foraging and predator avoidance in the keystone inshore mollusc Concholepas concholepas.

The growth of Artificial Light At Night (ALAN) is potentially having widespread effects on terrestrial and coastal habitats. In this study we addressed both the individual effects of ALAN, as well as its combined effect with predation risk on the behaviour of Concholepas concholepas, a fishery resource and a keystone species in the southeastern Pacific coast. We measured the influence of ALAN and predation risk on this mollusc's feeding rate, use of refuge for light and crawling out of water behaviour. These behavioural responses were studied using light intensities that mimicked levels that had been recorded in coastal habitat exposed to ALAN. Cues were from two species known to prey on C. concholepas during its early ontogeny: the crab Acanthocyclus hassleri and the seastar Heliaster helianthus. The feeding rates of C. concholepas were 3-4 times higher in darkness and in the absence of predator cues. In contrast, ALAN-exposed C. concholepas showed lower feeding activity and were more likely to be in a refuge than those exposed to control conditions. In the presence of olfactory predator cues, and regardless of light treatment, C. concholepas tended to crawl-out of the waterline. We provide evidence to support the hypothesis that exposure to either ALAN or predation risk can alter the feeding behaviour of C. concholepas. However, predator cue recognition in C. concholepas was not affected by ALAN in situations where ALAN and predator cues were both present: C. concholepas continued to forage when predation risk was low, i.e., in darkness and away from predator cues. Whilst this response means that ALAN may not lead to increased predation mortality in C. concholepas, it will reduce feeding activity in this naturally nocturnal species in the absence of dark refugia. Such results may have implications for the long-term health, productivity and sustainability of this keystone species.

The influence of microplastics pollution on the feeding behavior of a prominent sandy beach amphipod, Orchestoidea tuberculata (Nicolet, 1849).

Pollution by microplastics has become a global threat affecting coastal habitats such as sandy beaches and their resident macrofauna. The goal of this study was to assess the influence of microplastics on the feeding behavior and growth rate of a widespread sandy beach amphipod, Orchestoidea tuberculata. These organisms were exposed to artificial food prepared with Poly(styrene-co-divinylbenzene) microspheres (8 µm particle size) at 3 different concentrations (0%, 5% and 10%). The amphipods consumed significantly more food when the concentration of microplastics was 0% and significantly less when the concentration was 10%, both in trials in which they had a choice (preference experiments) and those in which they did not have a food choice. In contrast to this, the amphipod's absorption efficiency and estimated growth rates were not significantly affected by the concentration of microplastics. Combined, these results indicate that high microplastics concentrations (e.g. 10%) cause a reduction in the amphipod's consumption rates and, indirectly, may affect the role of this species as a main consumer of stranded seaweeds in sandy beaches ecosystems.

Artificial light pollution at night (ALAN) disrupts the distribution and circadian rhythm of a sandy beach isopod.

Coastal habitats, in particular sandy beaches, are becoming increasingly exposed to artificial light pollution at night (ALAN). Yet, only a few studies have this far assessed the effects of ALAN on the species inhabiting these ecosystems. In this study we assessed the effects of ALAN on Tylos spinulosus, a prominent wrack-consumer isopod living in sandy beaches of north-central Chile. This species burrows in the sand during daylight and emerges at night to migrate down-shore, so we argue it can be used as a model species for the study of ALAN effects on coastal nocturnal species. We assessed whether ALAN alters the distribution and locomotor activity of this isopod using a light system placed in upper shore sediments close to the edge of the dunes, mimicking light intensities measured near public lighting. The response of the isopods was compared to control transects located farther away and not exposed to artificial light. In parallel, we measured the isopods' locomotor activity in the laboratory using actographs that recorded their movement within mesocosms simulating the beach surface. Measurements in the field indicated a clear reduction in isopod abundance near the source of the light and a restriction of their tidal distribution range, as compared to control transects. Meanwhile, the laboratory experiments showed that in mesocosms exposed to ALAN, isopods exhibited reduced activity and a circadian rhythm that was altered and even lost after a few days. Such changes with respect to control mesocosms with a natural day/night cycle suggest that the changes observed in the field were directly related to a disruption in the locomotor activity of the isopods. All together these results provide causal evidence of negative ALAN effects on this species, and call for further research on other nocturnal sandy beach species that might become increasingly affected by ALAN.

Artificial light pollution influences behavioral and physiological traits in a keystone predator species, Concholepas concholepas.

Artificial Light At Night (ALAN) is an increasing global problem that, despite being widely recognized in terrestrial systems, has been studied much less in marine habitats. In this study we investigated the effect of ALAN on behavioral and physiological traits of Concholepas concholepas, an important keystone species of the south-eastern Pacific coast. We used juveniles collected in intertidal habitats that had not previously been exposed to ALAN. In the laboratory we exposed them to two treatments: darkness and white LED (Lighting Emitting Diodes) to test for the impacts of ALAN on prey-searching behavior, self-righting time and metabolism. In the field, the distribution of juveniles was observed during daylight-hours to determine whether C. concholepas preferred shaded or illuminated microhabitats. Moreover, we compared the abundance of juveniles collected during day- and night-time hours. The laboratory experiments demonstrated that juveniles of C. concholepas seek out and choose their prey more efficiently in darkened areas. White LED illuminated conditions increased righting times and metabolism. Field surveys indicated that, during daylight hours, juveniles were more abundant in shaded micro-habitats than in illuminated ones. However, during darkness hours, individuals were not seen to aggregate in any particular microhabitats. We conclude that the exposure to ALAN might disrupt important behavioral and physiological traits of small juveniles in this species which, as a mechanism to avoid visual predators, are mainly active at night. It follows that ALAN in coastal areas might modify the entire community structure of intertidal habitats by altering the behavior of this keystone species.

Living on a trophic subsidy: Algal quality drives an upper-shore herbivore's consumption, preference and absorption but not growth rates.

The transfer of seaweeds from subtidal bottoms to nearby intertidal rocky shores is a common but often overlooked phenomenon. Freshly detached seaweeds often represent critical trophic subsidies for herbivores living in upper-shore rocky intertidal areas, such as the marine snail Diloma nigerrima. This species relies on three species of seaweeds for food and displays feeding strategies to deal with a resource that is scarce and at times unpredictable. This study focused on the nutritional quality of freshly detached algae (Durvillaea antarctica, Lessonia spicata and Lessonia trabeculata) and measured Diloma nigerrima's algal consumption rates in trials with and without choice. Absorption efficiency and growth of individual snails fed on each alga were also measured. Durvillaea antarctica had the highest nutritional quality and was the most consumed algae in both single and multiple-choice trials. Absorption efficiency was also highest for D. antarctica but growth rates of snails fed with this species were similar to those fed with the other algae. Combined, these results suggest that D. nigerrima has the ability to discriminate among seaweeds based on their nutritional quality. A potential increase in oxygen uptake when D. nigerrima is consuming the preferred food item is also proposed as a plausible hypothesis to explain the mismatch between snails' preference and growth rate. These results aim to guide further studies on trophic subsidies and their role in coastal systems.