Explore the vast range of titles available.

Marine species exhibit a multitude of biological rhythms, in accordance with their complex ecosystem governed by sun, earth and moon trajectories. Because of the inclination of the earth’s axis, the high Arctic ecosystem is characterized by several months of permanent illumination during the polar day. The persistence of biological rhythms in this photic context remains unclear. Yet, this information is crucial for the understanding of polar ecosystems functioning, as well as to predict the impact of future climate changes. Particularly, the impact of extreme photoperiods on recent invasive species remains largely unknown. Here, we investigate how environmental cycles shape the behavior of a re-emerging polar resident, the mussel Mytilus sp. during polar day (17 April to 26 August 2020; Svalbard, Ny-Ålesund, 78°56ʹ N, 11°56ʹ E). Our results show that in the high Arctic polar day, mussels’ behavior is shaped by both the photoperiod and the diel sun trajectories above the horizon. In addition, mussels also exhibit tidal, semi-lunar, and lunar rhythms of valve opening amplitude. We argue that these rhythms may have ecosystems functioning implications, and that the mussels’ ability to deal with drastic light regimes may explain their northward expansion and new resettlement in high Arctic.

Many species synchronize their physiology and behavior to specific hours. It is commonly assumed that sunlight acts as the main entrainment signal for ∼24-h clocks. However, the moon provides similarly regular time information. Consistently, a growing number of studies have reported correlations between diel behavior and lunidian cycles. Yet, mechanistic insight into the possible influences of the moon on ∼24-h timers remains scarce. We have explored the marine bristleworm Platynereis dumerilii to investigate the role of moonlight in the timing of daily behavior. We uncover that moonlight, besides its role in monthly timing, also schedules the exact hour of nocturnal swarming onset to the nights’ darkest times. Our work reveals that extended moonlight impacts on a plastic clock that exhibits <24 h (moonlit) or >24 h (no moon) periodicity. Abundance, light sensitivity, and genetic requirement indicate that the Platynereis light receptor molecule r-Opsin1 serves as a receptor that senses moonrise, whereas the cryptochrome protein L-Cry is required to discriminate the proper valence of nocturnal light as either moonlight or sunlight. Comparative experiments in Drosophila suggest that cryptochrome’s principle requirement for light valence interpretation is conserved. Its exact biochemical properties differ, however, between species with dissimilar timing ecology. Our work advances the molecular understanding of lunar impact on fundamental rhythmic processes, including those of marine mass spawners endangered by anthropogenic change.

Synchronized mass coral spawning typically occurs several days after a full moon once a year. It is expected that spawning day is determined by corals sensing environmental change regulated by the lunar cycle (i.e., tide or moonlight); however, the exact regulatory mechanism remains unknown. Here, we demonstrate how moonlight influences the spawning process of coral, Dipsastraea speciosa. When corals in the field were shaded 1 and 3 d before the full moon or 1 d after the full moon, spawning always occurred 5 d after shading commenced. These results suggest moonlight suppresses spawning: a hypothesis supported by laboratory experiments in which we monitored the effects of experimental moonlight (night-light) on spawning day. Different night-light treatments in the laboratory showed that the presence of a dark period between day-light and night-light conditions eliminates the suppressive effect of night-light on spawning. In nature, moonrise gets progressively later during the course of the lunar cycle, shifting to after sunset following the day of the full moon. Our results indicate that this period of darkness between sunset and moonrise triggers synchronized mass spawning of D. speciosa in nature.

The lunar cycle has a major influence on the daily structure of marine ecosystems. As the moon orbits the earth across an approximate 29.5 day cycle, it influences these systems by altering nighttime light availability and shaping the strength and timing of tides. Numerous studies have documented the effects of the lunar cycle on large epipelagic fishes (here referring to tuna, billfish, sharks and rays), however, there has been no concerted effort to systematically compare these patterns across studies. Here, we review 190 studies documenting the effects of the lunar cycle on the ecology of large epipelagic fishes and discuss the potential underlying factors that contribute to the observed patterns. Most studies focused on fisheries science and movement ecology, examining metrics such as catch rate and depth of tagged individuals, respectively. A smaller proportion of studies delved into foraging behaviors and behavioral patterns. The effects observed varied among study types and taxa, yet vertical movement patterns consistently indicated a trend of deeper movements with increasing lunar illumination. Many factors likely contribute to this variation, including study specific methods (both field and analytical), other light sources or obstructions in the marine environment (such as bioluminescence and cloud cover), local site variation (such as local oceanography and prey distribution), species specific traits (such as distribution and foraging strategy) and individual traits (such as ontogenetic stage and body condition). We propose best practices for future studies on lunar effects, aimed at addressing this variation and promoting comparative analyses. Graphical abstract

Marine organisms adapt to complex temporal environments that include daily, tidal, semi-lunar, lunar and seasonal cycles. However, our understanding of marine biological rhythms and their underlying molecular basis is mainly confined to a few model organisms in rather simplistic laboratory settings. Here, we use new empirical data and recent examples of marine biorhythms to highlight how field ecologists and laboratory chronobiologists can complement each other's efforts. First, with continuous tracking of intertidal shorebirds in the field, we reveal individual differences in tidal and circadian foraging rhythms. Second, we demonstrate that shorebird species that spend 8–10 months in tidal environments rarely maintain such tidal or circadian rhythms during breeding, likely because of other, more pertinent, temporally structured, local ecological pressures such as predation or social environment. Finally, we use examples of initial findings from invertebrates (arthropods and polychaete worms) that are being developed as model species to study the molecular bases of lunar-related rhythms. These examples indicate that canonical circadian clock genes (i.e. the homologous clock genes identified in many higher organisms) may not be involved in lunar/tidal phenotypes. Together, our results and the examples we describe emphasize that linking field and laboratory studies is likely to generate a better ecological appreciation of lunar-related rhythms in the wild. This article is part of the themed issue ‘Wild clocks: integrating chronobiology and ecology to understand timekeeping in free-living animals’.

Lunar cycle and moonlight exposure have significant impacts on animal behaviour and physiology. The presence or absence of moonlight, along with predictable changes in brightness throughout the lunar cycle, can shape reproduction, foraging, communication, and other aspects of an animal’s world. While it has been shown that invertebrates use the moonlight for orientation, little is known on the effect of the lunar cycle on migratory birds. We found that the lunar cycle affected the nocturnal migration activity of a diurnal songbird species, the Eurasian Skylark Alauda arvensis. The occurrence of birds increased with moon fraction, moonlight intensity and duration, while abundance correlated positively with increasing moonlight intensity. Our findings of increased migration activity in bright nights around full moon contradict previous assumptions that small bird migrants would avoid such nights due to increased predation pressure and decreased visibility of stars for orientation. We argue that migrants relying on visual cues for orientation might favour moonlit nights, while future studies should also test whether the position of the moon can be used for navigation by birds.

Circadian clocks in terrestrial animals are encoded by molecular feedback loops involving the negative regulators PERIOD, TIMELESS or CRYPTOCHROME2 and positive transcription factors CLOCK and BMAL1/CYCLE. The molecular basis of circatidal (~12.4 hour) or other lunar-mediated cycles (~15 day, ~29 day), widely expressed in coastal organisms, is unknown. Disrupting circadian clockworks does not appear to affect lunar-based rhythms in several organisms that inhabit the shoreline suggesting a molecular independence of the two cycles. Nevertheless, pharmacological inhibition of casein kinase 1 (CK1) that targets PERIOD stability in mammals and flies, affects both circadian and circatidal phenotypes in Eurydice pulchra (Ep) , the speckled sea-louse. Here we show that these drug inhibitors of CK1 also affect the phosphorylation of Ep CLK and Ep BMAL1 and disrupt Ep CLK-BMAL1-mediated transcription in Drosophila S2 cells, revealing a potential link between these two positive circadian regulators and circatidal behaviour. We therefore performed dsRNAi knockdown of Epbmal1 as well as the major negative regulator in Eurydice , Epcry2 in animals taken from the wild. Epcry2 and Epbmal1 knockdown disrupted Eurydice ’s circadian phenotypes of chromatophore dispersion, tim mRNA cycling and the circadian modulation of circatidal swimming, as expected. However, circatidal behaviour was particularly sensitive to Epbmal1 knockdown with consistent effects on the power, amplitude and rhythmicity of the circatidal swimming cycle. Thus, three Eurydice negative circadian regulators, Ep CRY2, in addition to Ep PER and Ep TIM (from a previous study), do not appear to be required for the expression of robust circatidal behaviour, in contrast to the positive regulator Ep BMAL1. We suggest a neurogenetic model whereby the positive circadian regulators Ep BMAL1-CLK are shared between circadian and circatidal mechanisms in Eurydice but circatidal rhythms require a novel, as yet unknown negative regulator.

The lunar cycle is known to affect the behaviour of strictly nocturnal species, but for diurnal species that are periodically active during the night, this has been less investigated. Nocturnal bird song is relatively common in diurnal species, yet research on this behaviour accounts for little of the research on avian vocalisations. This is surprising given that diurnal species are adapted for bright environments and therefore may be particularly sensitive to change in the lunar cycles. We used automated bioacoustic recorders and automatic song detection software to measure nocturnal song rate in a diurnal bird where both sexes sing, the willie wagtail (Rhipidura leucophrys). We deployed recorders at eight locations across four naturally dark sites resulting in 457 h of nocturnal audio. We confirmed anecdotal evidence suggesting that willie wagtails are prolific nocturnal singers during the breeding season and demonstrate that while both male and females sing during the day, nocturnal song is largely sung by males. Moreover, we show that nocturnal song increased with lunar illumination, contrasting with previous research on other diurnal species that sing at night. Our data allow us to hypothesise possible functions for nocturnal song in this species, such as territory defence or mate attraction.

Planktonic foraminifera preserved in marine sediments archive the physical and chemical conditions under which they built their shells. To interpret the paleoceanographic information contained in fossil foraminifera, the recorded proxy signals have to be attributed to the habitat and life cycle characteristics of individual species. Much of our knowledge on habitat depth is based on indirect methods, which reconstruct the depth at which the largest portion of the shell has been calcified. However, habitat depth can be best studied by direct observations in stratified plankton nets. Here we present a synthesis of living planktonic foraminifera abundance data in vertically resolved plankton net hauls taken in the eastern North Atlantic during 12 oceanographic campaigns between 1995 and 2012. Live (cytoplasm-bearing) specimens were counted for each depth interval and the vertical habitat at each station was expressed as average living depth (ALD). This allows us to differentiate species showing an ALD consistently in the upper 100 m (e.g., Globigerinoides ruber white and pink), indicating a shallow habitat; species occurring from the surface to the subsurface (e.g., Globigerina bulloides, Globorotalia inflata, Globorotalia truncatulinoides); and species inhabiting the subsurface (e.g., Globorotalia scitula and Globorotalia hirsuta). For 17 species with variable ALD, we assessed whether their depth habitat at a given station could be predicted by mixed layer (ML) depth, temperature in the ML and chlorophyll a concentration in the ML. The influence of seasonal and lunar cycle on the depth habitat was also tested using periodic regression. In 11 out of the 17 tested species, ALD variation appears to have a predictable component. All of the tested parameters were significant in at least one case, with both seasonal and lunar cyclicity as well as the environmental parameters explaining up to > 50 % of the variance. Thus, G. truncatulinoides, G. hirsuta and G. scitula appear to descend in the water column towards the summer, whereas populations of Trilobatus sacculifer appear to descend in the water column towards the new moon. In all other species, properties of the mixed layer explained more of the observed variance than the periodic models. Chlorophyll a concentration seems least important for ALD, whilst shoaling of the habitat with deepening of the ML is observed most frequently. We observe both shoaling and deepening of species habitat with increasing temperature. Further, we observe that temperature and seawater density at the depth of the ALD were not equally variable among the studied species, and their variability showed no consistent relationship with depth habitat. According to our results, depth habitat of individual species changes in response to different environmental and ontogenetic factors and consequently planktonic foraminifera exhibit not only species-specific mean habitat depths but also species-specific changes in habitat depth.