Is Ambient Light during the High Arctic Polar Night Sufficient to Act as a Visual Cue for Zooplankton?

The light regime is an ecologically important factor in pelagic habitats, influencing a range of biological processes. However, the availability and importance of light to these processes in high Arctic zooplankton communities during periods of ''complete'' darkness (polar night) are poorly studied. Here we characterized the ambient light regime throughout the diel cycle during the high Arctic polar night, and ask whether visual systems of Arctic zooplankton can detect the low levels of irradiance available at this time. To this end, light measurements with a purpose-built irradiance sensor and coupled all-sky digital photographs were used to characterize diel skylight irradiance patterns over 24 hours at 79°N in January 2014 and 2015. Subsequent skylight spectral irradiance and in-water optical property measurements were used to model the underwater light field as a function of depth, which was then weighted by the electrophysiologically determined visual spectral sensitivity of a dominant high Arctic zooplankter, Thysanoessa inermis. Irradiance in air ranged between 1–1.5 x 10^-5 μmol photons m^-2 s^-1 (400–700 nm) in clear weather conditions at noon and with the moon below the horizon, hence values reflect only solar illumination. Radiative transfer modelling generated underwater light fields with peak transmission at blue-green wavelengths, with a 465 nm transmission maximum in shallow water shifting to 485 nm with depth. To the eye of a zooplankter, light from the surface to 75 m exhibits a maximum at 485 nm, with longer wavelengths (>600 nm) being of little visual significance. Our data are the first quantitative characterisation, including absolute intensities, spectral composition and photoperiod of biologically relevant solar ambient light in the high Arctic during the polar night, and indicate that some species of Arctic zooplankton are able to detect and utilize ambient light down to 20–30m depth during the Arctic polar night.     

A marine zooplankton community vertically structured by light across diel to interannual timescales

The predation risk of many aquatic taxa is dominated by visually searching predators, commonly a function of ambient light. Several studies propose that changes in visual predation will become a major climate-change impact on polar marine ecosystems. The High Arctic experiences extreme seasonality in the light environment, from 24 h light to 24 h darkness, and therefore provides a natural laboratory for studying light and predation risk over diel to seasonal timescales. Here, we show that zooplankton (observed using acoustics) in an Arctic fjord position themselves vertically in relation to light. A single isolume (depth-varying line of constant light intensity, the value of which is set at the lower limit of photobehaviour reponses of Calanus spp. and krill) forms a ceiling on zooplankton distribution. The vertical distribution is structured by light across timescales, from the deepening of zooplankton populations at midday as the sun rises in spring, to the depth to which zooplankton ascend to feed during diel vertical migration. These results suggest that zooplankton might already follow a foraging strategy that will keep visual predation risk roughly constant under changing light conditions, such as those caused by the reduction of sea ice, but likely with energetic costs such as lost feeding opportunities as a result of altered habitat use.     

Diving patterns of a Ross seal (Ommatophoca rossii ) near the eastern coast of the Antarctic Peninsula

In January 1987 we documented the diving patterns of a female Ross seal (Ommatophoca rossii) in the marginal pack-ice zone near the eastern coast of the Antarctic Peninsula for 2 days using a microprocessor-based time-depth recorder. The seal hauled out during the day and dived continually when in the water at night. Dives averaged 110 m deep and 6.4 min long; the deepest dive was 212 m and the longest 9.8 min. Dives were deepest near twilight and shallowest at night; this pattern suggests that the seal's prey, presumably mid-water squid and fish, may have been making vertical migrations or changing predator-avoidance behavior in response to diel light patterns. The dives of this Ross seal were substantially deeper, on average, than those of crabeater seals (Lobodon carcinophagus), which forage in the same areas on Antarctic krill (Euphausia superba). Received: 15 August 1996 / Accepted: 22 February 1997     

Cormorants dive through the Polar night

Most seabirds are visual hunters and are thus strongly affected by light levels. Dependence on vision should be problematic for species wintering at high latitudes, as they face very low light levels for extended periods during the Polar night. We examined the foraging rhythms of male great cormorants (Phalacrocorax carbo) wintering north of the Polar circle in West Greenland, conducting the first year-round recordings of the diving activity in a seabird wintering at high latitudes. Dive depth data revealed that birds dived every day during the Arctic winter and did not adjust their foraging rhythms to varying day length. Therefore, a significant proportion of the dive bouts were conducted in the dark (less than 1 lux) during the Polar night. Our study underlines the stunning adaptability of great cormorants and raises questions about the capacity of diving birds to use non-visual cues to target fish.     

The diel activity pattern of mountain hare (Lepus timidus) on managed heather moorland in Scotland

The research presented in this paper provides an insight into the behavioral ecology of mountain hares on heather moorland in the Lammermuir Hills of southeast Scotland. We examine the seasonal and diel activity patterns using camera traps over a period of 12 months. The rate of camera detections was calculated for the different divisions of the 24‐hr cycle (daylight, dusk, night, and dawn). During autumn and winter (October–February), the activity pattern was crepuscular with greater activity at dusk than at dawn. Daylight activity was relatively low, and there was a regular pattern of small peaks of activity during the night. In spring and summer (March–September), peaks of crepuscular activity remained evident but daylight activity was much more prevalent than during autumn and winter, and night activity was lower. We discuss the problematic definition of twilight and present an explanation for seasonal changes in the pattern of diel activity that is linked to the reproductive cycle of the mountain hare.     

Temperature effects on low-light vision in juvenile rockfish (Genus <Emphasis Type="Italic">Sebastes</Emphasis>) and consequences for habitat utilization

The absolute low-light sensitivity of four congeneric species of rockfish (genus Sebastes) was studied from analysis of electroretinograms measured in living fish. The purpose was: (1) to determine if temperature sensitive noise in rod photoreceptors affects the absolute limit to low-light sensitivity at environmentally realistic temperatures and light levels, and (2) to examine whether interspecific variations in habitat utilization within rockfish communities correlate with differences in visual sensitivity. It was found that the low-light sensitivity of individual retinae is inversely dependent on temperature, decreasing tenfold with a 10°C increase in temperature. While in all four species, temperature had a similar effect on sensitivity, the absolute sensitivity levels were different. The four species could be divided into two groups based on measured sensitivity. Kelp and olive rockfish form a high-sensitivity group capable of responding to light levels approximately 50-fold lower than blue and black rockfish. The sensitivity groups correlated with reported diel activity patterns; the high-sensitivity group forages nocturnally, whereas members of the low-sensitivity group are quiescent during twilight and night and forage during the day.     

The status and future of the Lake Naivasha fishery,Kenya

I describe a laboratory system for investigating the role of light as a proximate cue for diel changes in locomotor activity and vertical location on the substrate of stream macro-invertebrates. The system consisted of computer-controlled halogen lamps positioned over a laboratory stream in which video-recordings were made of Stenonema modestum mayfly nymphs located on the undersides of unglazed tile substrates. Locomotor activity of study organisms in response to light changes were quantified during computer-programmed and reproducible light/dark (LD) cycles. The system provided the flexibility to simulate a variety of light environments so that the separate influences of light intensity and light change on diel activities of individuals and populations could be examined, which is difficult under natural light conditions. As a group, nymphs responded similarly to simulated twilight (light decrease from 7.9 × 10² to 6.9 × 10^–2 W cm^–2 at a constant –1.9 × 10^–3 s^–1 rate of relative light change) and to natural twilight, suggesting that proposed mechanisms of light control of diel activities in nature can be adequately tested in the simulated environment. However, locomotor activity and vertical movements among individual mayflies were highly variable under controlled conditions, suggesting that physiological differences influence their responses to environmental conditions.     

Colour As a Signal for Entraining the Mammalian Circadian Clock

Twilight is characterised by changes in both quantity (“irradiance”) and quality (“colour”) of light. Animals use the variation in irradiance to adjust their internal circadian clocks, aligning their behaviour and physiology with the solar cycle. However, it is currently unknown whether changes in colour also contribute to this entrainment process. Using environmental measurements, we show here that mammalian blue–yellow colour discrimination provides a more reliable method of tracking twilight progression than simply measuring irradiance. We next use electrophysiological recordings to demonstrate that neurons in the mouse suprachiasmatic circadian clock display the cone-dependent spectral opponency required to make use of this information. Thus, our data show that some clock neurons are highly sensitive to changes in spectral composition occurring over twilight and that this input dictates their response to changes in irradiance. Finally, using mice housed under photoperiods with simulated dawn/dusk transitions, we confirm that spectral changes occurring during twilight are required for appropriate circadian alignment under natural conditions. Together, these data reveal a new sensory mechanism for telling time of day that would be available to any mammalian species capable of chromatic vision.     

Aboveground activity rhythm in Arctic black-capped marmot (Marmota camtschatica bungei Katschenko 1901) under polar day conditions

Daily aboveground activity of wild black-capped marmots of Yakutia (Marmota camtschatica bungei) was recorded under ‘polar day’ conditions at 71°56’ N and 127°19’ E (north of the Polar Circle). From the beginning of May until the end of August, the sun was permanently above or close to the horizon. However under this condition of continuous lighting, the aboveground activity of these arctic hibernating mammals was periodic. Onset and end of activity showed marked changes throughout the seasons. Activity time increased strongly from hibernation emergence until the end of July and then decreased slowly until onset of hibernation. Below daily mean temperatures of 5 °C, activity started when the sun was 35° above the horizon, and ended when it dropped below 28°. When daily mean temperatures were above 5 °C, activity onset was synchronised with a solar altitude around 17–18° and activity ended at 10°. Activity onset was more precise relative to the solar altitude than the end of activity. This may be explained by late feeding bouts, following a midday thermal stress. In absence of rapid natural light-dark (LD) transitions that occur at civil twilight, our results suggest that the activity pattern of black-capped marmots may be synchronised by the light cycle through the solar altitude and ambient temperature.     

Effects of Whaling on the Structure of the Southern Ocean Food Web: Insights on the “Krill Surplus” from Ecosystem Modelling

The aim of this study was to examine the ecological plausibility of the “krill surplus” hypothesis and the effects of whaling on the Southern Ocean food web using mass-balance ecosystem modelling. The depletion trajectory and unexploited biomass of each rorqual population in the Antarctic was reconstructed using yearly catch records and a set of species-specific surplus production models. The resulting estimates of the unexploited biomass of Antarctic rorquals were used to construct an Ecopath model of the Southern Ocean food web existing in 1900. The rorqual depletion trajectory was then used in an Ecosim scenario to drive rorqual biomasses and examine the “krill surplus” phenomenon and whaling effects on the food web in the years 1900–2008. An additional suite of Ecosim scenarios reflecting several hypothetical trends in Southern Ocean primary productivity were employed to examine the effect of bottom-up forcing on the documented krill biomass trend. The output of the Ecosim scenarios indicated that while the “krill surplus” hypothesis is a plausible explanation of the biomass trends observed in some penguin and pinniped species in the mid-20^th century, the excess krill biomass was most likely eliminated by a rapid decline in primary productivity in the years 1975–1995. Our findings suggest that changes in physical conditions in the Southern Ocean during this time period could have eliminated the ecological effects of rorqual depletion, although the mechanism responsible is currently unknown. Furthermore, a decline in iron bioavailability due to rorqual depletion may have contributed to the rapid decline in overall Southern Ocean productivity during the last quarter of the 20^th century. The results of this study underscore the need for further research on historical changes in the roles of top-down and bottom-up forcing in structuring the Southern Ocean food web.     

Diel food selection of pelagic Arctic charr, Salvelinus alpinus (L.), and brown trout, Salmo trutta L., in Lake Atnsjø, SE Norway

Patterns of diel food selection in pelagic Arctic charr, Salvelinus alpinus (L.) and brown trout, Salmo trutta L. were investigated in Lake Atnsjo, SE Norway, by gillnet sampling during July-September 1985. Arctic charr feed almost exclusively on zooplankton both day and night, while brown trout had a diurnal shift in diet. For this species zooplankton made up a considerable part of the diet in the daytime, while at night the diet consisted mainly of surface insect and chironomid pupae. Both species had a selective feeding mode on zooplankton during the day and night. Arctic charr had a higher gill raker number and a denser gill raker spacing compared with brown trout. Still, the differences in prey size between the two species were small. We argue that the observed differences in food selection between Arctic charr and brown trout can be explained by differing abilities to detect food items under low light conditions.     

To what extent do environmental factors affect the long-distance nocturnal post-fledging movements of the Reed Warbler?

We studied the effects of weather and the lunar cycle on long-distance nocturnal pre-migratory flights of Reed Warblers (Acrocephalus scirpaceus). Noturnal tape luring was used to capture the birds, and the study was carried out in a habitat atypical of this species on the Courish Spit (southeastern Baltic) between1999 and 2002. A total of 443 juvenile Reed Warblers were captured during 120 nights of trapping. Based on data on the moult and body condition of the birds, it was possible to identify 163 individuals as being on post-fledging movements. More than half of the birds (54.0%) were captured at the end of night during the nautical and civil twilight period; the remaining individuals were caught during the astronomical twilight period and darkest periods of night. Many birds performed pre-migratory flights in the middle of the night, the period during which a large part of the moon was visible. We suggest that the increased visibility under the full moon may provide conditions in which Reed Warblers increase the distance or intensity of nocturnal post-fledging movements. During the entire course of the night, the birds generally preferred to fly under rainless conditions, limited cloud cover and/or in still air. The birds performed flights under winds stronger than 2 m s⁻¹ when the wind was blowing along the axis of the spit. We also suggest that the birds generally fly along the spit. We found a weak but significant relationship between the numbers of Reed Warblers captured during post-fledging movements and the individual weather parameters and their interaction. Our data suggest that endogenous stimuli rather than weather parameters or lunar cycle phase determine the decision of Reed Warblers to undertake pre-migratory long-distance flights at night.     

Estimation of the Horizon in Photographed Outdoor Scenes by Human and Machine

We present three experiments on horizon estimation. In Experiment 1 we verify the human ability to estimate the horizon in static images from only visual input. Estimates are given without time constraints with emphasis on precision. The resulting estimates are used as baseline to evaluate horizon estimates from early visual processes. Stimuli are presented for only ms and then masked to purge visual short-term memory and enforcing estimates to rely on early processes, only. The high agreement between estimates and the lack of a training effect shows that enough information about viewpoint is extracted in the first few hundred milliseconds to make accurate horizon estimation possible. In Experiment 3 we investigate several strategies to estimate the horizon in the computer and compare human with machine “behavior” for different image manipulations and image scene types.     

Living planktonic foraminifera in the Fram Strait (Arctic): absence of diel vertical migration during the midnight sun

The timing of vertical migration in planktonic foraminifera (ex. ontogenetic, diel) is still an open debate. This work aims to investigate the diel vertical migration (DVM) of Neogloboquadrina pachyderma (N. pachyderma) and Turborotalita quinqueloba (T. quinqueloba) in the Arctic during the midnight sun. N. pachyderma and T. quinqueloba dominate the total assemblage in the cold Polar Water and warmer North Atlantic Water masses, respectively. Foraminifera were collected at several depths along the Fram Strait. Afterwards sampling was performed at the same station for 24 h at continuous and discrete time intervals. Results show no evidence of planktonic foraminifera DVM since there was no significant variability in the abundance and size distribution during the 24-h collection period. This finding provides information to improve the interpretation of foraminifera in paleoclimatic works. This is especially relevant in the Fram Strait as paleoclimatic studies in this region are fundamental to investigating the history of the Atlantic water inflow into the Arctic Ocean.     

Diel vertical migration and feeding of chaetognaths in coastal waters of the eastern Mediterranean

This study investigates the diel vertical distribution and the diet of the most important chaetognath species found in the 0–50 m surface layer of a coastal area in the eastern Mediterranean during a 24-hour period in July 2004. Among the recorded chaetognaths, Sagitta enflata was the most abundant species (41.6%), followed by S. minima (32.5%) and S. serratodentata (20.8%). Those three species exhibited a “twilight migration” pattern, with only small differences among them. Vertical separation was found between S. enflata and S. minima. Both species preyed mainly on cladocerans, although copepods were the most abundant group in the zooplankton assemblage. The chaetognath species followed partially the diel vertical migration of their prey. S. enflata showed high feeding intensity at different times in both day and night, while S. minima fed more intensively at midday (12:00) and at dusk (20:00), and S. serratodentata in the morning (08:00). It seems that in order to coexist in an area of low productivity the chaetognath species follow the basic ecological rules of space, time and food-type separation, in order to reduce the inter- and intra-specific competition. The high preference of S. minima and especially of S. enflata for the cladocerans made them probably the most important predators of cladocerans during summer.     

Diel variation in availability and vulnerability of fish to a survey trawl

The effect diel variation in availability and vulnerability has on the fish capture process of a groundfish survey trawl was examined from data collected during an experiment measuring fish escapement underneath the footgear. The survey trawl was significantly more efficient at night in capturing American plaice, Hippoglossoides platessoides, and yellowtail flounder, Limanda ferruginea, but no difference was found in catches of Atlantic cod, Gadus morhua. This increase in capture efficiency at night is related to an increase in availability and vulnerability associated with diel variation in light intensity. No evidence of diel vertical migration was found and changes in availability were linked to visual dependent gear avoidance by the fish. Regardless of light conditions, the survey trawl had a very low capture efficiency for juveniles of the three species studied. Escapement of this size group was extremely high during both day and night periods due to a fast towing speed, use of large bobbin gear, and possible mesh selection.     

An acoustic tracking of a megamouth shark, Megachasma pelagios: a crepuscular vertical migrator

A 4.9 m TL megamouth shark, only the sixth specimen known to science, was tracked continuously for 50.5 h, during which it exhibited distinct vertical migrations at the dawn and dusk transitions. The male shark was captured on 21 October 1990 in a drift gill net off Dana Point, California, restrained overnight in a harbor, and released at sea the next afternoon. Horizontally, the shark moved slowly southward, covering 62 km on a relatively straight path with no significant diel changes. For the major part of the tracking, its rate of movement was 1.15 km h^–1, as determined from positions at 15 min intervals. Considering a probable head current of 10–25 cm sec^–1, its estimated through-the-water swimming speed was more likely 1.5–2.1 km h^–1 (X¯ = 1.8, representing 0.1 body lengths sec^–1). Vertically, the shark stayed shallow at night (12–25 m depth range, X¯ = 17) and deep during the days (120–166 m, X¯ = 149) but still well above the bottom at 700–850 m. The four twilight depth-change events were very distinct and always spanned the times of sunset or sunrise. The ascent and descent profiles are a reasonble match to isolumes on the order of 0.4 lux for an extinction coefficient (0.07) calculated from water transparency measurements. Furthermore, the steepest parts of the shark's profiles correspond closely to the times of maximum rate-of-change of illumination. These findings suggest that, except during nights, the shark's chosen depth was to a large degree determined by light level.     

Trophic behavior of Euphausia superba Dana in laboratory conditions

Summary Experiments conducted at US Palmer Station, 1980 on antarctic krill, Euphausia superba, attempted 1) to quantify and describe behavioral features of ingestion, food clearance and egestion, and 2) to test the hypotheses that feeding and swarming are mutually exclusive events, and that feeding is a cyclical, diel phenomenon. Ingestion was quantified in a large flow-through aquarium. Food clearance and egestion were estimated visually and fluorometrically in E. superba individually kept in one liter jars. Ingestion was directly proportional to chlorophyll concentration (0.65–11.5 gChl/l) and did not change significantly as a function of krill density (200–9000 krill/m³). Ingestion and egestion did not show significant diel trends or mean day-night differences in a 16 h light-8 h dark cycle. Our results suggested that feeding may occur in nature: at swarm densities, within a wide range of phytoplankton concentrations, and may be sustained throughout the diel cycle. We propose that feeding and swarming are co-occurring events. The theoretical basis at the individual and group levels, and its implications, are briefly discussed.In memory of Mary Alice McWhinnie (1922–1980)This research was supported by the University of Chile, National Science Foundation and Chilean Antarctic Institute     

Hormone‐mediated foraging strategies in an uncertain environment: Insights into the at‐sea behavior of a marine predator

1. Hormones are extensively known to be physiological mediators of energy mobilization and allow animals to adjust behavioral performance in response to their environment, especially within a foraging context.
2. Few studies, however, have narrowed focus toward the consistency of hormonal patterns and their impact on individual foraging behavior. Describing these relationships can further our understanding of how individuals cope with heterogeneous environments and exploit different ecological niches.
3. To address this, we measured between‐ and within‐individual variation of basal cortisol (CORT), thyroid hormone T3, and testosterone (TEST) levels in wild adult female Galápagos sea lions (Zalophus wollebaeki) and analyzed how these hormones may be associated with foraging strategies. In this marine predator, females exhibit one of three spatially and temporally distinct foraging patterns (i.e., “benthic,” “pelagic,” and “night” divers) within diverse habitat types.
4. Night divers differentiated from other strategies by having lower T3 levels. Considering metabolic costs, night divers may represent an energetically conservative strategy with shorter dive durations, depths, and descent rates to exploit prey which migrate up the water column based on vertical diel patterns.
5. Intriguingly, CORT and TEST levels were highest in benthic divers, a strategy characterized by congregating around limited, shallow seafloors to specialize on confined yet reliable prey. This pattern may reflect hormone‐mediated behavioral responses to specific risks in these habitats, such as high competition with conspecifics, prey predictability, or greater risks of predation.
6. Overall, our study highlights the collective effects of hormonal and ecological variation on marine foraging. In doing so, we provide insights into how mechanistic constraints and environmental pressures may facilitate individual specialization in adaptive behavior in wild populations.