A field survey to investigate why nightjars frequent roads at night

Nightjars throughout the Afrotropical Region regularly frequent roads at night and during the twilight of dusk and dawn. Most species exhibit this behaviour and numerous individuals are killed by road traffic. Many theories have been advanced to explain the presence of nightjars on roads at night, but very little fieldwork has been carried out to test them. For a period of a year we carried out a weekly survey of nightjars on a selected road network near Harare in Zimbabwe. All nightjar encounters were documented in relation to variations in road surface, road width, adjacent habitat, arc of sky visible and other variables, such as the time of night, moon phase and weather conditions. None of these factors provided a complete explanation for the presence of nightjars on the roads surveyed. Neither were any of the nightjars seen dust bathing, taking grit or picking up insects from the road. Many were seen actively hunting flying insects, especially at dusk, and palpation of the stomachs of those caught confirmed that they had fed well during the evening hours. They were clearly using the road as an observation platform for hunting during the first few hours after sunset and then as a convenient place for resting and digesting. A nightjar sitting on an open road is certainly in a good position to see flying insects silhouetted against the twilight sky.     

The costs of becoming nocturnal: feeding efficiency in relation to light intensity in juvenile Atlantic Salmon

1. Most animals are active by day or by night, but not both; juvenile salmonids are unusual in that they switch from being predominantly diurnal for most of the year to being nocturnal in winter. They are visual foragers, and adaptations for high visual acuity at daytime light intensities are generally incompatible with sensitive night vision. Here we test whether juvenile Atlantic Salmon Salmo salar are able to maintain their efficiency of prey capture when switching between diurnal and nocturnal foraging.
2. By testing the ability of the fish to acquire drifting food items under a range of manipulated light intensities, we show that the foraging efficiency of juvenile salmon is high at light intensities down to those equivalent to dawn or dusk, but drops markedly at lower levels of illumination: even under the best night condition (full moon and clear sky), the feeding efficiency is only 35% of their diurnal efficiency, and fish will usually be feeding at less than 10% (whenever the moon is not full, skies are overcast or when in the shade of bankside trees). Fish were unable to feed on drifting prey when in complete darkness.
3. The ability of juvenile salmon to detect prey under different light intensities is similar to that of other planktivorous or drift-feeding species of fish; they thus appear to have no special adaptations for nocturnal foraging.
4. While winter drift abundance is slightly higher by night than by day, the difference is not enough to compensate for the loss in foraging efficiency. We suggest that juvenile salmon can nonetheless switch to nocturnal foraging in winter because their food requirements are low, many individuals adopting a strategy in which intake is suppressed to the minimum that ensures survival.     

Guiding lights: Foraging responses of juvenile nocturnal orb‐web spiders to the presence of artificial light at night

The reach of artificial light at night (ALAN) is growing rapidly around the globe, including the increasing use of energy‐efficient LED lights. Many studies document the physiological costs of light at night, but far fewer have focused on the potential benefits for nocturnal insectivores and the likely ecological consequences of shifts in predator–prey relationships. We investigated the effects of ALAN on the foraging behaviour and prey capture success in juvenile Australian garden orb‐web spiders (Eriophora biapicata). Laboratory experiments demonstrated that juvenile spiders were attracted to LED lights when choosing foraging sites, but prey availability was a stronger cue for remaining in a foraging site. Field experiments revealed a significant increase in prey capture rates for webs placed near LED lights. This suggests that any physiological costs of light at night may be offset by the foraging benefits, perhaps partially explaining recently observed increases in the size, fecundity and abundance of some orb‐web spider species in urban environments. Our results highlight the potential long‐term consequences of night lighting in urban ecosystems, through the impact of orb‐web spiders on insect populations.     

The Use of Automated Bioacoustic Recorders to Replace Human Wildlife Surveys: An Example Using Nightjars

To be able to monitor and protect endangered species, we need accurate information on their numbers and where they live. Survey methods using automated bioacoustic recorders offer significant promise, especially for species whose behaviour or ecology reduces their detectability during traditional surveys, such as the European nightjar. In this study we examined the utility of automated bioacoustic recorders and the associated classification software as a way to survey for wildlife, using the nightjar as an example. We compared traditional human surveys with results obtained from bioacoustic recorders. When we compared these two methods using the recordings made at the same time as the human surveys, we found that recorders were better at detecting nightjars. However, in practice fieldworkers are likely to deploy recorders for extended periods to make best use of them. Our comparison of this practical approach with human surveys revealed that recorders were significantly better at detecting nightjars than human surveyors: recorders detected nightjars during 19 of 22 survey periods, while surveyors detected nightjars on only six of these occasions. In addition, there was no correlation between the amount of vocalisation captured by the acoustic recorders and the abundance of nightjars as recorded by human surveyors. The data obtained from the recorders revealed that nightjars were most active just before dawn and just after dusk, and least active during the middle of the night. As a result, we found that recording at both dusk and dawn or only at dawn would give reasonably high levels of detection while significantly reducing recording time, preserving battery life. Our analyses suggest that automated bioacoustic recorders could increase the detection of other species, particularly those that are known to be difficult to detect using traditional survey methods. The accuracy of detection is especially important when the data are used to inform conservation.     

Torpor in dark times: patterns of heterothermy are associated with the lunar cycle in a nocturnal bird

Many studies have shown that endotherms become more heterothermic when the costs of thermoregulation are high and/or when limited energy availability constrains thermoregulatory capacity. However, the roles of many ecological variables, including constraints on foraging opportunities and/or success, remain largely unknown. To test the prediction that thermoregulatory patterns should be related to foraging opportunities in a heterothermic endotherm, we examined the relationship between the lunar cycle and heterothermy in Freckled Nightjars (Caprimulgus tristigma), which are visually orienting, nocturnal insectivores that are dependent on ambient light to forage. This model system provides an opportunity to assess whether variation in foraging opportunities influences the expression of heterothermy. The nightjars were active and foraged for insects when moonlight was available but became inactive and heterothermic in the absence of moonlight. Lunar illumination was a much stronger predictor of the magnitude of heterothermic responses than was air temperature (T(a)). Our data suggest that heterothermy was strongly related to variation in foraging opportunities associated with the lunar cycle, even though food abundance appeared to remain relatively high throughout the study period. Patterns of thermoregulation in this population of Freckled Nightjars provide novel insights into the environmental and ecological determinants of heterothermy, with the lunar cycle, and not T(a), being the strongest predictor of torpor use.     

Dietary Benefits of Twilight Foraging by the Insectivorous Bat Hipposideros speoris1

Although bats are nocturnal, many species emerge from roosts to forage during twilight, despite a presumed high risk of predation at this time. Here, we describe twilight foraging by a maternity colony of Schneider's leafnosed bat (Hipposideros speoris) in the dry zone of Sri Lanka and determine the dietary benefits of such behavior. Bats usually began foraging during dusk, sometimes before sunset, and also foraged during twilight in the morning. Mean use of available twilight by four radio‐tagged bats was 75 percent. Twilight foraging made up, on average, 47 percent of the total foraging time of these bats (range = 25–96%), although twilight consisted of only 12 percent of the available time between sunset and sunrise the next morning. Eight species of potential predators (7 birds and 1 mammal) were observed within a 1 km radius of the colony, of which 5 species are predicted to regularly capture bats. Bats took a wide diversity of prey (11 insect orders, including at least 27 families, and spiders) that ranged in wing length from 2.0 to 54.0 mm. Major orders in the diet were Coleoptera, Lepidoptera, and Diptera. Prey of secondary importance included Hemiptera, Hymenoptera, Isoptera, and Neuroptera. Bats captured large numbers of insects that were only available or had marked peaks in abundance during twilight. These groups included small, swarming insects (especially flies) that have peaks in flight activity at dusk and dawn, large diurnal species (especially dragonflies) that have crepuscular activity, and winged termites that emerge in swarms at dusk. Access to these insects was a clear benefit of twilight foraging.     

The Effect of Moonlight on Scopoli's Shearwater Calonectris diomedea Colony Attendance Patterns and Nocturnal Foraging: A Test of the Foraging Efficiency Hypothesis

Moonlight is known to affect the nocturnal behaviour and activity rhythms of many organisms. For instance, predators active at night may take advantage from increased visibility afforded by the moon, while prey might regulate their activity patterns to become less detectable. Many species of pelagic seabirds attend their colony only at night, in complete darkness, avoiding approaching their nest sites under moonlight. This behaviour has been most often interpreted as an antipredator adaptation (‘predation avoidance’ hypothesis). However, it may also reflect a lower foraging efficiency during moonlit nights (‘foraging efficiency’ hypothesis). Indeed, moonlight may reduce prey availability because preferred seabird prey is known to occur at higher depths in moonlit nights. Using high‐accuracy behavioural information from data loggers, we investigated the effect of moonlight on colony attendance and at‐sea nocturnal foraging in breeding Scopoli's shearwaters Calonectris diomedea. We found that birds departing for self‐feeding trips around the full moon performed longer trips than those departing around the new moon. On nights when the moon was present only partly, nest burrow entrances took place largely in the moonless portion of the night. Moreover, contrary to predictions from the ‘foraging efficiency’ hypothesis, nocturnal foraging activity increased according to moonlight intensity, suggesting that birds increased their foraging activity when prey became more detectable. This study strengthens the idea that colony attendance behaviour is strictly controlled by moonlight in shearwaters, which is possibly related to the perception of a predation risk.     

Foraging rhythm of great crested grebes Podiceps cristatus adjusted to diel variations in the vertical distribution of their prey Osmerus eperlanus in a shallow eutrophic lake in The Netherlands

Summary Great crested grebes Podiceps cristatus only foraged for an hour or two during dawn and again during dusk on Lake IJsselmeer in August-September. During this time of the year the adult birds are in wing moult and temporarily unable to fly. The food of grebes consisted almost exclusively of smelt Osmerus eperlanus, the most numberous pelagic fish. Simultaneous sonar registrations and trawl net fishing showed that smelt moved to the water surface during the twilight periods. During day and night they were concentrated near the bottom. We argue that grebes have the best foraging opportunities during twilight when much of their prey is near the surface, where light intensities allow the fish to be detected and captured. When the smelt are in the upper water layers the distance to the covered to get the prey (i.e. diving time and cost) is also least.     

A telemetry study on the diurnal distribution and activity of adult pikeperch, Sander lucioperca (L.), in a riverine environment

The diurnal movements and spatial distribution of adult pikeperch, Sander lucioperca, in the Elbe River, Czech Republic was observed using radio telemetry. The hypothesis that light intensity, within four different intervals (dawn, day, dusk, night), would determine the spatial distribution of pikeperch in a riverine environment were tested across a time span of 12 months. During the day, fish were located in deep water of the main channel, moving towards shallower waters during twilight and residing in the littoral zone, closest to the riverbanks, at night. Movement activity followed the behavioural pattern in a drainage canal with maximum at twilight and minimum at night. This suggests that nocturnal positions of adult pikeperch in the shallows were not associated with hunting but more likely with resting. Handling editor: J. Cambray     

Foraging strategies of Eriophora transmarina and Nephila plumipes (Araneae: Araneoidea): Nocturnal and diurnal orb-weaving spiders

Abstract The foraging behaviour, web characteristics and prey availability of two sympatric orb-weaving spiders, Nephila plumipes and Eriophora transmarina (Araneae: Araneoidea), are compared. The spiders are similarly sized but have different temporal foraging patterns. Nephila plumipes spins a relatively permanent web and captures most of its prey during the day. Eriophora transmarina only forages at night, spinning a new web every night and usually dismantling it at dawn. These different foraging activities are most likely to be responsible for the observed differences in the types and rates of prey capture: E. transmarina captured mostly Lepidoptera that were more abundant at night than during the day, while N. plumipes captured mostly Hymenoptera that were more abundant during the day than at night. While nocturnal E. transmarina have less time available for foraging than the diurnal N. plumipes, the former has a substantially higher nocturnal prey capture rate. We argue that the difference between the species in their prey capture rates are likely to be due to differences in the architecture of their webs.     

Lunar influences in the diet habits and selectivity of larval clingfish Gobiesox marmoratus

The influence of the lunar cycle on the feeding habits, feeding success and selectivity was tested in situ on larval stages of the clingfish Gobiesox marmoratus (Gobiesocidae). Gobiesox marmoratus larvae and their prey were collected in the water column during the lunar cycle in austral spring 2015 and 2016, in shallow waters (<30 m depth) of El Quisco, central Chile. Feeding incidence was high (80–100%) throughout the moon cycle. The diet was composed of 32 prey items, being gastropod larvae and invertebrate eggs the most important. Prey composition varied among lunar phases with the lowest prey richness during new moon. During 2016, G. marmoratus larvae ingested the lowest number, but the largest prey during new moon. In spring 2015, there were no significant changes in the abundance of each prey taxon in the water column among moon phases, but in spring 2016 there were significant differences in the abundance of cypris and copepod nauplii, particularly between full and new moon. Feeding selectivity index showed that gastropod larvae were positively selected throughout the lunar cycle during spring 2015. In spring 2016, invertebrate eggs were selectively ingested in full moon and third quarter, but at a new moon G. marmoratus larvae selected gastropod larvae. Mean temperature of the water column and its vertical gradient and nocturnal cloud cover influences the feeding success of larval G. marmoratus. Lunar illumination favoured only an increase of richness of prey items. Therefore, nocturnal cloud cover precludes a greater influence of lunar illumination in the larval trophic ecology of this crypto-benthic fish.     

Effects of light dynamics on coral spawning synchrony

Synchrony of spawning in many hermatypic corals, typically a few nights after the full moon, is putatively dependent on solar and lunar light cycles in conjunction with other possible cues such as tides and temperature. We analyze here the contributions of separate components of light dynamics, because the effects of twilight and lunar skylight on coral spawning synchrony have previously been conflated and the alternative hypothesis that these components have differential contributions as proximate cues has not been tested. Moonlight-dependent changes in spectra during twilight, rates of decreasing twilight intensities, and changes in lunar photoperiod were experimentally decoupled using programmed light-emitting diodes and compared for their separate effects on spawning synchrony in Acropora humilis. Effects on synchrony under the control of synthetic lunar cues were greatest in response to changes in lunar photoperiod; changes in light intensities and spectra had lesser influence. No significant differences among treatment responses were found at the circa-diel time scale. We conclude that spawning synchrony on a particular lunar night and specific time of night is a threshold response to differential periods of darkness after twilight that is primarily influenced by lunar photoperiod and secondarily by discrete optical components of early nocturnal illumination.     

Diel and lunar variation in diving behavior of rough-toothed dolphins (Steno bredanensis) off Kauaʻi,Hawaiʻi

Observational studies describe rough-toothed dolphins (Steno bredanensis) actively foraging during the day on epipelagic species. Using data from depth-transmitting satellite tags deployed on nine individuals off Kauaʻi, we investigated diving behavior and the effects of lunar phase and solar light levels on vertical movements. Overall, tagged rough-toothed dolphins primarily used near-surface waters, spending between 83.6% and 93.7% of their time in the top 30 m of the water column. When diving, grand mean, median, and maximum dive depths were 76.9 m, 67.5 m, and 399.5 m, although individuals were in water with depths from approximately 700–1,450 m. Dive rates varied by time of day, being lowest during the day and at dawn and highest at dusk and night. Dives were deepest (M = 133.7 m, SD = 52.6 m, median = 106.5 m) and longest (M = 4.0 min, SD = 0.4 min, median = 4.0 min) at dusk, suggesting dolphins were taking advantage of prey rising to the surface in response to reduced light levels. Lunar phase indirectly affected diving, with deeper and longer dives occurring with increasing illumination. The variations in dive behavior across solar and lunar cycles indicate diving patterns shift based on the distribution of prey.     

Laboratory video recordings and underwater visual observations combined to reveal activity rhythm of red-spotted grouper and banded wrasse, and their natural assemblages

The diel activity rhythm of red-spotted grouper Epinephelus akaara was studied both in captivity and in the wild. Behavior of solitary grouper (58 to 397?mm in total length) in a tank was video recorded using infrared illuminators under 11L/10D and two 1.5-h twilight transition periods, and was compared to that of banded wrasse Halichoeres poecilopterus, a typical diurnal fish. Underwater observations using SCUBA were also conducted in their natural habitat to reveal the behavioral activity together with a visual census of adjacent fish and crustacean assemblages. Red-spotted grouper showed a strong nocturnal activity in a tank regardless of body size as opposed to the strongly diurnal banded wrasse. Activity of groupers in natural waters was high at dawn and dusk, low at noon, and only a few individuals were observed at night. Visual census in the habitat revealed that fish abundance and species richness was highest at noon, lowest at night, and intermediate at dawn and dusk. The opposite trend was found in crustacean assemblages. Absence of groupers at night may reflect their nocturnal feeding migration away from the study area. Alternatively, the crepuscular activity of groupers in the wild is suggested to be an adaptation to feed on small fishes that shift between daytime activity and nighttime rest and/or on nocturnal crustaceans that show the opposite activity pattern.     

Rising rates of starch degradation during daytime and trehalose 6-phosphate optimize carbon availability

Many plants, including Arabidopsis (Arabidopsis thaliana), accumulate starch in the light and remobilize it to support maintenance and growth at night. Starch synthesis and degradation are usually viewed as temporally separate processes. Recently, we reported that starch is also degraded in the light. Degradation rates are generally low early in the day but rise with time. Here, we show that the rate of degradation in the light depends on time relative to dawn rather than dusk. We also show that degradation in the light is inhibited by trehalose 6-phosphate, a signal for sucrose availability. The observed responses of degradation in the light can be simulated by a skeletal model in which the rate of degradation is a function of starch content divided by time remaining until dawn. The fit is improved by extension to include feedback inhibition of starch degradation by trehalose 6-phosphate. We also investigate possible functions of simultaneous starch synthesis and degradation in the light, using empirically parameterized models and experimental approaches. The idea that this cycle buffers growth against falling rates of photosynthesis at twilight is supported by data showing that rates of protein and cell wall synthesis remain high during a simulated dusk twilight. Degradation of starch in the light may also counter over-accumulation of starch in long photoperiods and stabilize signaling around dusk. We conclude that starch degradation in the light is regulated by mechanisms similar to those that operate at night and is important for stabilizing carbon availability and signaling, thus optimizing growth in natural light conditions.

Starch degradation in the light is regulated by similar mechanisms to those operating at night, stabilizing carbon availability, and thereby optimizing growth in natural light conditions     

Temperature- and light-mediated predation by perch on vendace larvae

The perch foraging pattern in a Finnish lake (64) N) was crepuscular and they moved into littoral areas at dusk and returned to deeper waters by dawn. Predation on vendace larvae was guided by environmental cues, mainly temperature and light periodicity, which affected the density and foraging activity of perch in the littoral area. The availability and the vulnerability of different food categories were both responsible for the perch diet. The hypothesis that perch have a generally decisive role in the regulation of vendace larval abundance through predation at the time of high vendace larvae concentration in the littoral area was not supported. We emphasize the need to evaluate biological phenomena, such as predator–prey relationships, in the light of the relevant physical variables, and to do so within appropriate time and space scales, instead of relying on indirect estimates of the predator and prey population abundance and using their reciprocity as evidence of the interaction.     

DNA diet profiles with high‐resolution animal tracking data reveal levels of prey selection relative to habitat choice in a crepuscular insectivorous bird

Given the global decline of many invertebrate food resources, it is fundamental to understand the dietary requirements of insectivores. We give new insights into the functional relationship between the spatial habitat use, food availability, and diet of a crepuscular aerial insectivore, the European Nightjar (Caprimulgus europaeus) by relating spatial use data with high‐throughput sequencing (HTS) combined with DNA metabarcoding. Our study supports the predictions that nightjars collect a substantial part of their daily nourishment from foraging locations, sometimes at considerable distance from nesting sites. Lepidopterans comprise 65% of nightjars'' food source. Nightjars tend to select larger species of Lepidoptera (>19 mm) which suggests that nightjars optimize the efficiency of foraging trips by selecting the most energetically favorable—larger—prey items. We anticipate that our findings may shed additional light on the interactions between invertebrate communities and higher trophic levels, which is required to understand the repercussions of changing food resources on individual‐ and population‐level processes.     

Photoperiodism and the photic environment of the pitcher-plant mosquito,Wyeomyia smithii

Summary Wyeomyia smithii Coq. (Diptera: Culicidae) overwinters as a larva in a state of diapause which is initiated, maintained, and terminated by photoperiod. Both in the dawn and in the dusk, diapausing larvae are photoperiodically most sensitive to blue light (390–450 nm) with a shoulder in response in the bluegreen and green (480–540 nm) region of the spectrum. The saturation curves for response to blue light in the dusk has a steeper slope than for response to blue-green and green light in the dusk, suggesting two distinct pigments or pigment complexes underly photoperiodic response in W. smithii.The photic environment of W. smithii during twilight is rich in yellow-green light but sufficient light is available at 390–540 nm to trigger photoperiodic response early during morning civil twilight and to sustain response until late in evening civil twilight. Comparison of action spectra with spectra of available light indicates that the zenith angles of the sun which would result in 50% response are 95°48 and 94°52 in the dawn and dusk, respectively. Using these zenith angles to approximate daylength in nature provides a resonable prediction of development in the field.The flux density of photons necessary to elicit 50% development a 454 nm is about 9×10⁷ photons/cm² s in the dawn and 3×10⁸ photons/cm² s in the dusk. This high degree of sensitivity enables W. smithii to cue to the rapidly changing light intensity which occurs around the nautical-civil twilight transition. At the same time, the chromophore is not likely to be stimulated by the light of the full moon.     

Contrasting diel activity and feeding patterns of four species of carnivorous stoneflies

1. The hypothesis under test was that larvae of Dinocras cephalotes (Curtis), Perla bipunctata Pictet, Isoperla grammatica (Poda), and Perlodes microcephalus (Pictet) differed markedly in their diel activity and feeding patterns. Mature larvae collected about 1 month prior to adult emergence were used either for gut analyses or for observations of their activity and feeding patterns in three stream tanks with natural substrata and glass bottoms, so that activity could be observed above and below the substratum. A dull red light was used for observations in the dark. Food (larvae of Ephemeroptera, Simuliidae, and Chironomidae) was provided in excess. 2. Larvae for gut analyses were taken 1 h before dusk or dawn (n = 30 larvae per species for each day or night sample). The only species with food in the gut for the day samples was P. microcephalus. All species fed at night, the mean number of prey per larva being very similar for D. cephalotes, P. bipunctata, and I. grammatica but significantly higher for P. microcephalus. Most prey were insect larvae, especially Simuliidae and Chironomidae. 3. Diel activity patterns of single larvae differed interspecifically but not intraspecifically. Larvae of D. cephalotes and P. bipunctata were rarely active during the day; their activity increased at dusk and decreased at dawn, and was highest during the night; their success at prey capture was highest at dusk and dawn, with an ambush rather than a search strategy. Isoperla grammatica was rarely active during the day, most active at dusk and dawn when prey capture was highest, using a search strategy, and less active for the rest of the night. Perlodes microcephalus was active during the day, but only below the substratum, and very active from dusk to dawn with a high prey‐capture success, using a search strategy. These experiments provided support for the hypothesis under test. The discussion concludes that the results could also help to explain known differences in growth rate and the length of the life‐cycle in these four species.     

Nightjars may adjust breeding phenology to compensate for mismatches between moths and moonlight

Phenology match–mismatch usually refers to the extent of an organism's ability to match reproduction with peaks in food availability, but when mismatch occurs, it may indicate a response to another selective pressure. We assess the value of matching reproductive timing to multiple selective pressures for a migratory lunarphilic aerial insectivore bird, the whip‐poor‐will (Antrostomus vociferus). We hypothesize that a whip‐poor‐will's response to shifts in local phenology may be constrained by long annual migrations and a foraging mode that is dependent on both benign weather and the availability of moonlight. To test this, we monitored daily nest survival and overall reproductive success relative to food availability and moon phase in the northern part of whip‐poor‐will's breeding range. We found that moth abundance, and potentially temperature and moonlight, may all have a positive influence on daily chick survival rates and that the lowest chick survival rates for the period between hatching and fledging occurred when hatch was mismatched with both moths and moonlight. However, rather than breeding too late for peak moth abundance, the average first brood hatch date actually preceded the peak moth abundance and occurred during a period with slightly higher available moonlight than the period of peak food abundance. As a result, a low individual survival rate was partially compensated for by initiating more nesting attempts. This suggests that nightjars were able to adjust their breeding phenology in such a way that the costs of mismatch with food supply were at least partially balanced by a longer breeding season.