The effect of the lunar cycle on fecal cortisol metabolite levels and foraging ecology of nocturnally and diurnally active spiny mice

We studied stress hormones and foraging of nocturnal Acomys cahirinus and diurnal A. russatus in field populations as well as in two field enclosures populated by both species and two field enclosures with individuals of A. russatus alone. When alone, A. russatus individuals become also nocturnally active. We asked whether nocturnally active A. russatus will respond to moon phase and whether this response will be obtained also in diurnally active individuals. We studied giving-up densities (GUDs) in artificial foraging patches and fecal cortisol metabolite levels. Both species exhibited elevated fecal cortisol metabolite levels and foraged to higher GUDs in full moon nights; thus A. russatus retains physiological response and behavioral patterns that correlate with full moon conditions, as can be expected in nocturnal rodents, in spite of its diurnal activity. The endocrinological and behavioral response of this diurnal species to moon phase reflects its evolutionary heritage.     

The effect of rainfall on population dynamics in Sahara-Sahel rodents

In arid and semi-arid regions, rainfall is scarce, limiting primary productivity and animal reproduction. As long-term population monitoring is limited in remote arid areas, indirect and remote technicals are needed. We investigated if and how populations of rodents in Sahara-Sahel responded to past events of rainfall. Using short field surveys and remotely sensed rainfall data, spanning between years 2010 to 2015, we tested if rainfall prior to field surveys affected populations of Gerbillus rodents. Generalized additive analysis showed that amount of moonlight (the effect of number of days away from full moon) negatively correlated with number of trapped animals. When controlling for moonlight and geographic and temporal variation, rainfall up to 1 year prior to surveys positively and rainfall 2 years prior to surveys negatively correlated with number of trapped gerbils. We suggest that the effect of increased number of gerbils resulted from reproduction and population density increase after bursts of primary productivity. Negative correlation with rainfall could be related to increased predation or other ecological effects (e.g., resource collapse, pathogens spread) on prey population densities and activity levels. Our results suggest multiphase delayed effect of gerbils population response to rainfall implying interactive model of population regulation in rodent communities on Sahara-Sahel. Presented indirect method and results are readily applicable to population monitoring and management of remote and understudied areas.     

Influence of environmental factors on the numbers and activity of wintering Lapwings and Golden Plovers

Variations in the numbers and activity of Lapwings Vanellus vanellus and Golden Plovers Pluvialis apricaria using 322 ha of lowland farmland were examined in relation to temperature, rainfall, wind speed and moonphase. The study was undertaken over two winters (1987/88 and 1988/89), with maximum counts of 4000 Lapwings and 820 Golden Plovers. Patterns of occupancy of the study area and seasonal occurrence are described. Temperature and the phase of the lunar cycle seemed to be particularly important in affecting the distribution and activity of the birds. More Lapwings occurred in the study area on the colder and on the wetter days in several months, and also around the full moon period when feeding activity was particularly prevalent. A higher proportion of Lapwings tended to feed on colder days and around the new moon period, while daytime loafing increased around the full moon period. These results support the ideas that birds need to feed more in cold conditions, when energy demands are high, and that night feeding frequently occurs during the full moon period, although the reasons for this remain unclear.     

Non-additivity among multiple cues of predation risk: a behaviorally-driven trophic cascade between owls and songbirds

Non-lethal effects of predators on prey are initiated in the form of responses to direct and indirect cues of predation risk. Like their lethal equivalents, non-lethal effects may affect species further down the food web initiating a behaviorally-driven trophic cascade. I presented a direct cue of predation risk, owl vocalizations, to white-footed mice ( Peromyscus leucopus ) during either a new or full moon (indirect cue). Mice reduced their activity in space by nearly two-thirds in response to playbacks of owl vocalizations during a full moon. However, neither moonlight (full vs new) nor the presence/absence of owl calls had an effect on space use when each cue varied singly. Previous studies have demonstrated a tight correlation between spatial activity in mice as used in the current experiment and nest predation rates on ground-nesting birds. Because moonlight is a ubiquitous deterrent of activity in nocturnal rodents I used of long-term nesting records the veery ( Catharus fuscescens ) to test whether nest predation rates were correlated negatively with moonlight. For half the lunar cycle (∼full moon to new moon) predation rates decreased with moonlight as predicted. During the second half of the lunar cycle predation and moonlight did not correlate as expected, but this was likely due to the depletion of vulnerable nests after a period of in which predation rates were at their maximum near the full moon. These studies suggest that the non-lethal effects of predatory risk on mice (i.e. changes in space use) cascade to affect their prey. Through the mechanism of reduced space use by rodents, perceived predation risk has the potential to significantly and indirectly affect songbird nest predation rates.     

Responses of a top and a meso predator and their prey to moon phases

We compared movement patterns and rhythms of activity of a top predator, the Iberian lynx Lynx pardinus, a mesopredator, the red fox Vulpes vulpes, and their shared principal prey, the rabbit Oryctolagus cuniculus, in relation to moon phases. Because the three species are mostly nocturnal and crepuscular, we hypothesized that the shared prey would reduce its activity at most risky moon phases (i.e. during the brightest nights), but that fox, an intraguild prey of lynx, would avoid lynx activity peaks at the same time. Rabbits generally moved further from their core areas on darkest nights (i.e. new moon), using direct movements which minimize predation risk. Though rabbits responded to the increased predation risk by reducing their activity during the full moon, this response may require several days, and the moon effect we observed on the rabbits had, therefore, a temporal gap. Lynx activity patterns may be at least partially mirroring rabbit activity: around new moons, when rabbits moved furthest and were more active, lynxes reduced their travelling distances and their movements were concentrated in the core areas of their home ranges, which generally correspond to areas of high density of rabbits. Red foxes were more active during the darkest nights, when both the conditions for rabbit hunting were the best and lynxes moved less. On the one hand, foxes increased their activity when rabbits were further from their core areas and moved with more discrete displacements; on the other hand, fox activity in relation to the moon seemed to reduce dangerous encounters with its intraguild predator.     

Interaction effects between weather and space use on harvesting effort and patterns in red deer

Most cervid populations in Europe and North America are managed through selective harvesting, often with age‐ and sex‐specific quotas, with a large influence on the population growth rate. Less well understood is how prevailing weather affects harvesting selectivity and off‐take indirectly through changes in individual animal and hunter behavior. The behavior and movement patterns of hunters and their prey are expected to be influenced by weather conditions. Furthermore, habitat characteristics like habitat openness are also known to affect movement patterns and harvesting vulnerability, but how much such processes affect harvest composition has not been quantified. We use harvest data from red deer (Cervus elaphus) to investigate how weather and habitat characteristics affect behavioral decisions of red deer and their hunters throughout the hunting season. More specifically, we look at how sex and age class, temperature, precipitation, moon phase, and day of week affect the probability of being harvested on farmland (open habitat), hunter effort, and the overall harvest numbers. Moon phase and day of week were the strongest predictors of hunter effort and harvest numbers, with higher effort during full moon and weekends, and higher numbers during full moon. In general, the effect of fall weather conditions and habitat characteristics on harvest effort and numbers varied through the season. Yearlings showed the highest variation in the probability of being harvested on farmland through the season, but there was no effect of sex. Our study is among the first to highlight that weather may affect harvesting patterns and off‐take indirectly through animal and hunter behavior, but the interaction effects of weather and space use on hunter behavior are complicated, and seem less important than hunter preference and quotas in determining hunter selection and harvest off‐take. The consideration of hunter behavior is therefore key when forming management rules for sustainable harvesting.     

Temporal activity patterns of North China leopards and their prey in response to moonlight and habitat factors

The nocturnal activities of predators and prey are influenced by several factors, including physiological adaptations, habitat quality and, we suspect, corresponds to changes in brightness of moonlight according to moon phase. In this study, we used a dataset from 102 camera traps to explore which factors are related to the activity pattern of North China leopards (Panthera pardus japonensis) in Shanxi Tieqiaoshan Provincial Nature Reserve (TPNR), China. We found that nocturnal activities of leopards were irregular during four different lunar phases, and while not strictly lunar philic or lunar phobic, their temporal activity was highest during the brighter moon phases (especially the last quarter) and lower during the new moon phase. On the contrary, roe deer (Capreolus pygargus) exhibited lunar philic activity, while wild boar (Sus scrofa) and tolai hare (Lepus tolai) were evidently lunar phobic, with high and low temporal activity during the full moon, respectively. In terms of temporal overlap, there was positive overlap between leopards and their prey species, including roe deer and tolai hare, while leopard activity did not dip to the same low level of wild boar during the full moon phase. Human activities also more influenced the temporal activity of leopards and wild boar than other species investigated. Generally, our results suggested that besides moonlight risk index (MRI), cloud cover and season have diverse effects on leopard and prey nocturnal activity. Finally, distinct daytime and nighttime habitats were identified, with leopards, wild boar, and tolai hare all using lower elevations at night and higher elevations during the day, while leopards and roe deer were closer to secondary roads during the day than at night.     

Dodging the moon: The moon effect on activity allocation of prey in the presence of predators

Animals can adapt their activity patterns throughout the circadian cycle. Prey may use moonlight as a predation risk cue and allocate their activity to lower risk periods. Here, we assessed moon transit influence on the activity allocation of nocturnal mammalian prey, in the presence of a predator (pumas, Puma concolor), during different moon phases, through camera trapping in Central Amazon. Thirty camera traps were installed 2 km apart from each other in each of our three study sites. Prey record distributions were assessed across the moon cycle, and their daily activity patterns were described in each moon phase. The record distributions of pacas (Cuniculus paca) (N = 262) and armadillos (Dasypus sp.) (N = 244) were concentrated in darker nights, contrasting with red brocket deer (Mazama americana) (N = 123) and pumas (N = 31), whose records were evenly distributed through the moon cycle. Yet, every prey was found to avoid the brighter times of the night regardless of the moon phase. These findings suggest prey can shift the temporal distribution of their activities under different moon phases when predators are present, perhaps in response to predation risk variations.     

Singing in the moonlight: dawn song performance of a diurnal bird varies with lunar phase

It is well established that the lunar cycle can affect the behaviour of nocturnal animals, but its potential to have a similar influence on diurnal species has received less research attention. Here, we demonstrate that the dawn song of a cooperative songbird, the white-browed sparrow weaver (Plocepasser mahali), varies with moon phase. When the moon was above the horizon at dawn, males began singing on average 10 min earlier, if there was a full moon compared with a new moon, resulting in a 67% mean increase in performance period and greater total song output. The lack of a difference between full and new moon dawns when the moon was below the horizon suggests that the observed effects were driven by light intensity, rather than driven by other factors associated with moon phase. Effects of the lunar cycle on twilight signalling behaviour have implications for both pure and applied animal communication research.     

Moon and sun compasses in sandhoppers rely on two separate chronometric mechanisms

The relationship between the chronometric system of compensation for the apparent movement of the sun and that for the moon has been the subject of several, never proven, hypotheses. Our studies on sandhoppers have demonstrated that the chronometric mechanism of the moon compass is separate from that of the sun compass. They show (i) that a period of seven days in constant darkness has no influence on the capacity for orientation, either solar or lunar, and indicates the presence of one or more continuously operating timing mechanisms; (ii) that two different shifts in the light–dark phase have no effect on the chronometric mechanism of lunar orientation, but they do affect that of solar orientation; and (iii) that exposure to an artificial moon delayed by seven days with respect to the natural cycle causes the expected change in the mean direction of individuals tested under the natural moon, but not of those tested under the sun.     

Rainfall as a breeding stimulus and clutch size determinant in South African arid-zone birds

Rainfall is known to be a stimulus for initiating breeding in arid-zone birds, but its influence on clutch size is unknown for most southern African birds. The breeding activity of nine out of 11 species studied in the arid Nama Karoo biome of South Africa increased markedly after appreciable rainfall. Some resident insectivores did breed in the absence of rainfall, even under drought conditions, but also laid eggs within one week of small rain showers. Nomadic granivores required more substantial rainfall to stimulate what was usually a synchronized, population-wide breeding response. The timing and length of the breeding season was dependent on the integrated effect that rainfall and temperature had on the growing season of the vegetation. Over 50% of the 11 species studied showed evidence of an increase in clutch size following large rainfall events. Grey-backed Finchlark Eremopterix verticalis began laying larger clutches within one week of a rainfall stimulus, illustrating how rapid this response can be.     

Wheel-Running Activity Patterns of Five Species of Desert Rodents

In contrast to the extensive laboratory data on activity patterns in rodent species inhabiting temperate zones, much less is known about the activity patterns of desert rodents. In order to address this issue, we measured wheel-running activity patterns in males and females of five species of wild-trapped desert rodents (Dipodillus dasyurus, Gerbillus andersoni, Gerbillus pyramidum, Meriones shawi, and Acomys cahirinus) in long 'summer-like' and short, 'winter-like' day lengths. The specific goals of the present study were to characterize activity patterns in several desert rodent species in the laboratory and to determine if activity patterns are expressed in a seasonal or sexually dimorphic manner. Specifically, wheel-running was measured for 11 weeks in long days followed by 11 weeks in short days to test for photoperiodic entrainment as well as responsiveness to changes in the light-dark cycle. All animals exhibited rhythmic patterns of wheel-running with consistent onsets and offsets that had well-defined relations with the light-dark cycle. All individuals of G. andersoni showed nocturnal activity patterns. Most individuals of G. pyramidum had nocturnal activity patterns, but some individuals showed a short bout of activity at the beginning of the light period. Most individuals of D. dasyurus and M. shawi showed bimodal (i.e., nocturnal and diurnal) activity patterns, although some showed markedly nocturnal activity patterns. There was no sexual dimorphism in wheel running activity rhythms in any of the species examined. As expected, decreases in day length resulted in an overall increase in the duration of activity in all species. Collectively, these data provide an initial characterization of activity patterns within desert rodents in a controlled laboratory setting.     

Celestial moderation of tropical seabird behavior

Most animals, including birds, have cyclic life histories and numerous studies generally conducted on captive animals have shown that photoperiod is the main factor influencing this periodicity. Moon cycles can also affect periodic behavior of birds. Few studies have investigated the influence of these environmental cues in natural settings, and particularly in tropical areas where the change in photoperiod is slight and some bird species keep cyclic behaviors. Using miniaturized light sensors, we simultaneously investigated under natural conditions the influence of photoperiod and moon phases on the migration dates and at-sea activity of a tropical seabird species, the Barau's petrel, throughout its annual cycle. Firstly, we found that birds consistently started their pre- and post-breeding migrations at precise dates corresponding in both cases to a day-duration of 12.5 hours, suggesting a strong influence of the photoperiod in the regulation of migration behavior. We also found that mean population arrival dates to the colony changed from year to year and they were influenced by moon phases. Returns at their colonies occurred around the last full moon of the austral winter, suggesting that moon cycle is used by birds to synchronize their arrival. Secondly, variations of day-time activity were sinusoidal and correlated to seasonal changes of daylength. We thus hypothesize that the photoperiod could directly affect the behavior of the birds at sea. Night-time at-sea activity exhibited a clear cycle of 29.2 days, suggesting that nocturnal foraging was highly regulated by moon phase, particularly during the non-breeding season. To our knowledge, this is the first study to document a mixed regulation of the behavior of a wild bird by photoperiod and moon phases throughout its annual cycle.     

Studies on the migratory characin, Prochilodus platensis Holmberg 1889, (Pisces, Characoidei) in the River Pilcomayo, South America

General studies cover the habitat and life cycle of a migratory population of Prochilodus platensis . Specific studies of this species involved the elucidation and evaluation of abiotic effects on local upstream migrations and preliminary age and growth estimations. Upstream movements appeared to be affected by water temperature and moon phase. Most of the fish moved upstream when the temperature was between about 14 and 18°C at the particular monitoring point. Catches of ascending fish were highest during the full moon quarter, less during the waxing phase and least during the waning and new moon quarters. Length frequency and scale check data allowed approximate, indirect estimations to be made of the modal lengths attained in each of the first five age-groups.     

Ungulate: vehicle collision rates are associated with the phase of the moon

The phase of the moon can affect activity patterns of nocturnal animals, and may also affect visibility for motorists. However, surprisingly little is known about whether the risk of a wildlife-vehicle collision (WVC) is associated with lunar phase. We investigated the relationship between frequency of WVC at night and lunar phase for four large ungulate species that account for a high proportion of serious WVC along roads in agricultural and forested landscapes of two continents: wild boar Sus scrofa, roe deer Capreolus capreolus, and red deer Cervus elaphus in Castile and Leon, Spain, and white-tailed deer Odocoileus virginianus in New York State, USA. Three of the four species most frequently collided with vehicles at night during the full moon phase of the lunar cycle; this pattern was evident throughout the year but was stronger during some months. For roe deer, the species for which WVC was most closely associated with the lunar cycle, the frequency of WVC was 71.3% greater during the full than new moon period. Our results indicate that rates of ungulate WVC at night cycle on a period of a lunar month, which has implications for traffic safety planning and for motor vehicle collision emergency response preparation.     

Full moon and crime.

The incidence of crimes reported to three police stations in different towns (one rural, one urban, one industrial) was studied to see if it varied with the day of the lunar cycle. The period of the study covered 1978-82. The incidence of crimes committed on full moon days was much higher than on all other days, new moon days, and seventh days after the full moon and new moon. A small peak in the incidence of crimes was observed on new moon days, but this was not significant when compared with crimes committed on other days. The incidence of crimes on equinox and solstice days did not differ significantly from those on other days, suggesting that the sun probably does not influence the incidence of crime. The increased incidence of crimes on full moon days may be due to "human tidal waves" caused by the gravitational pull of the moon.     

Strategies of habitat use by mountain gorillas

Patterns of home range use by a mountain gorilla group are examined here in relation to variation in food abundance and quality, diet quality and rainfall, to interactions between different social units and to the distribution of hazards (poaching risk). Variation in habitat quality influenced both long-term area occupation densities and shorter-term movement parameters (day journey length and the distance moved between consecutive feeding sites). Observed home range use patterns increased the gorillas' foraging efficiency. Rainfall influenced the evenness of home range use slightly, but this was the only seasonal variability. Interactions with other gorillas and poaching risk had proximate effects superimposed on the fundamental influence of food distribution patterns. Data from a larger sample of groups show a positive influence of group size on day journey length, but this was apparent only at very small and relatively large group sizes. The absolute distance travelled per day tends to be short even for large groups. The data support arguments that the costs of social foraging are low for mountain gorillas.     

On the role of phylogeny in determining activity patterns of rodents

Evolutionary plasticity is limited, to a certain extent, by phylogenetic constraints. We asked whether the diel activity patterns of animals reflect their phylogenies by analyzing daily activity patterns in the order Rodentia. We carried out a literature survey of activity patterns of 700 species, placing each in an activity time category: diurnal, nocturnal, or active at both periods (a-rhythmic). The proportion of rodents active at these categories in the entire order, was compared to the activity patterns of species of different families for which we had data for over ten species each: Dipodidae, Echimyidae, Geomyidae, Heteromyidae, Muridae, and Sciuridae. Activity times of rodents from different habitat types were also compared to the ordinal activity time pattern. We also calculated the probability that two random species (from a particular subgroup: family, habitat, etc.) will be active in the same period of the day and compared it to this probability with species drawn from the entire order. Activity patterns at the family level were significantly different from the ordinal pattern, emphasizing the strong relationship between intra-family taxonomic affiliation and daily activity patterns. Large families (Muridae and Sciuridae) analyzed by subfamilies and tribes showed a similar but stronger pattern than that of the family level. Thus it is clear that phylogeny constrains the evolution of activity patterns in rodents, and may limit their ability to use the time niche axis for ecological separation. Rodents living in cold habitats differed significantly from the ordinal pattern, showing more diurnal and a-rhythmic activity patterns, possibly due to physiological constraints. Ground-dwelling rodents differed significantly, showing a high tendency towards a-rhythmic activity, perhaps reflecting their specialized habitat. Electronic supplementary material Electronic supplementary material is available for this article at and accessible for authorised users.     

Why animals respond to the full moon: Magnetic hypothesis

The geomagnetic field is typically about 50 μT (range 20-90 μT). Geomagnetic activity generally decreases by about 4% for the seven days leading up to a full moon, and increases by about 4% after the full moon, lasting for seven days. Animals can clearly detect the changes in magnetic field intensity that occur at full moon, as it has been shown that variations of just a few tens of nT are adequate to form a useful magnetic ‘map’. We think that moonlight increases the sensitivity of animals' magnetoreception because the radical pair model predicts that magnetoreception is light dependent. In fact, there have been some reports of changes in the sensitivity of magnetoreception with lunar phase. We propose a hypothesis that animals respond to the full moon because of changes in geomagnetic fields, and that the sensitivity of animals' magnetoreception increases at this time.