Sex-specific timing of mate searching and territory prospecting in the nightingale: nocturnal life of females

Formal models have shown that diel variation in female mate searching is likely to have profound influence on daily signalling routines of males. In studies on acoustic communication, the temporal patterns of the receivers'' signal evaluation should thus be taken into account when investigating the functions of signalling. In bird species in which diel patterns of signalling differ between males singing to defend a territory or to attract a mate, the diel patterns of mate and territory prospecting are suggested to depend on the sex of the prospector. We simulated newly arriving female nightingales (Luscinia megarhynchos) by translocating radio-tagged females to our study site. The mate-searching females prospected the area mostly at night, visiting several singing males. The timing of female prospecting corresponded to the period of the night when the singing activity of unpaired males was higher than that of paired males. In contrast to females, territory searching males have been shown to prospect territories almost exclusively during the dawn chorus. At dawn, both paired and unpaired males sang at high rates, suggesting that in contrast to nocturnal singing, dawn singing is important to announce territory occupancy to prospecting males. In the nightingale, the sex-specific timing of prospecting corresponded to the differential signalling routines of paired and unpaired males. The temporal patterns in the behaviour of signallers and receivers thus appear to be mutually adapted.     

Seasonal and diel movement patterns of brown bears in a population in southeastern Europe

Most animals concentrate their movement into certain hours of the day depending on drivers such as photoperiod, ambient temperature, inter‐ or intraspecific competition, and predation risk. The main activity periods of many mammal species, especially in human‐dominated landscapes, are commonly set at dusk, dawn, and during nighttime hours. Large carnivores, such as brown bears, often display great flexibility in diel movement patterns throughout their range, and even within populations, striking between individual differences in movement have been demonstrated. Here, we evaluated how seasonality and reproductive class affected diel movement patterns of brown bears of the Dinaric‐Pindos and Carpathian bear populations in Serbia. We analyzed the movement distances and general probability of movement of 13 brown bears (8 males and 5 females) equipped with GPS collars and monitored over 1–3 years. Our analyses revealed that movement distances and probability of bear movement differed between seasons (mating versus hyperphagia) and reproductive classes. Adult males, solitary females, and subadult males showed a crepuscular movement pattern. Compared with other reproductive classes, females with offspring were moving significantly less during crepuscular hours and during the night, particularly during the mating season, suggesting temporal niche partitioning among different reproductive classes. Adult males, solitary females, and in particular subadult males traveled greater hourly distances during the mating season in May‐June than the hyperphagia in July–October. Subadult males significantly decreased their movement from the mating season to hyperphagia, whereas females with offspring exhibited an opposite pattern with almost doubling their movement from the mating to hyperphagia season. Our results provide insights into how seasonality and reproductive class drive intrapopulation differences in movement distances and probability of movement in a recovering, to date little studied, brown bear population in southeastern Europe.     

Predator-dependent diel migration by <Emphasis Type="Italic">Halocaridina rubra</Emphasis> shrimp (Malacostraca: Atyidae) in Hawaiian anchialine pools

Diel migration is a common predator avoidance mechanism commonly found in temperate water bodies and increasingly in tropical systems. Previous research with only single day and night samples suggested that the endemic shrimp, Halocaridina rubra, may exhibit diel migration in Hawaiian anchialine pools to avoid predation by introduced mosquito fish, Gambusia affinis, and perhaps reverse migration to avoid the predatory invasive Tahitian prawn, Macrobrachium lar. To examine this phenomenon in greater detail, we conducted a diel study of H. rubra relative abundance and size at 2-h intervals in three anchialine pools that varied in predation regime on the Kona-Kohala Coast of Hawai‘i Island. We found two distinct patterns of diel migration. In two pools dominated by visually feeding G. affinis, the abundance of H. rubra present on the pool bottom or swimming in the water column was very low during the day, increased markedly at sunset and remained high until dawn. In contrast, in a pool dominated by the nocturnal predator M. lar, H. rubra density was significantly lower during the night than during the day (i.e., a pattern opposite to that of shrimp in pools containing fish). In addition, we observed that the mean body size of the shrimp populations varied among pools depending upon predator type and abundance, but did not vary between day and night in any pools. Our results are consistent with the hypothesis that H. rubra diel migratory behavior and size distributions are influenced by predation regime and suggest that diel migration may be a flexible strategy for predator avoidance in tropical pools where it may be a significant adaptive response of endemic species to introduced predators.     

Rufous‐capped Warblers Basileuterus rufifrons show seasonal,temporal and annual variation in song use

In the majority of songbird species, males have repertoires of multiple song types used for mate attraction and territory defence. The wood‐warblers (family Parulidae) are a diverse family of songbirds in which males of many migratory species use different song types or patterns of song delivery (known as ‘singing modes’) depending on context. The vocal behaviour of most tropical resident warblers remains undescribed, although these species differ ecologically and behaviourally from migratory species, and may therefore differ in their vocal behaviour. We test whether male Rufous‐capped Warblers Basileuterus rufifrons use distinct singing modes by examining song structure and context‐dependent variation in their songs. We recorded multiple song bouts from 50 male Warblers in a Costa Rican population over 3 years to describe seasonal, diel and annual variation in song structure and vocal behaviour. We found that Rufous‐capped Warbler songs are complex, with many syllable types shared both within and between males’ repertoires. Males varied their song output depending on context: they sang long songs at a high rate at dawn and during the breeding season, and shortened songs in the presence of a vocalizing female mate. Unlike many migratory species, Rufous‐capped Warblers do not appear to have different singing modes; they did not change the song variants used or the pattern of song delivery according to time of day, season or female vocal activity. Our research provides the first detailed vocal analysis of any Basileuterus warbler species, and enhances our understanding of the evolution of repertoire specialization in tropical resident songbirds.     

Diel patterns of locomotor activity in Cydia pomonella: age and sex related differences and effects of insect hormone mimics

Abstract. The diel pattern of locomotor activity in the codling moth Cydia pomonella L. (Lepidoptera: Tortricidae) was investigated in the laboratory, using a computer-based infra-red actograph. The level of locomotor activity varied with sex and age. On the day of emergence, females were more active than males, but on days 2, 4, 6 and 12 males were approximately twice as active as females. Males reached their highest activity on day 4 and females on day 12. Both sexes were most active at dusk. Their activity patterns were fitted to a set of polynomial regression models. Treatment with 400 p.p.m. of Juvenile Hormone mimic, fenoxycarb, stimulated locomotor activity and provoked a marked activity peak at dawn in both virgin and mated females. A similar treatment with the ecdysone agonist tebufenozide showed a neutral effect on the locomotor activity of females. Possible implications of these findings are discussed in relation to the physiology of the moth and to its dispersal behaviour.     

Female-biased incubation and strong diel sex-roles in the Two-banded Plover Charadrius falklandicus

The relative contributions of males and females to incubation, and the diel schedules by which incubation is shared, are important breeding system traits. We used infra-red sensitive cameras to record incubation patterns at 13 nests of the Two-banded Plover Charadrius falklandicus in the Falkland Islands during both day and night. Because predation risk can affect incubation behaviour, we also recorded the diel pattern of nest predation in the wider study population. We found high nest attendance, female-biased incubation, and strong diel sex-roles, with females incubating during the day and males at night. We also found that incubation intermissions tended to be short but frequent, and were correlated strongly with the diel pattern of nest predations which occurred exclusively in the daylight hours (probably due to the absence of terrestrial mammals from the study site). Our results suggest that sex-roles are unusually strict in the Two-banded Plover, and that these strict sex-roles lead to inequality in incubation sharing and the level of exposure to sources of energetic cost such as disturbance by nest predators.     

Age‐ and sex‐related differences in the spatial ecology of a dichromatic tropical python (Morelia viridis)

Abstract: Despite outnumbering their temperate counterparts, tropical snake species have been poorly studied. Yet, the few tropical species that have been studied show a variety of behavioural traits beyond those described in temperate species. Here we reveal both age and sexual differences in the movements of tropical green pythons (Morelia viridis: Pythonidae). We radio‐tracked 27 individuals (17 females and 10 males) for up to 18 months, locating individuals during both the day and night. The home range size for adult females (mean ± SE of 6.21 ± 1.85 ha) was correlated with snout–vent length. Neither adult males nor juveniles had a stable home range. Adult females had stable home ranges that overlapped considerably with those of other females and yellow individuals. Multiple radio‐tracked adult males passed through the territory of radio‐tracked adult females during the study. Females of all sizes were more likely to change position each day than males. There were no differences between the sexes or size categories in the distances moved in most months, although the variation in movement distances was higher in the dry season than during the wet season. In the wet season (January–March) movement distances increased and these were size‐ and sex‐related. This increased activity may be associated with mate searching. Males of M. viridis may maximize their rate of encountering mature females by roaming rather than maintaining a home range. Juvenile green pythons moved distances equal to adult snakes in most months despite their comparatively small size.     

Stomach temperature telemetry reveals temporal patterns of foraging success in a free-ranging marine mammal

1. We studied feeding frequency in free-ranging grey seals using stomach temperature telemetry to test if previously reported sex differences in the diving, movement and diet were reflected in the temporal pattern of foraging success. 2. Data were retrieved from 21 of 32 grey seals from 1999 to 2001, totalling 343 days and 555 feeding events, with individual record length varying from 2 to 40 days (mean: 16.33 +/- 2.67 days/seal). 3. Seals fed on 57.8 +/- 6.46% of days sampled and had an average of 1.7 +/- 0.26 meals per day, but individual variability was apparent in the temporal distribution of feeding as evidenced by high coefficients of variation (coefficient of variation = 69.0%). 4. Bout analysis of non-feeding intervals of six grey seals suggests that feeding intervals of individuals were varied and probably reflect differences in prey availability. Grey seals tended to have many single feeding events with long periods separating each event, as would be expected for a large carnivore with a batch-reactor digestive system. 5. We found significant sex differences in the temporal distribution of feeding. The number of feeding events per day was greater in males (2.2 +/- 0.4 vs. 1.0 +/- 0.2), as was time associated with feeding per day (56.6 +/- 5.8 min vs. 43.9 +/- 9.4 min). 6. The number of feeding events varied with time of day with the least number occurring during dawn. Feeding event size differed significantly by time of day, with greater meal sizes during the dawn and the smallest meals during the night. 7. The length of time between meals increased with the size of the previous meal, and was significantly less in males (541.4 +/- 63.5 min) than in females (1092.6 +/- 169.9 min). 8. These results provide new insight into the basis of sex differences in diving and diet in this large size-dimorphic marine predator.     

Detecting diel patterns in the songs of Chipping Sparrows using citizen-science data

Previous studies have revealed diel patterns in the songs of Chipping Sparrows (Spizella passerina), with songs shorter in duration before dawn than after. However, the extent to which this phenomenon generalizes to the full geographic range of these sparrows is unclear, as is the question of whether citizen-science data can be used to detect diel patterns in song. We analyzed all available songs of Chipping Sparrows from the Macaulay Library and xeno-canto databases and compared the distributions of song features of recordings made at different times of day. We show that, across their entire geographic range in North and Central America, Chipping Sparrows sing shorter songs before sunrise (dawn song) than after sunrise (day song). Furthermore, we show that Chipping Sparrows shorten their songs by singing fewer syllables, not by singing faster: the number of syllables per song accounts for the observed difference in duration, not the syllable nor the intersyllable duration. Our results demonstrate that recordings from public repositories can be used to determine whether daily song patterns exist in species even in the absence of prior fieldwork, and we further propose that citizen-science recordings can be used to inform cross-species hypotheses and facilitate future studies to determine whether diel patterns in song are associated with differences in social behavior.     

The reproductive tactics and activity patterns of solitary carnivores: the Iriomote cat

Felids are generally considered to be crepuscular and nocturnal in their activity, but few studies have attempted to analyze the variability of their activity patterns. We studied the daily activity of the Iriomote cat Prionailurus iriomotensis by radio-tracking on Iriomote Island, Japan. The general activity patterns of Iriomote cats showed slightly prevailing activity during dark periods of the day with particular peaks at dawn and dusk or during the early hours of the night. However, these patterns were clearly dependent on the sex and reproductive status of the cat. Peaks of cats’ activity coincided with those of their main prey. On average, the cats were active for thirteen hours per day. During the mating season, the rhythm of activity in males followed that of breeding females, but not that of non-breeding ones. Males exhibited 11% higher total daily activity and longer active bouts during the mating period than in the remainder of the year. Breeding females had additional mid-day activity peak during the nursing period, but their total time of activity per day was 16% lower than in the period of kittens’ independence. Their active bouts were shorter and more frequent during nursing than at other times. These results suggest that lactating females perform frequent movements to and from the den site to care for kittens. During the non-nursing period, females increased their activity, possibly in response to lowered prey abundance and the need of intensive foraging to recover after lactation. Seasonal and sexual variation of activity patterns in the Iriomote cats confirmed the existence of different reproductive strategies of males and females of these solitary carnivores.     

Contrasting diel activity and feeding patterns of four instars of Rhyacophila dorsalis (Trichoptera)

1. Ontogenetic shifts in prey choice and predator behaviour can affect food‐web structure. Therefore, it is important to establish if the diet and feeding activity differ between life‐stages of the same species. This hypothesis was tested for second, third, fourth and fifth larval instars of Rhyacophila dorsalis by comparing their diel activity and feeding patterns. Second to fifth instars collected from two streams were used either for gut analyses or for observations of their activity and feeding patterns in three stream tanks. Food was provided in excess; being organisms living in bryophytes on top of a large stone in each tank, augmented by different‐sized larvae of Ephemeroptera, Simuliidae and Chironomidae. As few first instars for gut analyses were found in the field, the diet of first instars reared in the laboratory was also studied. 2. Larvae for gut analyses were taken 1 h before dusk or dawn (n = 50 larvae per instar for each day or night sample). First and second instars fed on the smaller food items with no significant day‐night differences in diet. Gut contents indicated a progressive trend from feeding chiefly at night in third instars to almost exclusively at night in fifth instars. Fourth and fifth instars fed on the larger food items, whilst the diet of the third instar larvae overlapped with that of both the earlier and later instars. 3. Diel activity patterns of single larvae differed between instars but not within each instar (n = 20 larvae per instar). Second instars were active throughout the 24 h, with peaks at dusk, around midnight, dawn and around midday. A similar pattern was shown by third instars but the peak of activity at midday was less than the other three peaks. Prey were captured only during these peaks for both instars. Fourth and fifth instars were most active, and fed only, at night. They used an ambush strategy to capture more active prey at dusk and dawn (e.g. Baetis, Gammarus), and a searching strategy to capture more sedentary prey during the night (e.g. chironomids, simuliids). These experiments provided support for the hypothesis under test. If competition and/or interference occur between instars, then it could be reduced between earlier and later instars because of differences in their diet and diel pattern of feeding activity.     

Diel activity pattern and predation rate of the generalist predator Dicyphus hesperus

Abstract This study examined the diel activity pattern and the effect of diel activity pattern on predation rate and prey finding of Dicyphus hesperus Knight (Heteroptera: Miridae). To determine the diel activity pattern of D. hesperus, starved females were placed on tomato leaflets Lycopersicon esculentum Mill. (Solanaceae) under zero, low, or high light intensities at 02:00, 08:00, and 14:00 h, respectively, and the amount of time spent walking or resting during a 30‐min interval was recorded. Predation rates of D. hesperus females on Ephestia kuehniella Zeller (Lepidoptera: Pyralidae) egg patches were determined under either a L16:D8 (long day) or L8:D16 (short day) diel period. Egg patches were removed from D. hesperus females after either 8 or 16 h of dark or 16 or 8 h of light, and the number of eggs consumed was counted. Dicyphus hesperus females spent more time searching for prey at night than during the day. Females ate eggs at a higher rate during the night than during the day. Overall, D. hesperus females had higher predation rates when reared under a long day diel cycle compared with females reared under a short day diel cycle. More females reared at the L16:D8 diel cycle found the egg patch during the night than during the day. There was no difference in egg patch finding between night and day for females reared at L8:D16. Overall, L16:D8 reared females found more egg patches than females reared at L8:D16. Therefore, D. hesperus females are more active and find and consume prey at a higher rate at night than day.     

Activity Pattern of Polecats Mustela putorius L. in Relation to Food Habits and Prey Activity

The activity pattern of polecats Mustela putorius L. was studied by radiotracking five males and four females in the wetlands of western France from Jul. 1988 to Mar. 1992. Polecats were nocturnal throughout the year with no differences between males and females. The diel activity remained moderate (31 %). Seasonal food habits were characterized by the exploitation of anurans in spring, while rodents were preyed upon from summer to winter. Seasonal changes in the activity rhythm of polecats appeared synchronized with the activity of the main prey. It is suggested that the activity patterns exhibited in different periods of the year are a result of changes in feeding tactics.     

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.     

Thermal biology, torpor use and activity patterns of a small diurnal marsupial from a tropical desert: sexual differences

Many small desert dasyurids employ torpor almost daily during winter, because cold nights and low food availability impose high energetic costs. However, in Western Australia the arid zone extends into tropical, coastal regions, where winter temperature conditions are far less severe. We studied the thermal biology and activity patterns of free-ranging kaluta (~27 g), a dasyurid restricted to these tropical spinifex deserts, during the Austral winter (June–July) and in addition quantified activity patterns in captivity. Unlike most dasyurids, wild and captive kalutas were almost exclusively diurnal and retreated into underground burrows during the night. Despite being active during the warmer part of the day, kalutas entered torpor daily. However, torpor patterns differed remarkably between males and females. While females spent most of the night torpid at body temperatures (T _b) as low as 21°C, close to soil temperature, males entered multiple short and shallow bouts (T _b > 25°C) during the night. Males also maintained higher T _bs during the early morning when active, occupied larger home ranges and covered greater distances while foraging than females. Hence, males appear to expend more energy than the similar-sized females both while foraging and during the rest phase. We propose that physiological as well as behavioural preparations for the September mating season that culminate in a complete male die-off might already impose energetic costs on males during winter.     

Habitat selection by the swamp wallaby (Wallabia bicolor) in relation to diel period,food and shelter

Patterns of resource selection by animals may be influenced by sex, and often change over a 24‐h period. We used a dry sclerophyll landscape managed for commercial timber production to investigate the effects of sex and diel period on habitat selection by the swamp wallaby (Wallabia bicolor). We predicted that selection would be (i) affected by both sex and diel period; and (ii) positively related to lateral cover during the day, but to food resources at night. Non‐metric multidimentional scaling indicated that some of the available habitats differed markedly with respect to visibility (an indicator of lateral cover), fern cover, forb cover, wallaby density and a forage quality index, providing the basis for non‐random habitat selection. At the landscape scale, wallabies showed strong selection for 5‐year‐old regenerating sites, selected against 10‐year‐old regenerating sites and unharvested forest, and avoided recently harvested (3–10 months post‐harvest) sites completely. At the scale of individual home ranges, a pooled male and female sample demonstrated selection for unharvested forest over recently harvested sites during both diurnal and nocturnal periods. A separate analysis showed that both sex and diel period influenced the selection of 5‐ and 10‐year‐old sites and the surrounding unharvested forest. Using a novel approach, we demonstrated that diurnal habitat selection by both sexes was negatively correlated with visibility, representing stronger selection for areas with more lateral cover. Nocturnal selection by females was positively correlated with values of a forage quality index, but this was not the case for males. We hypothesise that the observed patterns of selection were driven by the need to find food and avoid predators, but were also affected by the different reproductive strategies of males and females. Our results demonstrate the importance of incorporating factors such as sex and diel period into analyses of habitat selection.     

A 'polarisation sun-dial' dictates the optimal time of day for dispersal by flying aquatic insects

1. Daily changes in the flight activity of aquatic insects have been investigated in only a few water beetles and bugs. The diel flight periodicity of aquatic insects and the environmental factors governing it are poorly understood. 2. We found that primary aquatic insects belonging to 99 taxa (78 Coleoptera, 21 Heteroptera) fly predominantly in mid‐morning, and/or around noon and/or at nightfall. There appears to be at least four different types of diurnal flight activity rhythm in aquatic insects, characterised by peak(s): (i) in mid‐morning; (ii) in the evening; (iii) both in mid‐morning and the evening; (iv) around noon and again in the evening. These activity maxima are quite general and cannot be explained exclusively by daily fluctuations of air temperature, humidity, wind speed and risks of predation, which are all somewhat stochastic. 3. We found experimental evidence that the proportion (%) P(θ) of reflecting surfaces detectable polarotactically as ‘water’ is always maximal at the lowest (dawn and dusk) and highest (noon) angles of solar elevation (θ) for dark reflectors while P(θ) is maximal at dawn and dusk (low solar elevations) for bright reflectors under clear or partly cloudy skies. 4. From the temporal coincidence between peaks in the diel flight activity of primary aquatic insects and the polarotactic detectability P(θ) of water surfaces we conclude that the optimal times of day for aquatic insects to disperse are the periods of low and high solar elevations θ. The θ‐dependent reflection–polarisation patterns, combined with an appropriate air temperature, clearly explain why polarotactic aquatic insects disperse to new habitats in mid‐morning, and/or around noon and/or at dusk. We call this phenomenon the ‘polarisation sun‐dial’ of dispersing aquatic insects.     

Does sex matter? Gender‐specific responses to forest fragmentation in Neotropical bats

Understanding the consequences of habitat modification on wildlife communities is central to the development of conservation strategies. However, albeit male and female individuals of numerous species are known to exhibit differences in habitat use, sex‐specific responses to habitat modification remain little explored. Here, we used a landscape‐scale fragmentation experiment to assess, separately for males and females, the effects of fragmentation on the abundance of Carollia perspicillata and Rhinophylla pumilio, two widespread Neotropical frugivorous bats. We predicted that sex‐specific responses would arise from higher energetic requirements from pregnancy and lactation in females. Analyses were conducted independently for each season, and we further investigated the joint responses to local and landscape‐scale metrics of habitat quality, composition, and configuration. Although males and females responded similarly to a fragmentation gradient composed by continuous forest, fragment interiors, edges, and matrix habitats, we found marked differences between sexes in habitat use for at least one of the seasons. Whereas the sex ratio varied little in continuous forest and fragment interiors, females were found to be more abundant than males in edge and matrix habitats. This difference was more prominent in the dry season, the reproductive season of both species. For both species, abundance responses to local‐ and landscape‐scale predictors differed between sexes and again, differences were more pronounced in the dry season. The results suggest considerable sex‐mediated responses to forest disruption and degradation in tropical bats and complement our understanding of the impacts of fragmentation on tropical forest vertebrate communities.     

Spatial structure of the Caribbean lobster (Metanephrops binghami) in the Colombian Caribbean Sea

Crustaceans of the genus Metanephrops are of great commercial value in some tropical and subtropical regions. With the potential development of a new deep lobster fishery in the Colombian Caribbean Sea, the objective of this work is to describe for first time the patterns of spatial and bathymetric distribution, and diel migratory periodicity of the Caribbean lobster (M. binghami). Data were collected by trawling in depths between 200 and 550 m (100 m strata intervals) in the Colombian Caribbean Sea. Higher biomass and size of these crustaceans were found between 250 and 350 m, with a maximum at about 300 m. The study offers diel patterns of M. binghami, which suggests nocturnal activity and burrowing during daylight hours.