Effects of light and prey availability on nocturnal, lunar and seasonal activity of tropical nightjars

Nightjars and their allies represent the only major group of visually hunting aerial insectivores with a crepuscular and/or nocturnal lifestyle. Our purpose was to examine how both light regime and prey abundance in the tropics, where periods of twilight are extremely short, but nightjar diversity is high, affect activity across different temporal scales. We studied two nightjar species in West African bush savannah, standard‐winged nightjars Macrodipteryx longipennis Shaw and long‐tailed nightjars Caprimulgus climacurus Vieillot. We measured biomass of potential prey available using a vehicle mounted trap and found that it was highest at dusk and significantly lower at dawn and during the night. Based on direct observations, both nightjars exhibit the most intense foraging behaviour at dusk, less intense foraging at dawn and least at night, as predicted by both prey abundance and conditions for visual prey detection. Nocturnal foraging was positively correlated with lunar light levels and ceased below about 0.03 mW m^?2. Over the course of a lunar cycle, nocturnal light availability varied markedly, while prey abundance remained constant at dusk and at night was slightly higher at full moon. Both species increased twilight foraging activity during new moon periods, compensating for the shorter nocturnal foraging window at that time. Seasonally, the pattern of nocturnal light availability was similar throughout the year, while prey availability peaked shortly after onset of the wet season and then slowly decreased over the following four months. The courtship and breeding phenology of both species was timed to coincide with the peak in aerial insect abundance, suggesting that prey availability rather than direct abiotic factors act as constraints, at least at the seasonal level. Our findings illustrate the peculiar constraints on visually orienting aerial nocturnal insectivores in general and tropical nightjars in particular and highlight the resulting nocturnal, lunar and seasonal allocation of activities.     

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.     

The food of the Afrotropical nightjars

Jackson, H.D. 2000. The food of the Afrotrapical nightjars. Ostrich 71(3&4): 408-415. A synthesis of 722 published and unpublished records of nightjar stomach contents from Afrotropical specimens was undertaken. Coleoptera were found in 84.6% of the stomachs, often in substantial numbers, beetles providing these birds with their staple diet. Lepidoptera occurred in 34.9% of the stomachs and were often present in large numbers; moths are an extremely important part of the nightjar diet in the Afrotropics, especially during winter. Orthoptera, Hemiptera and Dictyoptera, although present in 24.7, 16.3 and 11.5% of the stomachs, respectively, usually occurred in rather small numbers. Hymenoptera and lsoptera were often present in great numbers, yet were found in only 10.4 and 10.1% of the stomachs, respectively, so a limited number of individual nightjars had found emerging swarms of alate ants or termites in season. Neuroptera, Diptera, Odonata and Dermaptera were found in a few stomachs (2.4, 1.2, 0.6 and 0.3%, respectively) but these insects clearly play a very small part in the diet of nightjars in the Afrotropics. Similarities and differences in diet between some closely related nightjar species are discussed. Grit was found in 16 stomachs only, so deliberate ingestion of stones to aid digestion is regarded as extremely unlikely among nightjars in the Afrotropics. The contents of a full stomach may account for as much as 20-25% of the body mass of a nightjar.     

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     

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.     

Vocal individuality of large-tailed nightjar (Caprimulgus macrurus) in Peninsular Malaysia

All four nightjar species resident in Peninsular Malaysia are vocally distinct, yet they remain little studied. Conventional field methods based on visual cues to study diurnal species may be impractical for nightjars. Alternatively, aural survey can potentially be applied on nightjars provided that individuality in their vocalisations can be proven. Our study aimed to determine the vocal individuality of the common, large-tailed nightjar (Caprimulgus macrurus) in oil palm smallholdings and an isolated forest patch located in Selangor, Peninsular Malaysia. From the call recordings obtained from 22 individuals, results of the Kruskal-Wallis analysis revealed significant differences in all the nine vocal parameters (call length, interquartile range bandwidth, low, high, average, centre and peak frequencies as well as first and third quartile frequencies) measured among individual nightjars (p < 0.001) regardless of study sites. Discriminant Function Analysis showed that more than 94.5% of original grouped cases were correctly classified. This implied that the majority of vocalizations can be assigned to individual birds based on the parameters measured. This study demonstrated the occurrence of vocal individuality in the large-tailed nightjar and such a finding pertaining to distinct vocalisations at the individual level will compensate for the limited access to visual cues in field surveys, as with the case for all nocturnal birds.     

What Drives Wolves: Fear or Hunger? Humans,Diet, Climate and Wolf Activity Patterns

Activity patterns of animals depend on environmental and intrinsic factors. Studies undertaken across the current wolf ( Canis lupus ) range suggested a number of variables that may be correlated with activity patterns of wolves. These factors vary locally and there has been no attempt so far at defining those that ubiquitously impact wolf behaviour. I compared 11 studies (from Alaska to Israel) to assess the influence of (1) public road density, (2) human population density, (3) human-caused mortality, (4) proportion of domestic animals in wolf diet, (5) proportion of forest, (6) latitude and (7) mean annual temperature on nocturnal wolf activity and movements. Nocturnal activity was mainly correlated to the proportion of domestic animals in the diet and the density of public roads, whereas nocturnal movements were mainly correlated to latitude. The importance of latitude indicates that sun periodicity might represent an important signal (`Zeitgeber') for circadian rhythms in wolves. Environmental constraints such as high temperatures during the day and a higher hunting success in crepuscular periods probably limit the ability of wolves to avoid humans by nocturnal behaviour. I therefore suggest that in regions where wolves hunt wild prey, they experience a trade-off between predation risk by humans and increased hunting success during twilight hours.     

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     

Nightjar diversity and microhabitat use in Nechisar National Park,Ethiopia

Knowledge of the distribution and ecology of East African nightjars is, to a large extent, unknown. We collected ecological information on the diversity and microhabitat use of nightjars in Nechisar National Park in January 2015 by executing field captures and observations. We also attempted to find a live specimen of the Nechisar Nightjar Caprimulgus solala. During the course of 18 nights we observed five nightjar species, captured 49 individuals and observed that nightjars were closely associated with protective landscape elements. In spite of the intensity of our survey, we were not able to find a living specimen of the Nechisar Nightjar, which could indicate the species is either migratory or extinct.     

Little evidence of a road‐effect zone for nocturnal,flying insects

Roads and traffic may be contributing to global declines of insect populations. The ecological effects of roads often extend far into the surrounding habitat, over a distance known as the road‐effect zone. The quality of habitat in the road‐effect zone is generally degraded (e.g., due to edge effects, noise, light, and chemical pollution) and can be reflected in species presence, abundance, or demographic parameters. Road‐effect zones have been quantified for some vertebrate species but are yet to be quantified for insects. Investigating the road‐effect zone for insects will provide a better understanding of how roads impact ecosystems, which is particularly important given the role insects play as pollinators, predators, and prey for other species. We quantified the road‐effect zone for nocturnal flying insects along three major freeways in agricultural landscapes in southeast Australia. We collected insects using light traps at six points along 2‐km transects perpendicular to each highway (n = 17). We sorted the samples into order, and dried and weighed each order to obtain a measure of dry biomass. Using regression models within a Bayesian framework of inference, we estimated the change in biomass of each order with distance from the road, while accounting for environmental variables such as temperature, moon phase, and vegetation structure. The biomass of nine of the ten orders sampled did not change with distance from the freeway. Orthoptera (i.e., grasshoppers and crickets) was the only order whose biomass increased with distance from the freeway. From our findings, we suggest that the impacts of roads on insects are unlikely extending into the surrounding landscape over a distance of 2 km. Therefore, if there are impacts of roads on insects, these are more likely to be concentrated at the road itself, or on finer taxonomic scales such as family or genus level.     

Observations of the flight behaviour of the army worm moth, Spodoptera exempta, at an emergence site using radar and infra-red optical techniques

ABSTRACT.

• 1 Studies were made in Kenya of the flight behaviour of African armyworm moths which had emerged from areas previously infested with‘gregarious’caterpillars. The use of radar and an infra-red optical detector permitted quantitative, direct observations of the flying moths, over all of their flight altitudes, without disturbing their behaviour.
• 2 Almost all of the successfully emergent moths climbed to altitudes of several hundred metres above ground level and migrated from the emergence sites. Their migratory flights sometimes started on the night of emergence, but on other occasions the moths remained roosting in trees until dawn, then engaged in short dispersal flights, concealed themselves during the day. and commenced migration in mass flights at dusk the following night.
• 3 The onset of these‘dusk flights’occurred when the irradiance level fell on average to 2.7 × 10^?5Wm^?2 nm^?1 in the 450–800 nm range (in the photometric units appropriate for human vision this corresponds approximately to 2 lux). The‘dawn flights’began with the first sustained increase in irradiance at dawn, and terminated at the end of dawn twilight.
• 4 Migratory flight in the strong, easterly winds which usually occurred during the first half of the night resulted in rapid, down-wind displacement to the west. Observations of groups of flying moths passing successively over two radars demonstrated that these migrations could cover at least 20 km.
• 5 In the second half of the night, winds were usually weak and variable, and up-wind or cross-wind directions of displacement were sometimes observed.
• 6 Moths were observed to disperse rapidly during their migration, so that the mass influxes which lead to outbreaks must be a consequence of subsequent reconcentration. The importance of meso-scale wind convergence zones in reconcentrating flying moths, and the role of rainfall in inducing descent, and possibly landing, are discussed.

     

A review of Afrotropical nightjar mortality,mainly road kills

Afrotropical nightjars have evolved a number of adaptations that enable them to cope well with various natural mortality factors facing them. They are, however, extremely vulnerable to the many human factors that affect their lives. Three areas of human activity are of major concern: road traffic, habitat destruction and food gathering. Suitable legislation, with strict enforcement, should ensure that nightjar populations are not harvested excessively for the table. The many National Parks, Game Reserves and other protected areas throughout the Afrotropics should ensure that sufficient breeding and feeding habitats are available for the various nightjar species. There does not appear to be a solution to the road traffic problem, so road kills will almost certainly be the major mortality factor affecting most adult nightjars. A subjective assessment was made of the road mortality suffered by each species. The forest species (Brown Nightjar, Veles binotatus, Bates's Nightjar, Caprimulgus batesi and Prigogine's Nightjar, C. prigoginei) appear to be unaffected. The level of road mortality was rated as very low for Swamp Nightjar, C. natalensis, Nechisar Nightjar, C. solala, Golden Nightjar, C. eximius and Egyptian Nightjar, C. aetfvptius; as low for Montane Nightjar, C. poliocephalus, Slender-tailed Nightjar, C. clarus, Star-spotted Nightjar, C. stellatush and Red-necked Nightjar, C. ruficollis; as moderate for Fiery-necked (Pectoral) Nightjar, C. pectoralis, Donaldson-Smith's Nightjar, C. donaldsoni, Dark Nightjar, C. fraenatus and Nubian Nightjar, C. nubicus; as high for Freckled Nightjar, C. tristigma, Rufous-cheeked Nightjar, C. rufigena, Plain Nightjar, C. inornatus and Standard-winged Nightjar, Macrodipteryx longipennis; and as very high for Long-tailed Nightjar, C. climacurus, Square-tailed Nightjar, C. fossii, European Nightjar, C. europaeus and Pennant-winged Nightjar, Semeiophorus vexillarius. Throughout the Afrotropical Region nightjars die on the roads in great numbers and for some species the road toll is a major mortality factor.     

Grizzly bear movements relative to roads: application of step selection functions

Access management is among the most important conservation actions for grizzly bears in North America. In Alberta, Canada, nearly all grizzly bear mortalities are caused by humans and occur near roads and trails. Consequently, understanding how bears move relative to roads is of crucial importance for grizzly bear conservation. We present the first application of step‐selection functions to model habitat selection and movement of grizzly bears. We then relate this to a step‐length analysis to model the rate of movement through various habitats. Grizzly bears of all sex and age groups were more likely to select steps closer to roads irrespective of traffic volume. Roads are associated with habitats attractive to bears such as forestry cutblocks, and models substituting cutblocks for roads outperformed road models in predicting bear selection during day, dawn, and dusk time periods. Bear step lengths increased near roads and were longest near highly trafficked roads indicating faster movement when near roads. Bear selection of roads was consistent throughout the day; however, time of day had a strong influence over selection of forest structure and terrain variables. At night and dawn, bears selected forests of intermediate age between 40 and 100 yr, and bears selected older forests during the day. At dawn, bears selected steps with higher solar radiation values, whereas, at dusk, bears chose steps that were significantly closer to edges. Because grizzly bears use areas near roads during spring and most human‐caused mortalities occur near roads, access management is required to reduce conflicts between humans and bears. Our results support new conservation guidelines in western North America that encourage the restriction of human access to roads constructed for resource extraction.     

Diurnal body temperature patterns in free‐ranging populations of two southern African arid‐zone nightjars

Endotherms allocate large amounts of energy and water to the regulation of a precise body temperature (T_b), but can potentially reduce thermoregulatory costs by allowing T_b to deviate from normothermic levels. Many data on heterothermy at low air temperatures (T_a) exist for caprimulgids, whereas data on thermoregulation at high T_a are largely absent, despite members of this taxon frequently roosting and nesting in sites exposed to high operative temperatures. We investigated thermoregulation in free‐ranging rufous‐cheeked nightjars Caprimulgus rufigena and freckled nightjars Caprimulgus tristigma in the southern African arid zone. Individuals of both species showed labile T_b fluctuating around a single modal T_b (T_b‐mod). Average T_b‐mod was 39.7°C for rufous‐cheeked nightjars and 39.0°C for freckled nightjars. In both species, diurnal T_b increased with increasing T_a. At T_a ≥ 38°C, rufous‐cheeked nightjar mean T_b increased to 42°C, equivalent to 2.3°C above T_b‐mod. Under similar conditions, freckled nightjar T_b was on average only 1.1°C above T_b‐mod, with a mean T_b of 40.0°C. Freckled nightjars are one of the most heterothermic caprimulgids investigated to date, but our data suggest that during hot conditions this species maintains T_b within a narrow range above T_b‐mod, possibly reflecting an evolutionary tradeoff between decreased thermal sensitivity to lower T_b but increased sensitivity to high T_b. These findings reveal how general thermoregulatory patterns at similar T_a can vary even among closely related species.     

Non‐Parametric Entrainment by Natural Twilight in the Microchiropteran Bat,Hipposideros Speoris Inside a Cave

The study aimed to determine the influence of repeated natural dawn and dusk twilight pulses in entraining the circadian flight activity rhythm of the microchiropteran bat, Hipposideros speoris, free‐running in constant darkness in a natural cave. The bats were exposed to repeated dawn or dusk twilight pulses at eight circadian phases. All bats exposed to dawn twilight pulses were entrained by advancing transients, and the stable entrainment was reached when the onset of activity occurred about 12 h before the lights‐on of the pulses, irrespective of the initial phase at which the bats were exposed to twilight. All bats exposed to dusk twilight pulses, however, were entrained by delaying transients, and the stable entrainment was reached when the onset of activity occurred about 1.6 h after the lights‐on of the pulses. The entrainment caused by dawn and dusk twilight pulses is discussed in the context of the postulated two photoreceptors: the short wavelength sensitive (S) photoreceptors mediating entrainment via dusk twilight, and the medium wavelength sensitive (M) photoreceptors mediating entrainment via dawn twilight.     

A review of the evidence for the translocation of eggs and young by nightjars (Caprimulgidae)

I undertook a review of evidence for the translocation of eggs and young by nightjars through a comprehensive search of the literature. I found that most of the evidence was based on hearsay, supposition, a misunderstanding of nightjar behaviour, or on the repetition of a story going back 200 years to Le Vaillant, via Audubon. There is no satisfactory direct evidence of any nightjar deliberately airlifting its eggs or young away from a disturbance. This conclusion is based on 10 studies by ornithologists across five continents, involving over 522 nests of 13 species. Accidental airlifting occurs occasionally when an egg or young chick gets stuck to the ventral plumage of the sitting adult. This is most likely to occur near the time of hatching. A nightjar can move an egg along the ground for a short distance, either by placing its lower mandible over the egg and then walking backwards while rolling it, or by rolling it forwards with its feet. Nightjars regularly move their chicks, especially after a disturbance. They do so by first moving away themselves and then calling. The chicks, who are highly mobile within hours of hatching, respond immediately by running to the parent birds.     

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.     

Trapped in the darkness of the night: thermal and energetic constraints of daylight flight in bats

Bats are one of the most successful mammalian groups, even though their foraging activities are restricted to the hours of twilight and night-time. Some studies suggested that bats became nocturnal because of overheating when flying in daylight. This is because--in contrast to feathered wings of birds--dark and naked wing membranes of bats efficiently absorb short-wave solar radiation. We hypothesized that bats face elevated flight costs during daylight flights, since we expected them to alter wing-beat kinematics to reduce heat load by solar radiation. To test this assumption, we measured metabolic rate and body temperature during short flights in the tropical short-tailed fruit bat Carollia perspicillata at night and during the day. Core body temperature of flying bats differed by no more than 2°C between night and daytime flights, whereas mass-specific CO(2) production rates were higher by 15 per cent during daytime. We conclude that increased flight costs only render diurnal bat flights profitable when the relative energy gain during daytime is high and risk of predation is low. Ancestral bats possibly have evolved dark-skinned wing membranes to reduce nocturnal predation, but a low degree of reflectance of wing membranes made them also prone to overheating and elevated energy costs during daylight flights. In consequence, bats may have become trapped in the darkness of the night once dark-skinned wing membranes had evolved.     

Non-parametric entrainment by natural twilight in the microchiropteran bat, Hipposideros speoris inside a cave

The study aimed to determine the influence of repeated natural dawn and dusk twilight pulses in entraining the circadian flight activity rhythm of the microchiropteran bat, Hipposideros speoris, free-running in constant darkness in a natural cave. The bats were exposed to repeated dawn or dusk twilight pulses at eight circadian phases. All bats exposed to dawn twilight pulses were entrained by advancing transients, and the stable entrainment was reached when the onset of activity occurred about 12 h before the lights-on of the pulses, irrespective of the initial phase at which the bats were exposed to twilight. All bats exposed to dusk twilight pulses, however, were entrained by delaying transients, and the stable entrainment was reached when the onset of activity occurred about 1.6 h after the lights-on of the pulses. The entrainment caused by dawn and dusk twilight pulses is discussed in the context of the postulated two photoreceptors: the short wavelength sensitive (S) photoreceptors mediating entrainment via dusk twilight, and the medium wavelength sensitive (M) photoreceptors mediating entrainment via dawn twilight.     

Food of the nightjars in Zimbabwe

Jackson, H.D. 2000. Food of the nightjars in Zimbabwe. Ostrich 71 (3&4): 404-407. Analysis of the stomach contents of 282 nightjars (Caprimulgidae) collected in Zimbabwe showed that the seven species concerned feed almost exclusively on nocturnal flying insects, predominantly on beetles (Caleoptera, 52.7%). especially lamellicom beetles (Scarabaeidae, 67.5% of all beetles), and to a lesser extent on moths (Lepidoptera, 12.7%), grasshoppers (Orthoptem, 7.2%). bugs (Hemiptera, 3.5%) and, when available, termites (Isoptera, 20.4%). There is some seasonal variation in diet, and prey selection varies between species. Deliberate ingestion of grit to aid digestion is unlikely.