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.     

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.     

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.     

Hunting success of lions affected by the moon's phase in a wooded habitat

Variation in moonlight affects foraging, hunting and vigilance behaviours in many nocturnal species. Here, we explore the effect of moonlight on the movement and hunting behaviour of African lions. Previous studies found bright moonlight is associated with reduced hunting success; however, those studies were largely undertaken in open habitats where predators are easily seen by prey species on moonlit nights. In this study, we explored whether moonlight affected hunting behaviour and success in a largely wooded environment. Measures of short‐term lion movements (distance moved, displacement and path tortuosity) derived from GPS telemetry data were used as indicators of movement behaviour. Field observations of belly distension were used to assess recent food intake. Lions had greater belly distension (indicating feeding success) on dark nights. However, this change in feeding success was not reflected by lion movement patterns—there was no evidence that these changed across moon phases. There was no evidence that lions used more covered habitats on brighter nights to facilitate concealment.     

Aerial insectivorous bat activity in relation to moonlight intensity

It is commonly assumed that aerial insectivorous bats in the tropics respond to moonlight intensity by decreasing their foraging activity during bright nights due either to an increase in predation risk, or to a reduction in insect availability.The effect of moonlight on bat activity can be measured both between nights and within a single night. However, few studies have simultaneously used both approaches, and most authors generally compare bat activity with lunar phases. Our main aim was to evaluate how moonlight influences aerial insectivorous bat activity at different time scales: between nights and within the same night. Activity of five bat species was measured using autonomous ultrasound recording stations and moonlight intensity percentages retrieved from the Moontool program nightly throughout a 53-day sampling period. Only one species (Myotis riparius) responded negatively to moonlight, while two species (Pteronotus parnellii and Saccopteryx leptura) increased their foraging activity in moonlight. For Cormura brevirostris and S. bilineata, moonlight intensity did not affect activity level. Bat activity was greater for all species at the beginning of the night, independent of the presence of the moon, indicating that foraging just after the sunset is adaptive. Thus, bat response to the effect of moonlight intensity is more apparent between nights than within a single night and may depend on species-specific traits, such as flight speed, flexibility in habitat use and body size.     

Patch use in time and space for a meso-predator in a risky world

Predator–prey studies often assume a three trophic level system where predators forage free from any risk of predation. Since meso-predators themselves are also prospective prey, they too need to trade-off between food and safety. We applied foraging theory to study patch use and habitat selection by a meso-predator, the red fox. We present evidence that foxes use a quitting harvest rate rule when deciding whether or not to abandon a foraging patch, and experience diminishing returns when foraging from a depletable food patch. Furthermore, our data suggest that patch use decisions of red foxes are influenced not just by the availability of food, but also by their perceived risk of predation. Fox behavior was affected by moonlight, with foxes depleting food resources more thoroughly (lower giving-up density) on darker nights compared to moonlit nights. Foxes reduced risk from hyenas by being more active where and when hyena activity was low. While hyenas were least active during moon, and most active during full moon nights, the reverse was true for foxes. Foxes showed twice as much activity during new moon compared to full moon nights, suggesting different costs of predation. Interestingly, resources in patches with cues of another predator (scat of wolf) were depleted to significantly lower levels compared to patches without. Our results emphasize the need for considering risk of predation for intermediate predators, and also shows how patch use theory and experimental food patches can be used for a predator. Taken together, these results may help us better understand trophic interactions.     

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.     

Prey refuges as predator hotspots: ocelot (Leopardus pardalis) attraction to agouti (Dasyprocta punctata) dens

We tested the hypothesis that prey refuges attract predators, leading to elevated predator activity in the vicinity of refuges. We used camera traps to determine whether the spatial activity of a predator, the ocelot (Leopardus pardalis), was biased toward refuge locations of its principal prey, the agouti (Dasyprocta punctata). We radio-tracked agoutis at night to locate active refuges and compared the activity of ocelots between these refuges and surrounding control grid locations. We found that ocelots visited the area near agouti refuges significantly more often and for longer periods of time than control locations, and that they actively investigated the refuge entrances. Both occupied and unoccupied refuges were visited, but the duration of inspection was longer at occupied refuges. As the ocelots could probably not see the agoutis within the refuges, olfaction likely cued foraging ocelots. Two refuges were repeatedly visited by the same ocelots on different days, suggesting spatial memory. Overall, our results suggest that predators can be attracted to prey refuges or refuging prey. The benefits to prey of staying nearby a refuge would thus be counterbalanced by higher likelihoods of predator encounter. This should stimulate prey to use multiple refuges alternatingly and to not enter or exit refuges at times of high predator activity.     

Artificial light at night reverses monthly foraging pattern under simulated moonlight

Mounting evidence shows that artificial light at night (ALAN) alters biological processes across levels of organization, from cells to communities. Yet, the combined impacts of ALAN and natural sources of night-time illumination remain little explored. This is in part due the lack of accurate simulations of the complex changes moonlight intensity, timing and spectra throughout a single night and lunar cycles in laboratory experiments. We custom-built a novel system to simulate natural patterns of moonlight to test how different ALAN intensities affect predator–prey relationships over the full lunar cycle. Exposure to high intensity ALAN (10 and 50 lx) reversed the natural lunar-guided foraging pattern by the gastropod mesopredator Nucella lapillus on its prey Semibalanus balanoides. Foraging decreased during brighter moonlight in naturally lit conditions. When exposed to high intensity ALAN, foraging increased with brighter moonlight. Low intensity ALAN (0.1 and 0.5 lx) had no impact on foraging. Our results show that ALAN alters the foraging pattern guided by changes in moonlight brightness. ALAN impacts on ecosystems can depend on lunar light cycles. Accurate simulations of night-time light cycle will warrant more realistic insights into ALAN impacts and also facilitate advances in fundamental night-time ecology and chronobiology.     

Amphipod (Gammarus minus) responses to predators and predator impact on amphipod density

Recent theoretical work suggests that predator impact on local prey density will be the result of interactions between prey emigration responses to predators and predator consumption of prey. Whether prey increase or decrease their movement rates in response to predators will greatly influence the impact that predators have on prey density. In stream systems the type of predator, benthic versus water-column, is expected to influence whether prey increase or decrease their movement rates. Experiments were conducted to examine the response of amphipods (Gammarus minus) to benthic and water-column predators and to examine the interplay between amphipod response to predators and predator consumption of prey in determining prey density. Amphipods did not respond to nor were they consumed by the benthic predator. Thus, this predator had no impact on amphipod density. In contrast, amphipods did respond to two species of water-column predators (the predatory fish bluegills, Lepomis macrochirus, and striped shiners, Luxilus chrysocephalus) by decreasing their activity rates. This response led to similar positive effects on amphipod density at night by both species of predatory fish. However, striped shiners did not consume many amphipods, suggesting their impact on the whole amphipod “population” was zero. In contrast, bluegills consumed a significant number of amphipods, and thus had a negative impact on the amphipod “population”. These results lend support to theoretical work which suggests that prey behavioral responses to predators can mask the true impact that predators have on prey populations when experiments are conducted at small scales. Received: 21 March 1997 / Accepted: 15 December 1997     

The fear diet: Risk,refuge, and biological control by omnivorous weed seed predators

Weed seed biocontrol by omnivorous mice and insects can limit weed seedbanks, but this ecosystem service can be difficult to predict given the broad diet breadth of seed predators and their potential for intraguild predation. Seed foraging behavior is further modified by fluctuating cues of predation risk from higher trophic levels and the availability of refuge habitat. Uncertainty about whether co-occurring insects and mice additively contribute to weed biocontrol or interfere with each other via intraguild predation limits our ability to recommend habitat management strategies that reliably promote seed destruction. Using seed removal assays, fluorescent powder tracking, and stable isotope analyses, we assessed effects of a predation risk cue (moonlight) on mouse foraging patterns in a patchwork of vegetated and exposed plots in a cultivated field. Mouse foraging activity decreased on exposed ground during the full moon, compared to dark nights, yet foraging movements were unaffected by moon cycle within refuge patches. Weed seed consumption was more than three times higher in cover than exposed soil, and 78% of that difference was attributable to invertebrate granivores. Mice and invertebrate granivores both exhibited higher foraging activity in cover, indicating co-occurrence of intraguild predators and prey. However, stable isotope analyses of fecal samples revealed that mice captured in refuge habitats fed at slightly lower trophic levels than those in exposed habitats (suggesting minimal intraguild predation in refuge habitat), and mouse diet was unaffected by moonlight. Despite increased availability of invertebrate prey in cover patches, mice do not appear to preferentially exploit prey when avoiding their own predators or interfere with weed seed predation. Therefore, functional redundancy of mice and invertebrate seed predators in cover crops and other refuge habitats may strengthen and stabilize weed seed biocontrol.     

Induced changes in island fox (Urocyon littoralis) activity do not mitigate the extinction threat posed by a novel predator

Prey response to novel predators influences the impacts on prey populations of introduced predators, bio-control efforts, and predator range expansion. Predicting the impacts of novel predators on native prey requires an understanding of both predator avoidance strategies and their potential to reduce predation risk. We examine the response of island foxes (Urocyon littoralis) to invasion by golden eagles (Aquila chrysaetos). Foxes reduced daytime activity and increased night time activity relative to eagle-na?ve foxes. Individual foxes reverted toward diurnal tendencies following eagle removal efforts. We quantified the potential population impact of reduced diurnality by modeling island fox population dynamics. Our model predicted an annual population decline similar to what was observed following golden eagle invasion and predicted that the observed 11% reduction in daytime activity would not reduce predation risk sufficiently to reduce extinction risk. The limited effect of this behaviorally plastic predator avoidance strategy highlights the importance of linking behavioral change to population dynamics for predicting the impact of novel predators on resident prey populations.     

Introduced species provide a novel temporal resource that facilitates native predator population growth

Non-native species are recognized as important components of change to food web structure. Non-native prey may increase native predator populations by providing an additional food source and simultaneously decrease native prey populations by outcompeting them for a limited resource. This pattern of apparent competition may be important for plants and sessile marine invertebrate suspension feeders as they often compete for space and their immobile state make them readily accessible to predators. Reported studies on apparent competition have rarely been examined in biological invasions and no study has linked seasonal patterns of native and non-native prey abundance to increasing native predator populations. Here, we evaluate the effects of non-native colonial ascidians (Diplosoma listerianum and Didemnum vexillum) on population growth of a native predator (bloodstar, Henricia sanguinolenta) and native sponges through long-term surveys of abundance, prey choice and growth experiments. We show non-native species facilitate native predator population growth by providing a novel temporal resource that prevents loss of predator biomass when its native prey species are rare. We expect that by incorporating native and non-native prey seasonal abundance patterns, ecologists will gain a more comprehensive understanding of the mechanisms underlying the effects of non-native prey species on native predator and prey population dynamics.     

Predator odor recognition and antipredatory response in fish: does the prey know the predator diel rhythm?

We studied in a laboratory experiment using stream tanks if two percid prey fish, the perch (Perca fluviatilis) and the ruffe (Gymnocephalus cernuus), can recognize and respond to increased predation risk using odors of two piscivores, the pike (Esox lucius) and the burbot (Lota lota). Burbot is night-active most of the year but pike hunts predominantly visually whenever there is enough light. Perch is a common day-active prey of pike and dark-active ruffe that of burbot. We predicted that besides recognizing the predator odors, the prey species would respond more strongly to odors of the predator which share the same activity pattern. Both perch and ruffe clearly responded to both predator fish odors. They decreased movements and erected the spiny dorsal fins. Fin erection showed clearly the black warning ornamentation in the fin and thus erected fin may function besides as mechanical defense also as warning ornament for an approaching predator. No rapid escape movements were generally observed. Both perch and ruffe responded more strongly to pike odor than to burbot. There were no clear differences in response between day and night. In conclusion, we were able to verify clear predator odor recognition by both prey fish. Both perch and ruffe responded to both predator odors and it seemed that pike forms a stronger threat for both prey species. Despite of diel activity differences both perch and ruffe used the same antipredatory strategies, but the day-active perch seemed to have a more flexible antipredatory behavior by responding more strongly to burbot threat during the night when burbot is active.     

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.     

Importance of visual cues of conspecifics and predators during the habitat selection of coral reef fish larvae

The study investigated visual recognition of conspecifics and predators by settlement-stage coral reef fish larvae in a set of three experiments using a dual-choice aquarium (Moorea Island). Experiments 1 and 2 were conducted under artificial light conditions. Experiment 3 was conducted under natural light during new and full moon nights. In experiment 1, five out of six species preferred conspecifics rather than heterospecifics (Acanthurus triostegus, Chromis viridis, Ostorhinchus angustatus, Stegastes fasciolatus, Valenciaenna strigata). In experiment 2, three out of six species were repulsed by predators (Mulloidichtys flavolineatus, O. angustatus, V. strigata). In experiment 3 (conducted on one species), A. triostegus was attracted to conspecifics during bright nights, but did not show such behavior during dark nights. Our study raises the question of trade-off for fish larvae to settle during the night with high light intensities to favor the visual recognition of conspecifics and predators, or during darker nights to reduce reef predation.     

Growth rate as a measure of food value in thaidid gastropods: Assumptions and implications for prey morphology and distribution

Rates of body growth were measured for three species of rocky intertidal gastropods: Thais (or Nucella) lamellosa (Gmelin), T. (or N.) canaliculata (Duclos), and T. (or N.) emarginata (Deshayes). Three size classes of each predator species were grown experimentally in cages at two tidal heights on four size classes of each prey species: Semibalanus cariosus (Pallas), Balanus glandula Darwin, and Mytilus edulis L. Growth was also assessed for Thais canaliculata feeding on Mytilus californianus Conrad, and Thais emarginata feeding on Chthamalus dalli Pilsbry.Rates of body growth varied as a function of predator size, prey size and prey species. With one exception (large Thais canaliculata) intermediate sized Balanus glandula promoted the most rapid growth for all sizes of all three species of Thais; thus these potentially competing predator species have the same highest ranked prey. Among prey promoting slower growth, those encountered commonly in the normal habitats of the predator promoted more rapid growth than those encountered rarely, suggesting that past evolutionary experience has influenced present food value (= growth potential) of prey. For some predators, the ranking of prey species changed with predator size; Chthamalus dalli and similar sized Balanus glandula promoted comparable growth in small Thais emarginata but Chthamalus dalli promoted much slower growth than similar sized Balanus glandula in larger Thais emarginata. Rank differences based on prey size in Balanusglandula became more pronounced and in some instances changed with increasing predator size; larger B. glandula promoted relatively faster growth for large snails than for small snails in all Thais species. Finally, growth rates were correlated with two important attributes of fitness in T. canaliculata and T. emarginata; in general, prey promoting more rapid growth also resulted in an earlier age of first reproduction for initially immature snails and higher rate of egg capsule production for mature individuals. Thus, these growth rates provide a basis from which to examine quantitatively patterns of prey selection from an energy- or growth-maximization perspective.These patterns of predator growth also permit inferences about the possible influence of predation on the evolution of prey morphology, life-history, and microhabitat distribution. Size refuges from Thais predation were confirmed for both Semibalanus cariosus and Mytilus californianus. Both Balanus glandula and Mytilus edulis, on the other hand, were vulnerable to predation by all sizes of Thais examined (> 10 mm) regardless of prey size; thus, neither of these species achieves a size refuge. In addition, on rocky shores, vertical distributions of the prey species reflect general preference patterns of their predators: higher value prey species, i.e. those with less well developed defensive morphologies (Balanus glandula, Mytilus edulis), generally occur higher on the shore. This pattern would be expected if preferred prey are consumed first lower on the shore where they are more available.     

Facilitation and interference among three predators affect their consumption of a stream-dwelling mayfly

1. We experimentally tested if a multiplicative risk model accurately predicted the consumption of a common mayfly at risk of predation from three predator species in New Zealand streams. Deviations between model predictions and experimental observations were interpreted as indicators of ecologically important interactions between predators. 2. The predators included a drift‐feeding fish [brown trout (T), Salmo trutta], a benthivorous fish [galaxiid (G), koaro, Galaxias brevipennis] and a benthic predatory stonefly (S; Stenoperla sp.) with Deleatidium sp. mayflies as prey. Eight treatments with all predator species combinations and a predator‐free control were used. Experiments were performed in aquaria with cobbles as predator refuges for mayflies and we measured the proportion of prey consumed after 6 h for both day and night trials. 3. Trout consumed a higher proportion of prey than other predators. For the two predator treatments we found less than expected prey consumption in the galaxiid + trout treatment (G + T) for both day and night trials, whereas a higher than expected proportion of prey was consumed during night time in the stonefly + trout (S + T) treatment. 4. The results indicate interference (G + T) and facilitation (S + T) between predators depending on predator identity and time of day. Thus, to make accurate predictions of interspecific interactions, it is necessary to consider the ecology of individual species and how differences influence the direction and magnitude of interactions.     

Population and behavioural responses of native prey to alien predation

The introduction of invasive alien predators often has catastrophic effects on populations of naïve native prey, but in situations where prey survive the initial impact a predator may act as a strong selective agent for prey that can discriminate and avoid it. Using two common species of Australian small mammals that have persisted in the presence of an alien predator, the European red fox Vulpes vulpes, for over a century, we hypothesised that populations of both would perform better where the activity of the predator was low than where it was high and that prey individuals would avoid signs of the predator’s presence. We found no difference in prey abundance in sites with high and low fox activity, but survival of one species—the bush rat Rattus fuscipes—was almost twofold higher where fox activity was low. Juvenile, but not adult rats, avoided fox odour on traps, as did individuals of the second prey species, the brown antechinus, Antechinus stuartii. Both species also showed reduced activity at foraging trays bearing fox odour in giving-up density (GUD) experiments, although GUDs and avoidance of fox odour declined over time. Young rats avoided fox odour more strongly where fox activity was high than where it was low, but neither adult R. fuscipes nor A. stuartii responded differently to different levels of fox activity. Conservation managers often attempt to eliminate alien predators or to protect predator-naïve prey in protected reserves. Our results suggest that, if predator pressure can be reduced, otherwise susceptible prey may survive the initial impact of an alien predator, and experience selection to discriminate cues to its presence and avoid it over the longer term. Although predator reduction is often feasible, identifying the level of reduction that will conserve prey and allow selection for avoidance remains an important challenge.