Variation in budburst phenology of Douglas-fir related to western spruce budworm (Lepidoptera: Tortricidae) fitness

Variation in budburst phenology among individual trees of interior Douglas-fir (Pseudotsuga menziesii var. glauca [Beissn.] Franco) may influence their susceptibility to western spruce budworm (Choristoneura occidentalis Freeman) defoliation. We tested the hypothesis that phenological asynchrony between Douglas-fir and the western spruce budworm is a mechanism of resistance using clones derived from parent trees that showed resistance versus susceptibility to C. occidentalis defoliation in the field. Susceptible clones had earlier budburst phenology compared with resistant clones when they were grown in a common greenhouse environment, demonstrating a genetic basis for parallel phenological differences exhibited by the parent trees. We tested the importance of phenological asynchrony as a factor influencing fitness of C. occidentalis using two different greenhouse bioassay experiments. One experiment compared western spruce budworm performance on equivalent phenological stages of susceptible and resistant clones by matching larval feeding to the columnar (fourth) bud development stage of each clone. Larvae reared on resistant clones had greater realized fitness (i.e., number of F1 offspring produced) than those reared on susceptible clones when the influence of variation in budburst phenology was minimized. In the other experiment, western spruce budworm larvae were placed on all trees on the same date when approximately 50% of all terminal buds in the population were in the yellow (second) budburst stage. Larvae reared on susceptible clones had greater realized fitness than those reared on resistant clones when the influence of phenological asynchrony was expressed. Our results suggest that resistant phenotypes of Douglas-fir have negative effects on survival and reproduction of C. occidentalis under the natural conditions that insects and trees experience in the field. Genetic variation among trees in budburst phenology has an important influence on interactions between the western spruce budworm and Douglas-fir.     

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.     

Moth wing scales slightly increase the absorbance of bat echolocation calls

Coevolutionary arms races between predators and prey can lead to a diverse range of foraging and defense strategies, such as countermeasures between nocturnal insects and echolocating bats. Here, we show how the fine structure of wing scales may help moths by slightly increasing sound absorbance at frequencies typically used in bat echolocation. Using four widespread species of moths and butterflies, we found that moth scales are composed of honeycomb-like hollows similar to sound-absorbing material, but these were absent from butterfly scales. Micro-reverberation chamber experiments revealed that moth wings were more absorbent at the frequencies emitted by many echolocating bats (40-60 kHz) than butterfly wings. Furthermore, moth wings lost absorbance at these frequencies when scales were removed, which suggests that some moths have evolved stealth tactics to reduce their conspicuousness to echolocating bats. Although the benefits to moths are relatively small in terms of reducing their target strengths, scales may nonetheless confer survival advantages by reducing the detection distances of moths by bats by 5-6%.     

Frequent Arousals from Winter Torpor in Rafinesque’s Big-Eared Bat (Corynorhinus rafinesquii)

Extensive use of torpor is a common winter survival strategy among bats; however, data comparing various torpor behaviors among species are scarce. Winter torpor behaviors are likely to vary among species with different physiologies and species inhabiting different regional climates. Understanding these differences may be important in identifying differing susceptibilities of species to white-nose syndrome (WNS) in North America. We fitted 24 Rafinesque’s big-eared bats (Corynorhinus rafinesquii) with temperature-sensitive radio-transmitters, and monitored 128 PIT-tagged big-eared bats, during the winter months of 2010 to 2012. We tested the hypothesis that Rafinesque’s big-eared bats use torpor less often than values reported for other North American cave-hibernators. Additionally, we tested the hypothesis that Rafinesque’s big-eared bats arouse on winter nights more suitable for nocturnal foraging. Radio-tagged bats used short (2.4 d ± 0.3 (SE)), shallow (13.9°C ± 0.6) torpor bouts and switched roosts every 4.1 d ± 0.6. Probability of arousal from torpor increased linearly with ambient temperature at sunset (P<0.0001), and 83% (n = 86) of arousals occurred within 1 hr of sunset. Activity of PIT-tagged bats at an artificial maternity/hibernaculum roost between November and March was positively correlated with ambient temperature at sunset (P<0.0001), with males more active at the roost than females. These data show Rafinesque’s big-eared bat is a shallow hibernator and is relatively active during winter. We hypothesize that winter activity patterns provide Corynorhinus species with an ecological and physiological defense against the fungus causing WNS, and that these bats may be better suited to withstand fungal infection than other cave-hibernating bat species in eastern North America.     

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.     

Behavioural Response of Bats to Perceived Predation Risk While Foraging

The ability to detect and respond to predation risk while foraging may have important fitness consequences for prey organisms. Anti‐predator behaviours may reduce the probability of mortality because of predation, but they may also be associated with reduced foraging efficiency. Several behaviours of bats have been suggested to serve as anti‐predator responses, and there is evidence that predation, particularly by avian predators such as owls, may be an important cause of bat mortality. Previous studies have attempted to determine whether predator presence affects bat behaviour when emerging from roost sites, but few have examined effects of predator presence on bat behaviour while foraging. In this study, we investigated whether foraging bats respond to predator cues by presenting bats with an acoustic cue simulating the presence of an owl. Within matched trials, which were conducted at different locations each of 18 nights, significantly fewer bat detections were recorded at owl playback stations than at control stations (no auditory cue), suggesting an avoidance response by bats. An acoustic control (i.e. station playing woodpecker calls), however, did not have significantly more detections than the stations playing the owl calls, suggesting that bats may simply be avoiding noise and more detailed investigation is warranted. Although evidence for owl predation on bats is minimal in North America, results of this study may indicate that the perceived presence of owls may represent a factor influencing the behaviour of bats while foraging.     

Numerical and Functional Responses of Forest Bats to a Major Insect Pest in Pine Plantations

Global change is expected to modify the frequency and magnitude of defoliating insect outbreaks in forest ecosystems. Bats are increasingly acknowledged as effective biocontrol agents for pest insect populations. However, a better understanding is required of whether and how bat communities contribute to the resilience of forests to man- and climate-driven biotic disturbances. We studied the responses of forest insectivorous bats to a major pine defoliator, the pine processionary moth pityocampa, which is currently expanding its range in response to global warming. We used pheromone traps and ultrasound bat recorders to estimate the abundance and activity of moths and predatory bats along the edge of infested pine stands. We used synthetic pheromone to evaluate the effects of experimentally increased moth availability on bat foraging activity. We also evaluated the top-down regulation of moth population by estimating T. pityocampa larval colonies abundance on the same edges the following winter. We observed a close spatio-temporal matching between emergent moths and foraging bats, with bat activity significantly increasing with moth abundance. The foraging activity of some bat species was significantly higher near pheromone lures, i.e. in areas of expected increased prey availability. Furthermore moth reproductive success significantly decreased with increasing bat activity during the flight period of adult moths. These findings suggest that bats, at least in condition of low prey density, exhibit numerical and functional responses to a specific and abundant prey, which may ultimately result in an effective top-down regulation of the population of the prey. These observations are consistent with bats being useful agents for the biocontrol of insect pest populations in plantation forests.     

Do the evolutionary interactions between moths and bats promote niche partitioning between bats and birds?

Ecological theory suggests that the coexistence of species is promoted by the partitioning of available resources, as in dietary niche partitioning where predators partition prey. Yet, the mechanisms underlying dietary niche partitioning are not always clear. We used fecal DNA metabarcoding to investigate the diets of seven nocturnal insectivorous bird and bat species. Low diet overlap (2%–22%) supported resource partitioning among all species. Differences in diet corresponded with species identity, prey detection method, and foraging behavior of predators. Insects with ultrasonic hearing capabilities were consumed significantly more often by birds than bats, consistent with an evolved avoidance of echolocating strategies. In turn, bats consumed a greater proportion of noneared insects such as spruce budworms. Overall, our results suggest that evolutionary interactions among bats and moths translate to dietary niche partitioning and coexistence among bats and nocturnal birds.     

High duty cycle pulses suppress orientation flights of crambid moths

Bat-and-moth is a good model system for understanding predator–prey interactions resulting from interspecific coevolution. Night-flying insects have been under predation pressure from echolocating bats for 65 Myr, pressuring vulnerable moths to evolve ultrasound detection and evasive maneuvers as counter tactics. Past studies of defensive behaviors against attacking bats have been biased toward noctuoid moth responses to short duration pulses of low-duty-cycle (LDC) bat calls. Depending on the region, however, moths have been exposed to predation pressure from high-duty-cycle (HDC) bats as well. Here, we reveal that long duration pulse of the sympatric HDC bat (e.g., greater horseshoe bat) is easily detected by the auditory nerve of Japanese crambid moths (yellow peach moth and Asian corn borer) and suppress both mate-finding flights of virgin males and host-finding flights of mated females. The hearing sensitivities for the duration of pulse stimuli significantly dropped non-linearly in both the two moth species as the pulse duration shortened. These hearing properties support the energy integrator model; however, the threshold reduction per doubling the duration has slightly larger than those of other moth species hitherto reported. And also, Asian corn borer showed a lower auditory sensitivity and a lower flight suppression to short duration pulse than yellow peach moth did. Therefore, flight disruption of moth might be more frequently achieved by the pulse structure of HDC calls. The combination of long pulses and inter-pulse intervals, which moths can readily continue detecting, will be useful for repelling moth pests.     

食鱼蝙蝠形态和行为特化研究

总结了食鱼蝙蝠种类、分布 ,及其形态结构、回声定位功能和捕食行为的研究成果。比较食鱼蝙蝠与近水面“拖网式”食虫蝙蝠在形态、回声定位信号及捕食行为方面的异、同 ,推测食鱼蝙蝠起源于“拖网式”食虫蝙蝠类 ;体形和回声定位信号的几种特异性是捕食行为进化压力 ,而环境是决定因素。     

Lunar phobia in bats and its ecological correlates: A meta-analysis

Animals show several behavioral strategies to reduce predation risks. Presumably, moonlight avoidance is a strategy used by some nocturnal species to reduce the risk of predation. In bats, some research indicates that foraging activity is negatively correlated with moonlight intensity, a phenomenon better known as lunar phobia. However, the currently available evidence is contradictory because some bat species reduce their activity during nights with more moonlight while the opposite occurs in other species. We quantitatively evaluated the strength and direction of the relationship between moonlight intensity and bat activity using a meta-analysis. We also looked at some ecological correlates of lunar phobia in bats. Specifically, we examined foraging habitat and latitude as potential moderators of the size of the lunar phobia effect. Our results show that, regardless of the method used to evaluate bat activity, the overall relationship between moonlight intensity and bat activity is significant and negative (r = ?0.22). Species foraging on the surface of the water (piscivores and insectivores; r = ?0.83) and forest canopy species (i.e., big frugivores; r = ?0.30) are more affected by moonlight than those with different foraging habitats (understory, subcanopy, open air). Latitude was positively correlated with lunar phobia (r = 0.023). The stronger lunar phobia for bats foraging on the water surface and in the forest canopy may suggest that the risk of predation is greater where moonlight penetrates more easily. The significant effect of latitude as a moderator of lunar phobia suggests that there is a weak geographic pattern, with this phobia slightly more common in tropical bats than in temperate species.     

Foraging and diet of the northern bat Eptesicus nilssoni in Sweden

Northern bats foraged predominantly in small individual and transient feeding sites usually in open places near trees and over water. Lakes were preferred as foraging habitat as compared to woodlands and farmlands. The pattern of habitat selection did not change drastically during the period of reproduction. The diet was probably unselective, consisting mainly of small dipterans and moths and also caddis flies, lacewings and mayflies (lake area) and dung beetles (farmland). Northern bats consumed insects of 3–30 mm body length, thus taking prey items of a broader size range than other bat species studied.     

Light,predation and the lekking behaviour of the ghost swift Hepialus humuli (L.) (Lepidoptera,Hepialidae)

We examined the timing of the crepuscular lekking flight of male ghost swift moths in southern Sweden with respect to variations in: (i) the quality of the visual mating signal; and (ii) the behaviour of potential vertebrate predators (mainly bats). The moths'' display flights started ca. 57 min after sunset, and occurred during 20–30 min at incident light intensities between 10.0 and 2.0 lux. Owing to the falling and more shortwave ambient light after sunset, the brightness contrast between the moth wings and the background (grass) increased steeply at the time of display onset. The silvery white male wing colour thereby seems to maximize conspicuousness, and may be a secondary adaptation that facilitates visibility at low light intensities. The display timing itself is probably determined by other factors, possibly predation. By displaying only for a short period at dusk, the moths seem to avoid most birds, which normally do not forage at these light levels, and gleaning bats, which typically do not start to feed until the light intensity has fallen even further. Nevertheless, aerial-hawking bats were often (54% of the evenings, n = 22) seen at the leks, and one species (Eptesicus nilssonii) frequently fed on the displaying moths (22% of the moths observed, n = 83). H. humuli represents an ancient clade among the Lepidoptera. By restricting its sexual behaviour to a short time window at dusk, when predation risk may be minimized (but still high), it may to some extent compensate for the lack of sophisticated predator defence systems such as aposematic or mimetic coloration, manoeuvrable flight, and ultrasonic hearing, which predominate among the more recent Lepidopteran clades. However, the time window solution restricts the moths'' activities considerably and the lack of defence still carries a considerable cost in terms of predation.     

Activity patterns of the Ryukyu flying fox on a subtropical island: responses to seasonal changes in night length

Almost all chiropteran species are nocturnal, but some species are occasionally active during the daytime. We conducted radio-tracking surveys and direct observations of the Ryukyu flying fox, Pteropus dasymallus, in two different habitats—urbanized and forested areas—on a subtropical island from April 2002 to January 2006. We recorded the departure time and return time from/to day roosts as well as behavioral time budgets during the night. The departure and return times shifted in correspondence with seasonal changes in sunset and sunrise times. The Ryukyu flying fox tended to depart earlier in summer when the night length was shorter, suggesting that it adjusts its active period by departing earlier. On the contrary, the amount of foraging performed by the bats in urbanized areas decreased in the summer when fruits of Ficus microcarpa were more abundant, suggesting that the bats adjust their behavioral time budgets in line with local food availability. Daytime activity was observed only in the forested area. In conclusion, the duration of Ryukyu flying fox activity was found to primarily depend on seasonal changes in the light–dark cycle, and this bat may adjust its behavioral time budget according to local food availability and the intensity of human activities.     

Impact of predation by a cave-dwelling bat, Rhinolophus ferrumequinum, on the diapausing population of a troglophilic moth, Goniocraspidum preyeri

The predation pressure of the greater horseshoe bat, Rhinolophus ferrumequinum, on the diapausing population of the noctuid moth, Goniocraspidum preyeri, was examined at an abandoned mine in central Japan. These bats did not prey on the moths in summer. The number of moths preyed on was largest in March. More than 60% of the diapausing moths were eaten by only three or four bats, which accounted for over 90% of all the moth deaths. This moth was an important source of food at the end of the hibernating period when the bat had already used most of its stored subcutaneous fat. The predation on G. preyeri may have raised the overwintering success rate of R. ferrumequinum.     

The Importance of Acacia Trees for Insectivorous Bats and Arthropods in the Arava Desert

Anthropogenic habitat modification often has a profound negative impact on the flora and fauna of an ecosystem. In parts of the Middle East, ephemeral rivers (wadis) are characterised by stands of acacia trees. Green, flourishing assemblages of these trees are in decline in several countries, most likely due to human-induced water stress and habitat changes. We examined the importance of healthy acacia stands for bats and their arthropod prey in comparison to other natural and artificial habitats available in the Arava desert of Israel. We assessed bat activity and species richness through acoustic monitoring for entire nights and concurrently collected arthropods using light and pit traps. Dense green stands of acacia trees were the most important natural desert habitat for insectivorous bats. Irrigated gardens and parks in villages and fields of date palms had high arthropod levels but only village sites rivalled acacia trees in bat activity level. We confirmed up to 13 bat species around a single patch of acacia trees; one of the richest sites in any natural desert habitat in Israel. Some bat species utilised artificial sites; others were found almost exclusively in natural habitats. Two rare species (Barbastella leucomelas and Nycteris thebaica) were identified solely around acacia trees. We provide strong evidence that acacia trees are of unique importance to the community of insectivorous desert-dwelling bats, and that the health of the trees is crucial to their value as a foraging resource. Consequently, conservation efforts for acacia habitats, and in particular for the green more densely packed stands of trees, need to increase to protect this vital habitat for an entire community of protected bats.     

Auditory changes in noctuid moths endemic to a bat-free habitat

If the ears of moths exist primarily to detect the echolocation calls of hunting bats, endemic moths in bat-free areas (i.e., species that have evolved in the absence of the selection pressure maintaining ears) should exhibit signs of deafness. Noctuid moths from the Pacific islands of French Polynesia, a site that has never possessed bats, were sampled and electrophysiologically analysed to test this hypothesis. The auditory sensitivities of seven endemic and twelve immigrant species, captured from the islands of Tahiti, Moorea, and Hiva Oa, were compared. Both classes possess neurally responsive ears, but endemics are significantly deafer than immigrants at frequencies above 35 kHz. This form of deafness is similar to other moths presumably released from the selection pressure of bat predation. I conclude that endemic moths at this site exhibit preliminary stages of deafness and that, considering their small cellular investment, ears in moths will be lost at a slower rate than more complex sensory organs.     

Auditory sensitivity of Hawaiian moths (Lepidoptera: Noctuidae) and selective predation by the Hawaiian hoary bat (Chiroptera: Lasiurus cinereus semotus).

The islands of Hawai'i offer a unique opportunity for studying the auditory ecology of moths and bats since this habitat has a single species of bat, the Hawaiian hoary bat (Lasiurus cinereus semotus), which exerts the entire predatory selection pressure on the ears of sympatric moths. I compared the moth wings discarded by foraging bats with the number of surviving moths on the island of Kaua'i and concluded that the endemic noctuid Haliophyle euclidias is more heavily preyed upon than similar-sized endemic (e.g. Agrotis diplosticta) and adventive (Agrotis ipsilon and Pseudaletia unipuncta) species. Electrophysiological examinations indicated that, compared with species less preyed upon, H. euclidias has lower auditory sensitivities to the bat's social and echolocation calls, which will result in shorter detection distances of the bat. The poor ears of H. euclidias suggest that this moth coevolved with the bat using non-auditory defences that resulted in auditory degeneration. This moth now suffers higher predation because it is drawn away from its normal habitat by the man-made lights that are exploited by the bat.     

Effects of artificial and western spruce budworm (Lepidoptera: Tortricidae) defoliation on growth and biomass allocation of Douglas-fir seedlings

Artificial defoliation has been used commonly to simulate defoliation by insect herbivores in experiments, in spite of the fact that obvious differences exist between clipping foliage and natural defoliation due to insect feeding. We used a greenhouse experiment to compare the effects of artificial and western spruce budworm (Choristoneura occidentalis Freeman) defoliation on the growth and biomass allocation of 3-yr old half-sib seedlings from mature Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco variety glauca] trees that showed phenotypic resistance versus susceptibility to budworm defoliation in the forest. Artificial clipping of buds mimicked the effects of budworm feeding on total seedling biomass when 50% of the terminal buds were damaged. However, artificial defoliation decreased seedling height, relative growth rate of height, and shoot: root ratio more than budworm defoliation, whereas budworm defoliation decreased stem diameter relative growth rate more than artificial defoliation. Half-sib seedling progeny from resistant maternal tree phenotypes had greater height, diameter, biomass, and shoot: root ratio than seedlings from susceptible phenotypes. We concluded that careful artificial defoliation could generally simulate effects of budworm defoliation on total biomass of Douglas-fir seedlings, but that the two defoliation types did not have equal effects on biomass allocation between shoot and root. Further, an inherently higher growth rate and a greater allocation of biomass to shoot versus root are associated with resistance of Douglas-fir trees to western spruce budworm defoliation.     

Summer tree roost selection by Rafinesque's big-eared bat

Roost requirements of most North American forest bats are well-documented, but questions remain regarding the ultimate mechanisms underlying roost selection. Hypotheses regarding roost selection include provision of a stable microclimate, space for large colonies, protection from predators, and proximity to foraging habitat, among others. Although several hypotheses have been proposed, specific mechanisms likely vary by species and geographic region. Rafinesque's big-eared bat (Corynorhinus rafinesquii) commonly roosts in trees with large basal hollows in the Coastal Plain of the southeastern United States. Our objective was to weigh evidence for hypotheses regarding selection of diurnal summer roosts by Rafinesque's big-eared bat at 8 study sites across the Coastal Plain of Georgia, USA. We used transect searches and radiotelemetry to locate roosts and measured 22 characteristics of trees, tree cavities, and surrounding vegetation at all occupied roosts and for randomly selected unoccupied trees. We evaluated 10 hypotheses using single-season occupancy models and used Akaike's information criterion to select the most parsimonious models. We located 170 tree roosts containing approximately 870 bats for our analysis. The best supported model predicted bat presence from cavity size, interior wall texture, and number of entrances. Because large cavities allow bats to fly and smooth walls impede attacks by terrestrial predators, our results are consistent with the hypothesis that bats select roosts that allow them to evade predators. However, data on predation rates are needed for a conclusive determination. Because trees suitable as roosts for Rafinesque's big-eared bat are rare in the landscape, protection of suitable forested wetland habitat is essential to provide current and long-term roost tree availability. © 2012 The Wildlife Society.