Activity patterns at the Arctic Circle: nocturnal eagle owls and interspecific interactions during continuous midsummer daylight

Circadian rhythms result from adaptations to biotic and abiotic environmental conditions that cycle through the day, such as light, temperature, or temporal overlap between interacting species. At high latitudes, close to or beyond the polar circles, uninterrupted midsummer daylight may pose a challenge to the circadian rhythms of otherwise nocturnal species, such as eagle owls Bubo bubo. By non‐invasive field methods, we studied eagle owl activity in light of their interactions with their main prey the water vole Arvicola amphibius, and their competitor the white‐tailed eagle Haliaeetus albicilla during continuous midsummer daylight on open, treeless islands in coastal northern Norway. We evaluated circadian rhythms, temporal overlap, exposure, and spatial distribution. The owls maintained a nocturnal activity pattern, possibly because slightly dimmer light around midnight offered favourable hunting conditions. The eagles were active throughout the 24‐h period as opposed to the strictly diurnal rhythm reported elsewhere, thus increasing temporal overlap and the potential for interference competition between the two avian predators. This may indicate an asymmetry, with the owls facing the highest cost of interference competition. The presence of eagles combined with constant daylight in this open landscape may make the owls vulnerable to interspecific aggression, and contrary to the available literature, eagle owls rarely exposed themselves visually during territorial calls, possibly to avoid detection by eagles. We found indications of spatial segregation between owls and eagles reflecting differences in main prey, possibly in combination with habitat‐mediated avoidance. Eagle owl vocal activity peaked in the evening before a nocturnal peak in visual observations, when owls were active hunting, consistent with the hypothesis of a dusk chorus in nocturnal bird species. The owls may have had to trade‐off between calling and foraging during the few hours around midnight when slightly dimmer light reduced the detection risk while also providing better hunting conditions     

Parallel declines in abundance of insects and insectivorous birds in Denmark over 22 years

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Occurrence of bats in northernmost Sweden (65°N) and their feeding ecology in summer

The population density of the northern bat Eptesicus nilssoni at 65° N was about one-fifth of that in southern Sweden (57° N). No other bat species was observed at 65° N. In one maternity colony, young northern bats were born in July and were flying by early August. The diet consisted mainly of small insects, predominantly dipterans, which were captured and eaten in the air. The bats maintained a nocturnal (22:00–02:30 h) foraging schedule in summer, partly reflecting the activity of insects, and spent about three hours away from the day roost each night. Owing to the short periods of darkness, the foraging flights usually started and ended in daylight. Habitat use reflected the occurrence of small insects near vegetation.     

Activity patterns of pipistrelle bats (Pipistrellus pipistrellus) in Oxfordshire

A maternity colony of pipistrelle bats ( Pipisfrellus pipistrellus ), in Oxfordshire, was monitored between 1 March 1989 and 6 October 1989. An infra-red 'automatic bat counter' was installed at the roost, to record the number of bats entering and leaving each minute throughout the night. Air temperature, light intensity at sunset, cloud cover, wind speed and rain were recorded on each night of monitoring. Insect abundance was estimated on 18 nights.
The nightly activity pattern was found to be unimodal in pregnancy, bimodal during lactation and unimodal post-weaning. The mean time that each bat spent outside the roost ranged from 103–483 min, with a mean of 321 min.
Ambient air temperature and length of night were significant factors affecting mean time spent outside the roost. The percentage of the night which the bats spent away from the roost ranged from 22 to 88%, with a mean of 64%. There was a significant positive correlation between ambient air temperature and percentage of the night spent away from the roost. Insect abundance showed no significant correlation with the time that bats spent outside the roost. Wind and rain had no apparent effect on time spent outside the roost.     

Activity patterns of Pipistrelle bats (Pipistrellus pipistrellus) in north-east Scotland

Pipistrellus pipistrellus emerge from their nursery roosts in north-east Scotland about 35 minutes after sunset, at light intensities of between 15 and 35 lux. Cloud cover, windspeed, ambient temperature, rain, light mist and moonlight have no apparent effect on the time or pattern of emergence. Throughout pregnancy and lactation, emergence lasts for about an hour. After weaning, when the adult females have left the roost, their young take about 40 minutes to emerge. The average rate of initial emergence is proportional to colony size, and the maximum rate of emergence occurs half way through the exodus.
During pregnancy in May and June most bats leave the roost once each night soon after dusk and return between midnight and dawn. After parturition in late June the activity pattern becomes bimodal and the numbers of bats outside the roost show peaks after dusk and immediately before dawn. There is intermittent activity in the vicinity of the roost all night and bats make two or three flights each night. After weaning in August the activity pattern gradually ceases to be bimodal, and the number of flights per bat falls to between one and two. The average time spent outside the roost varies between 2–5 and 5 hours during the summer. The recorded activity patterns of night-flying insects are all bimodal, with peaks after dusk and before dawn, corresponding with the maximum number of bats outside the roost during lactation.     

Seasonal patterns of habitat use by insectivorous bats in a subtropical African agro‐ecosystem dominated by macadamia orchards

We report on acoustic surveys of insectivorous bats conducted during seven months of the year using ANABAT recordings in two habitats (macadamia orchards and adjacent riparian bush) in a subtropical agro‐ecosystem in northern South Africa. We defined two functional foraging groups of bats based on their echolocation calls: (i) open‐air foragers (family Molossidae) having narrow‐band, low‐frequency, low duty cycle calls; and (ii) clutter‐edge foragers (families Miniopteridae and Vespertilionidae), having broad‐band, higher frequency, low duty cycle calls. Bat activity (number of bat passes) was not significantly influenced by habitat. Total bat activity and activity of both functional groups varied significantly between seasons, being highest in summer and autumn (coinciding with annual peaks in numbers of Twin spotted (Bathycoelia natalicola) and Green (Nezara spp) Stinkbugs, order Heteroptera, family Pentatomidae, and Macadamia Nut Borer moths, Cryptophlebia ombrodelta) and lower in winter and spring. No significant effect of moon phase was detected, either on total activity or activity of the two functional groups. We postulate that the significant pattern of seasonality of commuting and/or foraging activity of bats in macadamia orchards (which is more marked in open‐air foragers) may be driven by the seasonal abundance of pest insects such as stinkbugs and Macadamia Nut Borer moths.     

Microhabitat associations of terrestrial insectivorous birds in Amazonian rainforest and second‐growth forests

ABSTRACT Across the Neotropics, small‐bodied terrestrial insectivores are sensitive to forest fragmentation and are largely absent from second‐growth forests. Despite their sensitivity to forest structure, the microhabitat relationships of these birds have not been quantified. From July 1994 to January 1995 in central Amazonia, we characterized habitat at sites where nine species of terrestrial insectivores were observed foraging, as well as at randomly selected sites in continuous forest and two types of 10–15‐yr‐old second‐growth forest common in Amazonia (Vismia‐ and Cecropia‐dominated). We used factor analysis to find suites of correlated variables. From each factor, we selected a representative variable that was relatively easy to measure. We used Bayesian analysis to estimate means and standard deviations of these variables for each species and for each type of habitat. All nine focal species were associated with ranges of microhabitat variables, such as leaf litter depth and tree densities, often absent in second‐growth forests. At least in the early stages of regeneration, neither type of second‐growth forest provides suitable structure for the terrestrial insectivores in our study. The large leaves of Cecropia trees that make up the thick leaf litter may preclude the use of Cecropia‐dominated second growth by our focal species, many of which manipulate leaves when foraging. The leaf litter in Vismia‐dominated second growth was also thicker than sites used for foraging by our focal species. In addition, Vismia‐dominated growth had more small trees and small nonwoody vegetation, perhaps impeding movement by terrestrial birds. In continuous forest, our focal species foraged in microhabitats with characteristics that generally overlapped those of randomly selected sites. Thus, our results are consistent with the hypothesis that microhabitat differences make second‐growth forests unsuitable for our focal species.     

Phylogeographical patterns of two closely related desert shrubs,Nitraria roborowskii and N. sphaerocarpa (Nitrariaceae), from arid north‐western China

To investigate the influence of climate aridification and oscillations on the genetic diversity and evolutionary processes of organisms in the Quaternary in north‐western China, we selected Nitraria roborowskii and N. sphaerocarpa and examined the geographical apportionment of genetic variation in their vast range. In the study, 21 plastid haplotypes were identified in N. roborowskii based on two plastid DNA regions. We found significant genetic differentiation between populations in both N. roborowskii and N. sphaerocarpa. We also found similar phylogeographical patterns of allopatric divergence and regional range expansion in both species, but the degree of allopatric divergence in N. roborowskii was lower. Finally, areas to the south of the Alxa Desert, the Hexi Corridor and Ningxia Province were identified as possible refugia for N. roborowskii.     

Standard energetics of leaf-nosed bats (Hipposideridae): its relationship to intermittent- and protracted-foraging tactics in bats and birds

Basal rates of metabolism within the insectivorous genera Hipposideros and Ascelliscus, Old World leaf-nosed bats (Hipposideridae), ranged from 58% to 77% of the mammalian standard. The larger species, Hipposideros diadema and Hipposideros maggietaylori, effectively thermoregulated at ambient temperatures down to 9 degrees C, whereas two smaller species, Hipposideros galeritus and Hipposideros cervinus, occasionally permitted body temperatures to fall below 32 degrees C. The low basal rates of metabolism in hipposiderids correlated with a predatory life-style characterized by intermittent flight from a perch to capture insects, a correlation similar to that found in nonpasserine birds. Intermittent-foraging bats and nonpasserines collectively had basal rates of metabolism that averaged 75% of those that pursue insects during protracted flight. However, no difference in basal rate was found between protracted- and intermittent-foraging passerines, which had basal rates 1.8- and 2.4-times those of protracted-foraging and intermittent-foraging bats and nonpasserines, respectively. Bats, swifts, and caprimulgids that enter torpor have basal rates that are 85% of those of similar species that do not enter torpor. Body mass, order affiliation, foraging mode, and propensity to enter into torpor collectively account for 97% of the variation in basal rate of metabolism in insectivorous bats and birds. Foraging style therefore appears to be a factor contributing to the diversity in endotherm energetics. Minimal thermal conductance in the genus Hipposideros ranged from 75% to 102% of the mammalian standard. Birds have minimal thermal conductances that are 75% of mammals and intermittent foragers have minimal conductances that are 78% of protracted foragers.     

Energetic cost of hovering flight in nectar-feeding bats (Phyllostomidae: Glossophaginae) and its scaling in moths, birds and bats

Three groups of specialist nectar-feeders covering a continuous size range from insects, birds and bats have evolved the ability for hovering flight. Among birds and bats these groups generally comprise small species, suggesting a relationship between hovering ability and size. In this study we established the scaling relationship of hovering power with body mass for nectar-feeding glossophagine bats (Phyllostomidae). Employing both standard and fast-response respirometry, we determined rates of gas exchange in Hylonycteris underwoodi (7 g) and Choeronycteris mexicana (13–18 g) during hover-feeding flights at an artificial flower that served as a respirometric mask to estimate metabolic power input. The O₂ uptake rate (V˙ _o2) in ml g⁻¹ h⁻¹ (and derived power input) was 27.3 (1.12 W or 160 W kg⁻¹) in 7-g Hylonycteris and 27.3 (2.63 W or 160 W kg⁻¹) in 16.5-g Choeronycteris and thus consistent with measurements in 11.9-g Glossophagasoricina (158 W kg⁻¹, Winter 1998). V˙ _o2 at the onset of hovering was also used to estimate power during forward flight, because after a transition from level forward to hovering flight gas exchange rates initially still reflect forward flight rates. V˙ _o2 during short hovering events (<1.5 s) was 19.0 ml g⁻¹ h⁻¹ (1.8 W) in 16-g Choeronycteris, which was not significantly different from a previous, indirect estimate of the cost of level forward flight (2.1 W, Winter and von Helversen 1998). Our estimates suggest that power input during hovering flight P _h (W) increased with body mass M (kg) within 13–18-g Choeronycteris (n = 4) as P _h  = 3544 (±2057 SE) M ^{1.76 (±0.21 SE)} and between different glossophagine bat species (n = 3) as P _h  = 128 (±2.4 SE) M ^{0.95 (±0.034 SE)}. The slopes of three scaling functions for flight power (hovering, level forward flight at intermediate speed and submaximal flight power) indicate that: 1. The relationship between flight power to flight speed may change with body mass in the 6–30-g bats from a J- towards a U-shaped curve. 2. A metabolic constraint (hovering flight power equal maximal flight power) may influence the upper size limit of 30–35 g for this group of flower specialists. Mass-specific power input (W kg⁻¹) during hovering flight appeared constant with regard to body size (for the mass ranges considered), but differed significantly (P < 0.001) between groups. Group means were 393 W kg⁻¹ (sphingid moths), 261 W kg⁻¹ (hummingbirds) and 159 W kg⁻¹ (glossophagine bats). Thus, glossophagine bats expend the least metabolic power per unit of body mass supported during hovering flight. At a metabolic power input of 1.1 W a glossophagine bat can generate the lift forces necessary for balancing 7 g against gravitation, whereas a hummingbird can support 4 g and a sphingid moth only 3 g of body mass with the same amount of metabolic energy. These differences in power input were not fully explained by differences in induced power output estimated from Rankine-Froude momentum-jet theory. Accepted: 10 November 1998     

The patterns of genetic diversity in Carpentaria acuminata (Arecaceae), and rainforest history in northern Australia

Carpentaria acuminata occurs in monsoon rainforest and is endemic to the Northern Territory, Australia. The genetic diversity of C. acuminata populations was surveyed across the geographical range of the species using isozyme analysis. Genetic diversity within C. acuminata populations ( H _E = 0.143) was typical of rainforest species and woody angiosperms generally. Genetic diversity was not correlated with rainforest patch size. However, there was significant heterogeneity among populations ( F _ST = 0.379), with infrequent effective gene flow among populations ( Nm = 0.39). Genetic diversity was negatively correlated with increasing distance between neighbouring C. acuminata populations, but geographical distance was not a good predictor of genetic similarity. C. acuminata is a favoured food of mobile frugivores such as Torres Strait pigeons and flying foxes. The decreased diversity with decreasing density of populations indicated that seed dispersal by frugivores has been important for the maintenance of diversity in this species. Populations known to have originated on relatively young, Holocene landforms were not necessarily genetically depauperate. Gene flow by pollen is apparently limited because C. acuminata populations are significantly inbred regardless of genetic diversity ( F = 0.641). The distribution and diversity of rare alleles, i.e. those occurring in few populations, is consistent with the theory of rainforest contraction during the Pleistocene.     

Spatial and temporal variation in Nothofagus pumilio growth at tree line along its latitudinal range (35°40'–55° S) in the Chilean Andes

Aim To identify the dominant spatial and temporal patterns of Nothofagus pumilio radial growth over its entire latitudinal range in Chile, and to find how these patterns relate to temperature and precipitation variation from instrumental records. Location This study comprises 48 tree line or high elevation N. pumilio sites in the Chilean Andes between 35° 36′ and 55° S. Nothofagus pumilio is a deciduous tree species that dominates the upper tree line of the Chilean and Argentinean Andes in this latitudinal range. Methods At each of the sampled sites, two cores from 15 to 40 living trees were collected using increment borers. Cores were processed, tree rings were measured and cross‐dated, using standard dendrochronological procedures. Radii from nearby sites were grouped into 13 study regions. A composite tree‐ring width chronology was developed for each region in order to capture and integrate the common growth patterns. For the identification of the dominant patterns of growth, as well as temperature and precipitation variation, we used principal components (PCs) analysis. Correlation analysis was used for the study of the relationship of N. pumilio tree‐ring growth with temperature and precipitation records. Results Nothofagus pumilio tree line elevation is 1600 m in the northernmost region and gradually decreases to 400 m in the southernmost region. Despite local differences along the transect, the decrease in tree line elevation is fairly constant, averaging c. 60 m per degree of latitude (111 km). Tree growth at the northernmost regions shows a positive correlation with annual precipitation (PC1‐prec) and negative correlation with mean annual temperature (PC2‐temp), under a Mediterranean‐type climate where water availability is a major limiting factor. Conversely, tree growth is positively correlated with mean annual temperature (PC1‐temp) in the southern portion of the gradient, under a relatively cooler climate with little seasonality in precipitation. Main conclusions Our findings indicate that temperature has a spatially larger control of N. pumilio growth than precipitation, as indicated by a significant (P < 0.05) either positive or negative correlation of tree growth and PC1‐temp and/or PC2‐temp for nine of the 13 regional chronologies (69.2% of the total), whereas precipitation is significantly correlated with only two chronologies (15.4% of the total). Temporal patterns of N. pumilio tree growth reflected in PC1‐growth for the period between 1778 and 1996 indicate an increasing trend with above the mean values after 1963, showing high loadings in the southern part of the gradient. This trend may be explained by a well‐documented increase in temperature in southern Patagonia. Ongoing and future research on N. pumilio growth patterns and their relationship to climate covering the Chilean and Argentinean Andes will improve the understanding of long‐term climate fluctuations of the last three to four centuries, and their relationship to global change at a wide range of spatial and temporal scales.     

Space use and territoriality of wolverines (<Emphasis Type="Italic">Gulo gulo</Emphasis>) in northern Scandinavia

Space use and territoriality influence population structure and dynamics and is therefore an important aspect in understanding the ecology of animals. We investigated spatial and temporal space use of wolverines (Gulo gulo) in northern Scandinavia. We estimated home ranges of 24 radio-marked individuals (17 females and seven males). Male home ranges (mean 669 km²; SE = 211) were significantly larger than female home ranges (mean 170 km²; Wilcoxon–Mann–Whitney; P = 0.001) and encompassed or included parts of up to five different females. Home range sizes of reproducing (170 km²; SE = 51) and barren (171 km²; SE = 63) adult females did not differ. Wolverines in Scandinavia exhibit intrasexual territoriality, with male home ranges totally exclusive and female home ranges either exclusive or with little home range overlap. Overlap between wolverine territories is most likely explained by intrasexual tolerance and kinship.     

Quality of chilled and cold-stored (5 °C) canine spermatozoa submitted to different rapid cooling rates

The aim of this study was to evaluate the sperm quality in chilled canine semen using di?erent cooling rates from room temperature (23 °C) to 5 °C and subsequently cold-stored at 5 °C for up to 96 hours. In experiment 1, semen samples from five dogs were pooled, diluted in Tris-fructose-citrate extender with 20% egg yolk and split into four aliquots that were chilled to 5 °C using di?erent cooling rates of 2.25, 0.9, 0.45, and 0.2 (control) °C/min. In experiment 2, semen from five dogs was processed individually as described above and split into two aliquots that were chilled to 5 °C using rates of either 2.25 °C/min or 0.2 °C/min. In both experiments, the sperm quality (i.e., sperm motility and viability) was evaluated before cooling and after 0, 24, 48, 72, and 96 hours of storage at 5 °C. The total motility, progressive motility, and quality of movement parameters were assessed using computer-assisted analysis system, and the percentage of viable spermatozoa was determined using ?ow cytometry (H-42/PI//FITC-PNA). The cooling rate did not in?uence the sperm quality parameters at any of the evaluation times. All evaluated males showed the same response to chilling semen at a rapid cooling rate. Storage time negatively in?uenced (P < 0.05) sperm motility, regardless of the cooling rate used. In conclusion, canine sperm could be chilled and stored for 96 hours at 5 °C in a Tris-fructose extender with 20% egg yolk using rapid cooling rates, with values for sperm quality similar to those from a conventional protocol.     

Application of methyl jasmonate to grey willow (<Emphasis Type="Italic">Salix cinerea</Emphasis>) attracts insectivorous birds in nature

It has been suggested that insectivorous birds may be guided by herbivore-induced plant volatiles (HIPVs) to herbivore-rich trees with herbivorous damage. The HIPV production in plants is partly mediated by jasmonic acid signalling pathway. Methyl jasmonate (MeJA) was proved to be a suitable agent for induction of HIPVs similar to those induced by herbivorous insects in many plant species. We studied the effects of methyl jasmonate on volatile emission and natural enemy attraction using mature grey willow (Salix cinerea) under natural conditions in Czech Republic. We treated 12 experimental shrubs with 30 mM MeJA and completed the experiment with 12 control shrubs. We monitored attacks by natural predators with artificial plasticine caterpillars which were checked daily. Birds most often pecked the caterpillars exposed on MeJA-treated shrubs and this attractiveness differed significantly from control. Attractiveness of MeJA-treated shrubs did not differ significantly from control shrubs for arthropod predators. Spraying MeJA on grey willows resulted in significantly higher production of α-pinene, β-pinene, 3-carene, limonene and β-ocimene. There was a marginally significant positive correlation between the predation rate by birds and relative change in α-pinene emissions.     

Characterization of dinucleotide microsatellite loci in big brown bats (Eptesicus fuscus), and their use in other North American vespertilionid bats

We describe the development of six microsatellite loci for big brown bats, Eptesicus fuscus. Microsatellite markers were isolated from a small insert genomic library, and tested on a population of 44 animals from the Pend d’Oreille Valley, in southeastern British Columbia, Canada. These six loci were highly variable, with 12–23 alleles per locus, and observed and expected heterozygosities of greater than 79.5%. The six primer sets, and three others that were not variable in E. fuscus, were tested on 11 other species in the families Vespertilionidae and Antrozoidae. All the tested loci amplified highly variable products in at least several other species.     

Food habits of northern (Eptesicus nilssoni) and brown long-eared (Plecotus auritus) bats in Sweden

Food habits of sympatric northern and brown long-eared bats were examined by analysis of droppings collected in six maternity roosts of each species. Relatively large, nocturnal flying insects such as moths (Lepidoptera), June-beetles (Coleoptera: Scarabaeidae), crane-flies (Diptera: Tipulidae) and caddis-flies (Trichoptera), were eaten by both bat species, representing 47% (by volume) of the northern and 57% of the long-eared bat's diet. Small dipterans were important food items for the northern bat (47% of the diet) but not for the long-eared bat, which instead consumed large amounts of diurnal or predominantly non-flying taxa such as blow-flies (Diptera: Calliphoridae), earwigs (Dermaptera), spiders (Araneae) and harvestmen (Opiliones) (40% of the diet). The difference in food selection by the two species is probably due to the different foraging strategies used.