Bats, Chiroptera

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The Ecology of Bats (Chiroptera) of the Shahdagh National  Park of Azerbaijan

The Ecology of Bats (Chiroptera) of the Shahdagh National Park of Azerbaijan

In the Shahdag National park there are Chiroptera of the genus (Rhinolophus), family (Rhinolophidae) species–the lesser horseshoe bat (Rhinolophus hipposideros Bechstein, 1800). It should be recognised that this species are not particularly widespread, it can be found locally in Shamakhi and partlyIsmailli rayons in the landscapes of foothill and mountain forests. This species is very flexible, for several decades it disappeared in Belgium, the Netherlands and northern Germany (with the exception of Bavaria, Thuringia and Saxony), in a large part of Poland, in Switzerland and Austria is remained fragmentary, in Spain most of the colonies have disappeared in connection with the construction of new houses, in the north of France the number is significantly reduced. It is no coincidence that in recent years the number of the less horseshoe bat is reduced not only in our country but also in the world, therefore, as a rare and vanishing species is included in Red Book of some countries, including in Azerbaijan. According to Rahmatulina (1971, 1995, 2005), the greater horseshoe bat Rhinolophus ferrumequinum Schreber, 1774, is very small in number. On the territory of the Shahdag National park, it is confined to the foothills and low mountains, as well as to the plains, where there is suitable shelters for little animals:karst caves, crevices,erosion scars in the river cliffs, natural and artificial catacombs, appropriate human buildings. In the mountains on the territory of Oguz rayon this species is found up to 3200 m above sea level and is considered the most large representative of the horseshoe bats in Europe.
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The Diet Composition of Four Vesper Bats (Chiroptera: Vespertilionidae) from the Centre Region of Cameroon (Central Africa)

The Diet Composition of Four Vesper Bats (Chiroptera: Vespertilionidae) from the Centre Region of Cameroon (Central Africa)

species from the Centre region of Cameroon. The diets of these species have never been studied anywhere in Cameroon before, hence this study provides important baseline information about these species. This new information can be used for both conservation and to improve public awareness about bats in Cameroon. Lepidopterans were an important food source for vesper bats in our study area and further investigations are now required into the quantity and composition of invertebrates consumed by microchiropterans in order to assess their economic and ecological role in the Centre region of Cameroon. We observed large amounts of Diptera content in the diet of S. hirundo, and it is possible it could play a role in the control of certain insect vector such as mosquitoes. Elucidating the role of bats in mosquito vector control could possibly lead to benefits for both the conservation of bats and public health. Equally, the bat diet could help reduce dependence on toxic and dangerous pesticides especially in agriculture. This could be done through the erecting of bat houses to attract bats in farms such that these bats can feed on such pests and thus reduce or minimize the use of pesticides. Conservation effort for the studied species should focus on their foraging and/or roosting areas.
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What ears do for bats: a comparative study of pinna sound pressure transformation in chiroptera

What ears do for bats: a comparative study of pinna sound pressure transformation in chiroptera

and physiological specializations (e.g. Bruns et al. 1976) to detect fluttering insects. These specialisations allow decoding of frequency and amplitude modulations ‘imprinted’ at the rate of the insect wingbeat on the echo returning to the bat (e.g. Kober and Schnitzler, 1990). Using an analogy from the visual system, the hearing world of these bats probably has a slightly ‘monochromatic’ touch, and the characteristics of their pinnae reflect this view. We found an unmistakable ‘tuning’ of the external ears to the CF echolocation call components in Hipposideridae and Rhinolophidae (Fig. 8 D – F ) . The same pattern does not fit all species, however, for some exhibit peak pinna gain, others highest interaural intensity differences and still others steepest IID in the spectral range of the CF part of their echolocation calls. The IID frequency display showed the most consistent specializations, again substantiating the importance of interaural intensity differences for echolocation. However, bats in these families have conspicuous ear movements (e.g. Pye and Roberts, 1970), correlated with sound emissions. These movements seem to be crucial for target localization in the vertical plane (Mogdans et al. 1988). We predict that the movements will affect the IID available to the auditory system. An accurate reconstruction of IIDs at different frequencies and at different positions of the pinnae could provide an interesting comparison. Pteronotus parnellii shows an interesting convergence with hipposiderids and rhinolophids, coinciding with specializations in its auditory pathway, which is tuned to the dominant CF harmonic at 6 0 kHz (Pye, 1980b). The specializations reflect the use of Doppler-shifted echoes to ensure that returning echoes are at the frequency of greatest sensitivity of its auditory system. Our results indicate that the directionality of sound pressure transformation at the external ear and derived qualities such as the interaural intensity difference and pinna gain are also specialized compared with other mormoopids (Fig. 1 2 ) .
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The Detection of Partial Albinism at Three Species of Bats (Mammalia: Chiroptera) in European Part of Russia

The Detection of Partial Albinism at Three Species of Bats (Mammalia: Chiroptera) in European Part of Russia

Bats extremely rare have full or partial albinism [5]. Nowadays, it was registered only in 17 species in Europe [6], whereas worldwide albinism is noticed in 50 species [1] [2] [4] [7]-[14]. It is predominantly animals from Central and Latin America. Each of these species has from one to five individuals who has abnormal coat color that compose about 75 individuals.

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Energetic consequences of flight speeds of foraging red and hoary bats (Lasiurus borealis and Lasiurus cinereus; Chiroptera: Vespertilionidae)

Energetic consequences of flight speeds of foraging red and hoary bats (Lasiurus borealis and Lasiurus cinereus; Chiroptera: Vespertilionidae)

behaviour of moths, a factor influencing the bats’ foraging success (Acharya and Fenton, 1992). Although the rate at which L. cinereus attacked moths was independent of moth density at the lights (Hickey, 1993), two lines of evidence suggest that the insects attracted to the lights were important to the bats we studied. First, marked individuals returned night after night (both species) and season after season (both species, but mainly L. cinereus) to the same lights and concentrated their foraging there (Hickey and Fenton, 1990; Hickey, 1993). Most radio-tagged bats of both species roosted within 500 m of the foraging sites (Hickey and Fenton, 1990; Hickey, 1993). At 500 m, the two-way commuting costs are 58.73 J for L. cinereus and 33.13 J for L. borealis at the V and P values we obtained (Table 1); both values are much lower than the useful energy in a medium-sized moth. Second, monitoring bats using their echolocation calls repeatedly demonstrated that both species were rarely encountered away from the lights and feeding buzzes in unlighted areas were extremely uncommon (Hickey and Fenton, 1990; Hickey, 1993). We predict that bats foraging in lower prey-density situations will fly at speeds closer to predicted V mp and spend more time foraging
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Karyology of eight species of bats (Mammalia: Chiroptera) from Hainan Island, China

Karyology of eight species of bats (Mammalia: Chiroptera) from Hainan Island, China

Twenty-eight specimens of eight species of bats were captured on Hainan Island, China in February 2005 and September 2008 (Table 1). The identification of these bats was based on standard taxonomic pa- pers [18-21]. Voucher specimens are deposited in the College of Life Science, Guangzhou University, Guangzhou. Chromosome preparations were made from primary culture cells of lung and tail tissue us- ing the methods of Harada and Yosida [22]. G- and C-banding differential stainings were applied for se- lected species following the methods of Seabright [23] and Sumner [24], respectively. The nomenclature of chromosomes follows Levan et al. [25]. The diploid number (2n) was determined by observing 30 meta- phase cells in each specimen, and the fundamental number (FN) was defined as the total number of autosomal arms (Table 2).
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Host Switching in Lyssavirus History from the Chiroptera to the Carnivora Orders

Host Switching in Lyssavirus History from the Chiroptera to the Carnivora Orders

lineage (central United States), as was suggested by partial G gene sequencing (data not included). However, a second spill- over in an unknown region spread the RABV worldwide in a variety of carnivores from an unknown vector. It produced an RABV responsible for 32,000 human deaths per year (52a), constituting a historical spillover, and made rabies a public health problem. Spillovers of lyssaviruses from chiropters to other animals, mainly carnivores, may have occurred repeat- edly in history and still occur (10). However, it is not known why only rare spillovers have succeeded in being maintained through time or why others have been extinguished. The present work give strong evidence that what is today known as carnivoran rabies is indeed a result of a few successful episodes of host switching from bats.
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Bats Are an Untapped System for Understanding Microbiome Evolution in Mammals

Bats Are an Untapped System for Understanding Microbiome Evolution in Mammals

ABSTRACT Mammals evolved in a microbial world, and consequently, microbial symbionts have played a role in their evolution. An exciting new subdiscipline of metagenomics considers the ways in which microbes, particularly those found in the gut, have facilitated the ecological and phylogenetic radiation of mammals. How- ever, the vast majority of such studies focus on domestic animals, laboratory models, or charismatic megafauna (e.g., pandas and chimpanzees). The result is a plethora of studies covering few taxa across the mammal tree of life, leaving broad patterns of microbiome function and evolution unclear. Wildlife microbiome research urgently needs a model system in which to test hypotheses about metagenomic involvement in host ecology and evolution. We propose that bats (Order: Chiroptera) represent a model system ideal for comparative microbiome research, affording opportunities to examine host phylogeny, diet, and other natural history characteristics in relation to the evolution of the gut microbiome.
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Vocalizations, feeding and flight behaviour of nectar-feeding bats (Glossophaga soricina and Leptonycteris yerbabuenae)

Vocalizations, feeding and flight behaviour of nectar-feeding bats (Glossophaga soricina and Leptonycteris yerbabuenae)

New World leaf-nosed bats, family Phyllostomidae, are one of the most varied families within the order Chiroptera. The range of dietary specializations of these bats includes carnivory, insectivory, frugivory and nectarivory, speaking to the rich diversity within this family (Datzmann, von Helversen, & Mayer, 2010). Pallas’ long-tongued bat (Glossophaga soricina) and the lesser long-nosed bat (Leptonycteris yerbabuenae, previously sanborni) belong to the subfamily Glossophaginae, a group of bats that primarily feed on nectar and pollen. G. soricina are distributed throughout Central and South America (Barquez, Perez, Miller, & Diaz, 2008) while L. yerbabuenae are migratory bats that range from Central America to the southwest of the United States (Arroyo-Cabrales, Miller, Reid, Cuarón, & de Grammont, 2008). These species are sympatric where their ranges overlap in Mexico and feed from the same flowers of Bombacaceae, Agavacae and Cactacae species (Henry & Stoner, 2011). Some L. yerbabuenae migrate to Arizona in the spring; the females form large maternity roosts (10,000 to >100,000) to give birth and care for young while the males live separately in small groups. Females, juveniles and males migrate back south in the fall (Cole & Wilson, 2006).
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EMERGING VIRUS DISEASES: CAN WE EVER EXPECT  THE UNEXPECTED?

EMERGING VIRUS DISEASES: CAN WE EVER EXPECT THE UNEXPECTED?

Given the expanding confirmation of bats as stores of developing infections,22 it merits considering the advancement and assorted qualities of these warm blooded animals. About 1000 species are disseminated all through the world, with the larger part in territories near the equator where sustenance sources are generally copious. Having a place with the mammalian request Chiroptera, bats are extensively distinct into the Old World natural product eating bats (180 species, suborder Megachiroptera) and the microbats—approximately 800 species gathered into 17 families inside the suborder Microchiroptera. Insectivorous bats are on the whole microbats. Bats developed around 50 million years back, with the organic product bats advancing along an altogether different way to the bug eating species. Bats are found in most earthly natural surroundings, with species dissemination differing broadly, some being confined to a solitary island, others being found crosswise over mainlands. Among the last is Miniopterus schreibersii from which Negredo et al.23 detached Lloviu infection from a collapse north-eastern Spain: Schreiber's bats are found all through southern Europe, as far south as South Africa, and as far east as Japan.
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Tölch, Ulf
  

(2006):


	Bat Time Stories: Decision-making in spatio-temporally predictable environments.


Dissertation, LMU München: Fakultät für Biologie

Tölch, Ulf (2006): Bat Time Stories: Decision-making in spatio-temporally predictable environments. Dissertation, LMU München: Fakultät für Biologie

Foraging on nectar: A flower visiting bats ability to perceive differences in nectar volume Kuss et al. (2005). This recently developed algorithm will estimate the psychometric function and three important parameters of this function with their confidence intervals. The first parameter (threshold parameter) is the point at which the investigated subject can distinguish between the two stimuli and is denoted by the point on the psychometric function with the steepest slope. The second parameter is the slope at this point, which gives a measure for the reliability of sensory performance (Treutwein & Strassburger 1999). The third parameter is the lapse rate, which is inferred from the difference between perfect performance and the actual behaviour of animals at high stimulus intensities. It serves as a measure for the errors that are not of perceptional nature but are made due to lapses in attention or motivational problems. Markov Chain Monte Carlo (MCMC) sampling is applied for an estimation of these parameters. In this Bayesian approach to find the parameter estimates, the investigator has to state his/her prior beliefs about the parameter location in form of prior distributions. As prior functions we chose a beta distribution (2,50) for the lapse rate, normally distributed priors for the threshold, and the slope with a mean of 0 and a standard deviation of 1. For the MCMC sampling we performed 5000 runs with 50 leapfrog steps each. The leapfrog step size for the three parameters were: 0.05 (lapse rate), 0.08 (threshold location), 0.1 (slope) (the acceptance rate was at approximately 82%).
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Does sex matter? Gender specific responses to forest fragmentation in Neotropical bats

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

phyllostomid species at the BDFFP, were higher for females than males during the dry season at edges and matrix sites and, to a lesser extent, in continuous forest and fragment interiors. During the reproductive period, female bats face higher energetic demands than males (Barclay 1991), and for C. perspicillata, the most prominent pregnancy peak occurs during the dry season, whereas lactation peaks during the wet season (Bernard 2002, Durant et al. 2013). To compensate for increased energetic demands, females might forage preferentially in the most resource-rich areas (Barclay 1991, Encarnac ß~ ao et al. 2005), espe- cially in the dry season during which fruit availability is lower (Ramos Pereira et al. 2010). Similar to other studies across the species’ distribution (e.g., Mello et al. 2004, Durant et al. 2013), we have identified a second reproductive peak, of somewhat weaker intensity, in the wet season. Since this second peak takes place in the season of highest fruit availability (Ramos Pereira et al. 2010), it is unlikely to affect male–female ratios as much as the more pronounced reproductive peak in the dry season. Early succes- sional gap species of the genus Piper, the preferred food resource of C. perspicillata (Horsley et al. 2015) produce 2–10 times more fruits than shade tolerant or late successional forest species (Thies
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Of the 12,746 observations of individual Flying Foxes throughout the study area, 62.3% of bats (7,940 observed individuals) were noted to roost in rain trees (Albizia saman), 15.3% (1,950) in cluster fig trees (Ficus glomerata), 4.7% (610) in durian trees (Durio zibethinus), 3.5% (445) in Burmese ironwood trees (Xylia dolabriformis), 3% (380) in Para rubber trees (Hevea brasiliensis), 2.4% (300) in jackfruit trees (Artocarpus heterophyllus), 2.3% (290) in Shorea sp., 2% (250) in banyan trees (Ficus benghalensis), 1.9% (240) in beleric myrobolan trees (Terminalia bellerica), 1.6% (210) in betel nut palms (Areca catechu), and 1% (131) in santol trees (Sandoricum koetjape). In respect of the last species, however, unconfirmed reports from local people state that significant numbers of Flying Foxes roost in santol trees in Shaw-taw-maw and on The-byu island. To determine whether the bats’ roost tree preferences are similar to the percentages shown, it would be helpful to undertake an assessment of the available roosting space in each tree species within the study area.
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Update on rabies

Update on rabies

Abstract: Human rabies is almost invariably fatal, and globally it remains an important public health problem. Our knowledge of rabies pathogenesis has been learned mainly from studies performed in experimental animal models, and a number of unresolved issues remain. In contrast with the neural pathway of spread, there is still no credible evidence that hematogenous spread of rabies virus to the central nervous system plays a significant role in rabies pathogenesis. Although neuronal dysfunction has been thought to explain the neurological disease in rabies, recent evidence indicates that structural changes involving neuronal processes may explain the severe clinical disease and fatal outcome. Endemic dog rabies results in an ongoing risk to humans in many resource-limited and resource-poor countries, whereas rabies in wildlife is important in North America and Europe. In human cases in North America, transmission from bats is most common, but there is usually no history of a bat bite and there may be no history of contact with bats. Physicians may not recognize typical features of rabies in North America and Europe. Laboratory diagnostic evaluation for rabies includes rabies serology plus skin biopsy, cerebrospinal fluid, and saliva specimens for rabies virus antigen and/or RNA detection. Methods of postexposure rabies prophylaxis, including wound cleansing and administration of rabies vaccine and human rabies immune globulin, are highly effective after recognized exposure. Although there have been rare survivors of human rabies, no effective therapy is presently available. Therapeutic coma (midazolam and phenobarbital), ketamine, and antiviral therapies (known as the “Milwaukee protocol”) were given to a rabies survivor, but this therapy was likely not directly responsible for the favorable outcome. New therapeutic approaches for human rabies need to be developed. A better understanding of basic mechanisms involved in rabies pathogenesis may be helpful in the development of potential new therapies for the future.
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Foraging in a complex naturalistic environment: capacity of spatial working memory in flower bats

Foraging in a complex naturalistic environment: capacity of spatial working memory in flower bats

If bats did remember feeders visited then it might be expected that feeders emptied recently are better remembered and more successfully avoided than those feeders emptied longer ago (recency effect). After a visit to a feeder a bat has n opportunities during the next n visits of a trial to revisit this particular feeder. Accordingly, there are n–1 opportunities to return two or more visits later to an emptied feeder, and n–2 opportunities to return three or more visits later during each sequence of n visits following the first visit and so on. We calculated the number of opportunities that each bat had for each length of inter-visit interval from the number of visits made during each trial. We then summed the number of revisits and the number of opportunities over blocks of five inter-visit interval lengths and calculated the percentage of opportunities taken by each bat for each block of intervisit intervals (Shettleworth and Krebs, 1982). We performed these calculations separately for the initial trials 1–3 and the later trials 11 to 20 (Fig.·8A). A consequence of the initial win-stay strategy was a strong tendency in trials 1–3 to revisit feeders that had recently been emptied. So there were many revisits during short intervisit intervals during those trials. By trials 11–20, this win-stay effect was no longer detectable. Surprisingly, however, was the flat continuation of the curve in Fig.·8A. In general, recent events are often remembered better than events from longer ago, the so-called recency effect (Shettleworth, 1998). For our data, we had expected that for visits to feeders that had occurred longer ago (long inter-visit interval) bats would eventually show disproportionately higher rates of revisitation errors, indicating that they were forgetting their initial visit. However, the expected increase in the number of errors after long intervisit intervals is not apparent in the data (Fig.·8A). This differs from the findings of other authors performing similar experiments with other organisms (Fig.·8B, Discussion).
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Volitional control of social vocalisations and vocal usage learning in bats

Volitional control of social vocalisations and vocal usage learning in bats

Bats are gregarious, highly vocal animals that possess a broad repertoire of social vocalisations. For in-depth studies of their vocal behaviours, including vocal flexibility and vocal learning, it is necessary to gather repeatable evidence from controlled laboratory experiments on isolated individuals. However, such studies are rare for one simple reason: eliciting social calls in isolation and under operant control is challenging and has rarely been achieved. To overcome this limitation, we designed an automated setup that allows conditioning of social vocalisations in a new context and tracks spectro-temporal changes in the recorded calls over time. Using this setup, we were able to reliably evoke social calls from temporarily isolated lesser spear-nosed bats (Phyllostomus discolor). When we adjusted the call criteria that could result in a food reward, bats responded by adjusting temporal and spectral call parameters. This was achieved without the help of an auditory template or social context to direct the bats. Our results demonstrate vocal flexibility and vocal usage learning in bats. Our setup provides a new paradigm that allows the controlled study of the production and learning of social vocalisations in isolated bats, overcoming limitations that have, until now, prevented in-depth studies of these behaviours.
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Comparative Analyses of Vertebrate Gut Microbiomes Reveal Convergence between Birds and Bats

Comparative Analyses of Vertebrate Gut Microbiomes Reveal Convergence between Birds and Bats

Birds have the smallest genomes of all amniotes, and birds and bats both have more compact genomes than their flightless relatives, most likely due to extensive DNA loss— hypothesized to be related to powerful ongoing selection to reduce mass and enhance flight-related correlates of metabolism (31, 32). It is therefore plausible that pressure to decrease mass may extend to microbial biomass. Both birds and bats also have greater rates of intestinal paracellular absorption than nonflying vertebrates (28, 29), meaning that a higher proportion of simple nutrients are absorbed directly by the host, potentially decreasing the role for symbiotic microbial metabolism. Indeed, some studies of microbial biomass across animal hosts have reported that both birds and bats carry much lower numbers of microbial cells in feces than nonflying mammals (33). A selection toward microbial reduction may also explain in part why birds and bats both seem to have lost an association with Bacteroidetes but retained an association with Proteobacteria, a group of bacteria suggested to have high functional variability (34): to maximize microbial function while also reducing diversity and mass. Even if selective pressure to reduce overall microbial biomass did not, in itself, lead to the loss of tight evolutionary ties with specific symbionts, consistently lower resident microbial biomass might increase the proportion of transient environmental microbes relative to the total pool of microbes in the sample, decreasing the observed degree of host specificity on average. Indeed, this could also help explain the dominance of Proteobacteria, which make up a large proportion of the airborne microbiome (35), a source to which flighted animals are constantly exposed.
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Detection of Latin American Strains of Histoplasma in a Murine Model by Use of a Commercially Available Antigen Test

Detection of Latin American Strains of Histoplasma in a Murine Model by Use of a Commercially Available Antigen Test

Human infection with RFLP class 6 isolates has been de- scribed, including in AIDS patients with disseminated his- toplasmosis from Brazil (6) and Panama (8). Although histo- pathology, quantitative culture, and seropositivity for anti-H. capsulatum antibodies were included in these reports, His- toplasma antigen testing was not. Our experience with negative HAg testing in the infected zoo bats and the evolving knowl- edge that Histoplasma likely consists of at least 7 phylogenetic clades or species (7) have raised concern that HAg testing might only be effective in the detection of North American class 2 Histoplasma.
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Further Evidence for Bats as the Evolutionary Source of Middle East Respiratory Syndrome Coronavirus

Further Evidence for Bats as the Evolutionary Source of Middle East Respiratory Syndrome Coronavirus

What is less clear is whether this recombination occurred in bats or an intermediate host. Lineage 2c strains that use DPP4 have been reported in bats (25, 26), and there is also evidence of positive selection in the bat DPP4 that would indicate the existence of a large diversity of (as-yet-unknown) DPP4-competent strains (39). Just as detailed metagenomics studies have revealed the presence of several severe acute respiratory syndrome (SARS)-like bat CoVs that can use the human angiotensin converting enzyme 2 receptor and/or replicate efficiently in human cells (23, 24, 40–42), it seems likely that subsets of diverse MERS-CoV-like bat coronaviruses will also exist which are prepro- grammed to efficiently use the human DPP4 receptor. This would support the hypoth- esis that the recombination occurred in bats; however, the MERS-CoV spike seems to have adapted and acquired a preference for human DPP4 over the bat homologue (26, 43) making it difficult to conclude with certainty that the MERS-CoV spike has bat origins. Increased surveillance will be required to understand the full diversity of spike phenotypes circulating in bats or in intermediate hosts such as camels.
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Tight coordination of aerial flight maneuvers and sonar call production in insectivorous bats

Tight coordination of aerial flight maneuvers and sonar call production in insectivorous bats

We recorded 94 animal flight trials for each bat over a period of four consecutive days. Two ultrasound-sensitive microphones (UltraSound Advice, SM2 microphone with SP2 amplifier) were used to record the wideband sonar emissions (band-passed between the frequencies of 10 and 100 kHz, Wavetek-Rockland Dual Hi/Lo Filter) and recorded digitally (National Instruments PCI-6122, sampling rate 250 kHz). A set of 10 motion- tracking cameras (Vicon MX T40) tracked reflective markers placed on the bats at 300 Hz. Motion-tracking data were downloaded to a computer running Vicon Nexus software and exported to MATLAB (MathWorks) for further analysis. A set of markers, reflective tape cut into 6.35 mm diameter circles, were attached to the wings of the bats. On each wing, markers were placed at the thumb joint, the base of the wing near digit five, and at the wing tip on both the dorsal and ventral sides (Fig. 1B). Markers were fixed to the bat using tape adhesive and were replaced each day they fell off. A hemispherical marker was also attached to the body but was not reliably recorded by the motion-tracking system and was not used in analysis. Data synchronization was achieved using a trigger switch that broadcast a transistor – transistor logic (TTL) signal to each system. Each system was configured with an 8 s rolling buffer aligned to the onset of the TTL pulse. We did not analyze a set of trials in which there were tracking problems, reducing the total number of trials from 94 for each bat to 86, 85 and 78 for bats 1, 2 and 3, respectively. Trials in which the bats did not fly directly between the first net opening and the second net opening (indirect flight) were excluded from analysis. These trials were excluded if the time to cross the nets was larger than 1 s.d. above the median time and if the flight speed was below 2 s.d. of the median speed. Direct flights comprised 81.4%,
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