Genus Rhinolophus Research Articles

Complete mitochondrial genome of the big-eared horseshoe bat Rhinolophus macrotis (Chiroptera, Rhinolophidae)

We sequenced and characterized the complete mitochondrial genome of the big-eared horseshoe bat, Rhinolophus macrotis. Total length of the mitogenome is 16,848 bp, with a base composition of 31.2% A, 25.3% T, 28.8% C and 14.7% G. The mitogenome consists of 13 protein-coding genes, 2 rRNA (12S and 16S rRNA) genes, 22 tRNA genes and 1 control region. It has the same gene arrangement pattern as those of typical vertebrate mitochondrial genome. The results will contribute to our understanding of the taxonomic status and evolution in the genus Rhinolophus bats.

The Divergence of Echolocation Frequency in Horseshoe Bats: Moth Hearing, Body Size or Habitat?

A phylogenetic approach was used to test three hypotheses regarding the evolution of diversity in the echolocation frequencies used by horseshoe bats (family Rhinolophidae, genus Rhinolophus): 1) Allotonic Frequency Hypothesis (high frequency echolocation in the Rhinolophidae resulted from coevolution with moth hearing); 2) Allometry Hypothesis (echolocation frequency is negatively scaled with body size and evolutionary changes in echolocation frequencies are correlated with changes in body size in the Rhinolophidae); and 3) Foraging Habitat Hypothesis (evolution of echolocation frequency is associated with changes in habitat type). Both discrete and continuous character sets were used for ancestral state reconstructions and for investigating patterns of evolution between frequency and body size, and frequency and habitat type. Contrary to the prediction of the Allotonic Frequency Hypothesis, echolocation frequency in the Rhinolophidae did not increase over time, which would be expected if moth hearing and bat echolocation frequency coevolved. The number of extant species that exhibit calls within moth hearing ranges was not significantly different from the number of species that echolocate outside of moth hearing range. There was also no correlation between changes in frequency and changes in habitat type as predicted by the Foraging Habitat Hypothesis. Instead, the evolution of echolocation frequency within the Rhinolophidae was correlated with changes in body size as predicted by the Allometry Hypothesis.

A New Species of Rhinolophus (Chiroptera: Rhinolophidae) from China

A new species of the genus Rhinolophus is described from Yunnan Province, southwestern China. The new taxon belongs to the Rhinolophus "philippinensis-group" and is distinguished by differences in the nose-leaf structures, craniodental characteristics, and bacular features.

Karyotypic Variation in Rhinolophid and Hipposiderid Bats (Chiroptera: Rhinolophidae, Hipposideridae)

According to current phylogenetic hypotheses, the bats of the families Rhinolophidae and Hipposideridae are sister groups nested within the clade of Pteropodiformes. A conservative nature of karyotypic evolution was previously reported within the two families. Karyotypes with diploid number (2n) varying between 58 and 62 chromosomes were assumed to prevail among the rhinolophid species, whereas, karyotypes with 32 chromosomes were found in most of the hipposiderid bats. However, divergent lower or higher 2n numbers have been recorded in some species in both families. Variation is documented in the present paper by examination of non-differentially stained karyotypes in 10 species belonging to genera Rhinolophus and Hipposideros from western Africa and southern India. Among the species studied, the karyotypes with a 2n of 32, 36, 52, 56, 58, and 62, and with relatively stable number of autosomal arms (FNa = 60, 62, 64) were recorded.

Perch-hunting in insectivorous Rhinolophus bats is related to the high energy costs of manoeuvring in flight

Foraging behaviour of bats is supposedly largely influenced by the high costs of flapping flight. Yet our understanding of flight energetics focuses mostly on continuous horizontal forward flight at intermediate speeds. Many bats, however, perform manoeuvring flights at suboptimal speeds when foraging. For example, members of the genus Rhinolophus hunt insects during short sallying flights from a perch. Such flights include many descents and ascents below minimum power speed and are therefore considered energetically more expensive than flying at intermediate speed. To test this idea, we quantified the energy costs of short manoeuvring flights (<2 min) using the Na-bicarbonate technique in two Rhinolophus species that differ in body mass but have similar wing shapes. First, we hypothesized that, similar to birds, energy costs of short flights should be higher than predicted by an equation derived for bats at intermediate speeds. Second, we predicted that R. mehelyi encounters higher flight costs than R. euryale, because of its higher wing loading. Although wing loading of R. mehelyi was only 20% larger than that of R. euryale, its flight costs (2.61 ± 0.75 W; mean ± 1 SD) exceeded that of R. euryale (1.71 ± 0.37 W) by 50%. Measured flight costs were higher than predicted for R. mehelyi, but not for R. euryale. We conclude that R. mehelyi face elevated energy costs during short manoeuvring flights due to high wing loading and thus may optimize foraging efficiency by energy-conserving perch-hunting.

Molecular phylogenetics and historical biogeography of Rhinolophus bats

The phylogenetic relationships within the horseshoe bats (genus Rhinolophus) are poorly resolved, particularly at deeper levels within the tree. We present a better-resolved phylogenetic hypothesis for 30 rhinolophid species based on parsimony and Bayesian analyses of the mitochondrial cytochrome b gene and three nuclear introns (TG, THY and PRKC1). Strong support was found for the existence of two geographic clades within the monophyletic Rhinolophidae: an African group and an Oriental assemblage. The relaxed Bayesian clock method indicated that the two rhinolophid clades diverged approximately 35 million years ago and results from Dispersal Vicariance (DIVA) analysis suggest that the horseshoe bats arose in Asia and subsequently dispersed into Europe and Africa.

A New Species from Southwestern China in the Afro-Palearctic Lineage of the Horseshoe Bats (Rhinolophus)

A new species of horseshoe bat (Chiroptera: Rhinolophidae) is described from southwestern China. The presence of a wedge-shaped sella and pointed connecting process of the nose leaf aligns the new species to the landeri group in the Afro-Palearctic lineage of Rhinolophus. However, the new species is distinctly separable from these allopatrically distributed species by its noticeably larger body size. Other sympatric large-sized species of Rhinolophus have rounded connecting processes. Molecular systematic analyses based on mitochondrial cytochrome-b sequences confirmed the affinity of the new species to the Afro-Palearctic lineage, but in a clade most closely related to the ferrumequinum, fumigatus, and maclaudi groups. Of these species, only R. ferrumequinum ranges into Asia and overlaps in distribution with the new species. R. ferrumequinum is similar in general body size and external appearance; however, the new species is distinct in the characteristics of the nose leaf, skull, and baculum. The presence of a new species from southwestern China in the Afro-Palearctic lineage indicates a more complex historical biogeographic scenario within Rhinolophus than previously known. The difficulties found in allocating the new species to one of the phenetically described traditional species groups stress the convenience of using a phylogenetically based systematic organization of the genus Rhinolophus.

A New Species of Horseshoe Bat of the Genus Rhinolophus from China (Chiroptera: Rhinolophidae)

A new species of the Rhinolophus philippinensis group (Chiroptera: Rhinolophidae) is described from Guangdong, Guangxi, and Jiangxi Provinces in China. Rhinolophus huananus n. sp. is characterized by the horseshoe, as well as by external and cranial characteristics that separate it at the species level from the other members of the philippinensis group. One of the small species of the philippinensis group, R. huananus is intermediate in size between smaller R. siamensis and larger R. macrotis.

Bats, civets and the emergence of SARS.

Severe acute respiratory syndrome (SARS) was the first pandemic transmissible disease of previously unknown aetiology in the twenty-first century. Early epidemiologic investigations suggested an animal origin for SARS-CoV. Virological and serological studies indicated that masked palm civets ( Paguma larvata ), together with two other wildlife animals, sampled from a live animal market were infected with SARS-CoV or a closely related virus. Recently, horseshoe bats in the genus Rhinolophus have been identified as natural reservoir of SARS-like coronaviruses. Here, we review studies by different groups demonstrating that SARS-CoV succeeded in spillover from a wildlife reservoir (probably bats) to human population via an intermediate host(s) and that rapid virus evolution played a key role in the adaptation of SARS-CoVs in at least two nonreservoir species within a short period.

WING MEASUREMENT VARIATIONS IN THE FIVE EUROPEAN HORSESHOE BAT SPECIES (CHIROPTERA: RHINOLOPHIDAE)

Wing morphology is crucial for flight performance and foraging ecology in bats. We describe variations in 5 wing parameters within the 5 species of European horseshoe bats (genus Rhinolophus) based on data taken from 3,081 adult individuals. All 5 species belong to a single ecological guild. Measurements were taken from live bats in the field in southeastern Europe (Bulgaria, Greece, and Turkey), where all 5 species occur in sympatry. Examination of our data shows that the species and accordingly their wings differ substantially in size. Albeit grossly similar in form, we additionally found several size-independent differences in wing shape. For example, the smallest species, Rhinolophus hipposideros, and to a lesser extent also R. blasii, have extremely short hand wings, enabling highly maneuverable search flight close to vegetation. The largest species, R. ferrumequinum, and the 2nd largest one, R. mehelyi, have rather long hand wings, allowing fast and economic commuting flight over longer distances. We argue that both size and shape are likely to play a role for niche separation between species. We found both sexual and geographic variation within species. There was sexual dimorphism for most parameters, with females being larger than males. Populations of R. mehelyi in southeastern Europe had significant variation in wing measurements. This was not so for R. ferrumequinum and R. euryale. We give a discriminant function based on only 2 parameters that correctly assigned 98% of the 3,081 individuals to species. This function may prove useful for identification of museum specimens.

Bats and human emerging diseases

Recently, two independent research teams reported evidence of infection with SARS-like coronaviruses in insectivorous horseshoe bats (genus Rhinolophus) in China [1, 2]. SARS emerged in China in 2002, and eventually infected over 8000 people around the world, killing about 10% of them. Early epidemiological studies suggested that the human disease may have originated in Chinese live-animal food markets, and the hunt for a source of the virus quickly identified apparently healthy Himalayan palm civets (Paguma larvata) as prime suspects [3]. Other studies, however, failed to find any evidence of widespread infection in civets [4] or a variety of other animals traditionally used as food in China. Rather, palm civets, like human beings, seem to be only accidental hosts of the virus, becoming infected, again like people, when mixed with other species in markets. More structured hunts for the reservoir continued, although not before thousands of palm civets had been slaughtered.

Linking Bats to Emerging Diseases

A sars-like coronavirus found in 13 of 46 horseshoe bats (genus Rhinolophus ) (“Bats are natural reservoirs of SARS-like coronaviruses,” W. Li et al. , 28 Oct. 2005, p. [676][1]) led A. P. Dobson to conclude that these bats “have now been officially recorded as the natural reservoir host of

Review of Bats and SARS

Bats have been identified as a natural reservoir for an increasing number of emerging zoonotic viruses, including henipaviruses and variants of rabies viruses. Recently, we and another group independently identified several horseshoe bat species (genus Rhinolophus) as the reservoir host for a large number of viruses that have a close genetic relationship with the coronavirus associated with severe acute respiratory syndrome (SARS). Our current research focused on the identification of the reservoir species for the progenitor virus of the SARS coronaviruses responsible for outbreaks during 2002-2003 and 2003-2004. In addition to SARS-like coronaviruses, many other novel bat coronaviruses, which belong to groups 1 and 2 of the 3 existing coronavirus groups, have been detected by PCR. The discovery of bat SARS-like coronaviruses and the great genetic diversity of coronaviruses in bats have shed new light on the origin and transmission of SARS coronaviruses.

IDENTIFYING BATS FROM TIME-EXPANDED RECORDINGS OF SEARCH CALLS: COMPARING CLASSIFICATION METHODS

Recording ultrasonic echolocation calls of bats using bat-detectors is often used for wide-scale monitoring in studies on bat management and conservation. In Europe, the most important legal instrument for bat conservation is the Habitat Directive (43/92/EEC), which defines various levels of species (and habitat) protection for different bat species and/or genera. Thus for most management needs, the usefulness of bat-monitoring techniques depends on the possibility to determine to species/genus. We compared the discrimination performances of 4 statistical methods applied to identify bat species from their ultrasonic echolocation calls. In 3 different areas of northern Italy, we made recordings of 20 species of bat (60% of those occurring in Italy), 17 from the family Vespertilionidae and 3 from Rhinolophidae. Calls of bats identified to species level from morphological and genetic characters were time-expanded and recorded on release. We measured 7 variables from each call, and we developed classification models through both conventional tests (multiple discriminant analysis and cluster analysis) that were based on a classical statistical approach, and through 2 nonconventional classifiers (classification and regression trees, and neural networks) that relied on generalization and fuzzy reasoning. We compared the performance of the 4 techniques using the percentage of cases classified correctly in 5 classification trials at various taxonomic levels that were characterized by an increasingly difficult identification task: (1) family level (Rhinolophidae vs. Vespertilionidae), (2) species level within genus Rhinolophus, (3) genus level within Vespertilionidae, (4) species level within genus Myotis, and (5) all species. Multiple discriminant function analysis (DFA) correctly classified marginally more cases than artificial neural networks (ANN; 74–100% against 64–100%), especially at the species level (trial 4, species of genus Myotis; trial 5, all species). Both these techniques performed better than cluster analysis or classification and regression trees, the latter reaching only 56 and 41% in Myotis species and all species trials. Artificial neural networks do not yet seem to offer a major advantage over conventional multivariate methods (e.g., DFA) for identifying bat species from their ultrasonic echolocation calls.

Ground gleaning in horseshoe bats: comparative evidence from Rhinolophus blasii, R. euryale and R. mehelyi

The 71 species of horseshoe bat (genus Rhinolophus) use echolocation calls with long constant-frequency (CF) components to detect and localize fluttering insects which they seize in aerial captures or glean from foliage. Here we describe ground-gleaning as an additional prey-capture strategy for horseshoe bats. This study presents the first record and experimental evidence for ground-gleaning in the little-studied Blasius’ horseshoe bat (Rhinolophus blasii). The gleaning bouts in a flight tent included landing, quadrupedal walking and take-off from the ground. The bats emitted echolocation calls continuously during all phases of prey capture. Both spontaneously and in a choice experiment, all six individuals attacked only fluttering insects and never motionless prey. These data suggest that R. blasii performs ground-gleaning largely by relying on the same prey-detection strategy and echolocation behaviour that it and other horseshoe bats use for aerial hawking.

Remarks on some types of the genus Rhinolophus (Mammalia, Chiroptera)

Detailed morphological investigation of the type specimens of Rhinolophuspusillus, R. lepidus shortridgei, R. borneensis, R. affinis, R. thomasi septentrionalis and R. philippinensis mon- tanus have resulted in designation of lectotype specimens, taxonomical revisions and better under- standing of specific characters. With 19 figures.

Description of a New Species of Rhinolophus (Chiroptera: Rhinolophidae) from Malaysia

A new species of the genus Rhinolophus is described from Cameron Highlands, Peninsular Malaysia. The diagnostic characters are the shape of the noseleaf and external and cranial measurements. The new taxon belongs to the pusillus-group and based on the morphology of the posterior noseleaf, its closest relative is R. subbadius .

Origins of Horseshoe Bats ( Rhinolophus , Rhinolophidae) in Southern Africa: Evidence from Allozyme Variability

Morphological analyses indicate that horsehose bats in the genus Rhinolophus constitute a monophyletic group which most likely originated in southeastern Asia but which presently inhabits Oriental, Australian, Palaearctic, and Ethiopian zoogeographical provinces. Ten species occur in southern Africa, but it is uncertain which species represent dispersals from Eurasia through North Africa and which have resulted from speciation in Africa. Analyses of 34 allozyme encoding loci in these 10 species and in 2 southern African species of leafnose bats in the sister genus Hipposideros reveal the presence of at least three lineages of Rhinolophus in southern Africa. One lineage includes R. clivosus, R. darlingi, R. fumigatus, and R. hildebrandtii, all of which, except R. clivosus, are endemic to sub-Saharan Africa. Rhinolophus blasii is genetically allied with, but distinct from this group, and appears to be a recent migrant from another lineage centered on the Mediterranean. A third lineage, including at least R. capensis, R. denti, R. simulator, and R. swinnyi, is endemic to sub-Saharan Africa. The phylogenetic position of R. landeri is uncertain, most likely because of the small sample size used to estimate allelic frequencies for this species. The biochemical genetic definitions of these lineages largely agree with previous morphological analyses of Rhinolophus species. Divergences between species within two lineages (R. clivosus, R. darlingi, R. fumigatus, and R. hildebrandtii; and R. capensis, R. denti, R. simulator, and R. swinnyi) appear to reflect two bursts of speciation in the Plio-Pleistocene period within Africa.

Phased-array beam scanning as a possible aid to horizontal localization in horseshoe bats

It has recently been shown that oscillating ear movements in horseshoe bats (genus Rhinolophus) are necessary for localization of targets in the vertical dimension. Target elevation thus seems to be calculated from changes in interaural intensity ratios caused by ear movements during echo reception. This letter suggests that, if the radiated beam were spatially oscillated in a similar manner, target azimuth could be deduced from the corresponding oscillation in echo intensity. The potential for rapid phased-array beam steering exists in horseshoe bats because they emit their echolocation pulses through the nostrils.

Resource partitioning of sonar frequency bands in rhinolophoid bats.

In the Constant Frequency portions of the orientation calls of various Rhinolophus and Hipposideros species, the frequency with the strongest amplitude was studied comparatively. (1) In the five European species of the genus Rhinolophus call frequencies are either species-specific (R. ferrumequinum, R. blasii and R. euryale) or they overlap (R. hipposideros and R. mehelyi). The call frequency distributions are approximately 5-9 kHz wide, thus their ranges spead less than ±5% from the mean (Fig. 1). Frequency distributions are considerably narrower within smaller geographic areas. (2) As in other bat groups, call frequencies of the Rhinolophoidea are negatively correlated with body size (Fig. 3). Regression lines for the genera Rhinolophus and Rhinolophus, species from dryer climates have on the average higher call frequencies than species from tropical rain forests. (4) The Krau Game Reserve, a still largely intact rain forest area in Malaysia, harbours at least 12 syntopic Rhinolophus and Hipposiderso species. Their call frequencies lie between 40 and 200 kHz (Fig. 2). Distribution over the available frequency range is significantly more even than could be expected from chance alone. Two different null hypotheses to test for random character distribution were derived from frequency-size-relations and by sampling species assemblages from a species pool (Monte Carlo method); both were rejected. In particular, call frequencies lying close together are avoided (Figs. 4, 5). Conversely, the distribution of size ratios complied with a corresponding null hypothesis. This even distribution may be a consequence of resource partitioning with respect to prey type. Alternatively, the importance of these calls as social signals (e.g. recognition of conspecifics) might have necessitated a communication channel partitioning.