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List of Specimens
- Ara Monadjem, Peter John Taylor, Fenton (Woody) Cotterill, M. Corrie Schoeman, University of KwaZulu-Natal, South Africa
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Family Miniopteridae Long-fingered Bats
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Summary
The Miniopteridae were previously treated as a subfamily of the Vespertilionidae. However, this family differs in a number of morphological, embryological, immunological, and genetic ways from the Vespertilionidae (Hoofer and van den Bussche 2003, Eick et al. 2005). For example, the best-known character of this group is the uniquely elongated second phalanx of the third digit, which allows the wing to ‘bend’ back onto itself, hence the alternative name ‘bent-wing bats’. The second phalanx is more than three times the length of the first phalanx; in vespertilionid bats, by contrast, it is usually not more than twice the length of the preceding phalanx. The lengthening of this digit (Figure 246a) gives the wing its long and narrow shape, allowing it to fly swiftly and efficiently in open areas. This family typically also has a raised braincase (Figure 246b). Molecular studies have revealed that this group does indeed deserve familial status, having diverged from their closest relatives, the Vespertilionidae, about 38–49 million years ago (Miller-Butterworth et al. 2007). The Molossidae had diverged from these two lineages about 10 million years earlier.
This family is represented by the single genus Miniopterus. Of the six currently recognised African species, four have been recorded in southern Africa. Although members of this genus are readily identifiable in the field owing to the greatly elongated third finger, distinguishing between the species is often difficult. Recent molecular evidence reveals that Miniopterus is comprised of several overlooked cryptic species in Africa.
The five southern African species differ in body size, but this is only absolute for skull length, although there is overlap even in this measurement between any two taxa. They emit low duty-cycle, frequency-modulated (LD-FM) echolocation calls. Analysis of cytochrome-b sequences indicate that the largest and smallest of the three South African species, M. inflatus and M. fraterculus, are each other's closest relatives, while M. natalensis (which overlaps in size with both these species) is more distantly related (Miller-Butterworth et al. 2005). A more comprehensive phylogeny that includes all five southern African species shows that M. fraterculus and M. minor are sister taxa; these two in turn are sister to M. inflatus, with the newly described M. mossambicus being basal to all three of these species. The fifth species, M. natalensis, is more distantly related to the other four miniopterids (Monadjem et al. 2013a, 2019).
Suborder Vespertilioniformes
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Bat Biology
- Ara Monadjem, Peter John Taylor, Fenton (Woody) Cotterill, M. Corrie Schoeman, University of KwaZulu-Natal, South Africa
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Summary
This section gives an introduction to the biology of bats. The behaviour of individual species is discussed in the species accounts section. More detailed information, beyond this overview, can be obtained in the specialised and comprehensive literature, including Adams and Pedersen (2000, 2013), Kunz (1982, 1988), Kunz and Racey (1998) and Kunz and Fenton (2003). The illustrated syntheses of Hill and Smith (1984), Richarz and Limbrunner (1993), Altringham (1996), Neuweiler (2000) and Fenton and Simmons (2015) are readable, and they compile valuable information about the natural history of the Chiroptera.
Since 2010, when the first edition of our book appeared, the comprehensive six-volume Mammals of Africa has been published. The entire fourth volume is dedicated to bats and shrews (Happold and Happold 2013). Compiled by 25 authors, this is the most comprehensive synthesis of information on African bat ecology and general biology. The even more sumptuous Handbook of the Mammals of the World (Wilson and Reeder 2009–2019) comes in nine lavishly illustrated volumes that began with the carnivores in 2009. The ninth volume is entirely devoted to bats, and was published in late 2019.
OVERVIEW
As the only mammals capable of sustained flight, bats have evolved to exploit the nocturnal aerosphere. Their adaptive radiation into a diversity of ecological and behavioural niches includes adaptations to shelter in an impressive variety of daytime roosts. The unique and rich diversity of bats is also constrained tightly by the physiological limits placed on body size by flight. These factors are represented in the behaviour, reproduction and ecology of over 1,400 living species of the order Chiroptera. Dependence on sonar for nocturnal navigation and locating prey is an equally important determinant of the natural history of these fascinating mammals (Hill and Smith 1984, Fenton and Simmons 2015). The emerging synthesis in the concepts and methods of aeroecology (Kunz et al. 2008) recognises the keystone roles of bats as the apex frugivores, pollinators and predators in the aerosphere (Chilson et al. 2017, Frick et al. 2013).
MIGRATION
In temperate (and some subtropical) regions, cold winters force bats to migrate or hibernate. A few species migrate over longer distances (> 500 km), while many make short, local (< 50 km) or medium distance (< 500 km) movements between winter and summer roosts (Fleming and Eby 2003).
Family Nycteridae Slit-faced Bats
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Summary
In Africa, this family is represented by the single genus Nycteris, comprising at least 15 species (Monadjem 2019), of which nine have been recorded in the region. Members of this genus are immediately recognisable by the arrangement of their noseleafs, which are covered by flaps of skin, giving the appearance of a slit running down the length of the muzzle (Figure 170b). This genus has another unique feature: the last vertebra of the tail is bifurcated at its tip, creating a ‘T’ effect, which assists with supporting the tail membrane (Figure 170a) (Hill and Smith 1984, Neuweiler 1990). The wings are characteristically broad with rounded tips, adaptations for slow, manoeuvrable flight (Norberg and Rayner 1987).
African species were originally classified in the genus Petalia. Andersen (1912b) grouped the then known species into four ‘species groups’ based on tragus shape (Figure 172), size of the posterior lower premolar, and whether the upper incisors are bifid or trifid (Figure 171) (Rosevear 1965). This arrangement was amended by Aellen (1959) into five groups, and recent molecular evidence confirms this higher-level classification of Nycteris (Demos et al. 2019a).
Convergent evolution appears conspicuous in Nycteris, because species that appear very similar in external characters are in fact divergent, cryptic species. For example, the distinctive bacular morphology of the very similar N. woodi and N. parisii suggests that the latter is more closely related to N. macrotis, whilst bacular characters in N. woodi point to its affinities with N. thebaica (Thomas et al. 1994). Previously, multivariate analyses of external measurements led van Cakenberghe and de Vree (1985) to conclude that N. woodi and N. parisii were conspecific. A new phylogeny for Nycteris was published after the revisions to this book were completed, and shows deep divisions within the genus. This suggests that several species comprise complexes, including N. thebaica, N. hispida and N. macrotis (Demos et al. 2019a). Integrative taxonomy is required to resolve the relationships of these populations at a Pan-African scale.
Nycterids are aptly called ‘whispering’ bats, as they emit soft, low-intensity multi-harmonic low dutycycle frequency-modulated (LD-FM) echolocation calls (Neuweiler 1990). The distinctly elongated ears presumably assist with the detection of prey, which often consists of invertebrates that are taken off the ground (Bowie et al. 1999).
Index
- Ara Monadjem, Peter John Taylor, Fenton (Woody) Cotterill, M. Corrie Schoeman, University of KwaZulu-Natal, South Africa
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Acknowledgements
- Ara Monadjem, Peter John Taylor, Fenton (Woody) Cotterill, M. Corrie Schoeman, University of KwaZulu-Natal, South Africa
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Family Molossidae Free-Tailed Bats
- Ara Monadjem, Peter John Taylor, Fenton (Woody) Cotterill, M. Corrie Schoeman, University of KwaZulu-Natal, South Africa
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Summary
This family is represented globally by about 126 species in 22 genera, of which 20 species in seven genera have been recorded from southern Africa (Taylor et al. 2019c). Members of the Molossidae are immediately recognisable by the unique arrangement of the tail, which extends beyond the end of the tail membrane, giving rise to the common name of this group – free-tailed bats (Figure 196a). They have plain faces without noseleafs, but the upper lip is typically wrinkled in most molossids (except in the larger Tadarida species), giving them a bulldog appearance (Figure 196b). Pheromones are important in their social behaviour. Each species has a distinct, strong, often aromatic scent (Kingdon 1974). The family includes some of the commonest (Chaerephon pumilus), as well as rarest (Chaerephon gallagheri and Tadarida lobata) African bat species. Much of what we know about the more poorly known species reflects the devoted research of the late Randolph Peterson of the Royal Ontario Museum and the late David L. Harrison.
Some African molossids, notably Mops condylurus, are locally abundant, but a surprising number of species are among the rarest and most poorly known of mammals, a situation exemplified by Tadarida lobata and T. ventralis (Figures 195 and 241). Their inconspicuous roosts and high-flying habits make them challenging to locate and study. By their very nature, many locality records of molossids reflect serendipitous collecting events, which are so sparse that our knowledge of even the basic biology of most species has not improved since their original discoveries – even though several of these species were made known to science over 150 years ago. This is epitomised by the enigmatic circumstances in which two new species of forest-dwelling African molossids were discovered over a century ago:
On 8 September 1910, a violent storm blew down a large hollow tree in the Ituri forest near the American Museum Congo expedition camp of Herbert Lang and James Chapin. Living inside the tree were two species of molossid bats, both later described by J. A. Allen as new to science, one as Mops congicus and the other as Chaerephon russatus (Allen et al. 1917). Both species have continued to be among the rarest in collections (Peterson 1971: 297).
Our knowledge of the biology of both M. congicus and C. russatus is still poor; it resides primarily in the museum records from Africa's equatorial forest belt.
Museum Collections And Pioneering Researchers
- Ara Monadjem, Peter John Taylor, Fenton (Woody) Cotterill, M. Corrie Schoeman, University of KwaZulu-Natal, South Africa
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Summary
During the eighteenth and nineteenth centuries, early explorers and hunters such as Temminck and Wahlberg collected biological specimens in southern Africa, which were then sent back to the major museums of Europe, such as the Natural History Museum in London (previously known as the British Museum of Natural History). Within the past few decades, mammalogists including Knud Andersen, Wim Bergmans, J. L. Eger, D. L. Harrison, R. W. Hayman, J. Eric Hill, J. Edwards Hill, J. Kingdon, D. Kock, K. F. Koopman, R. L. Peterson, D. R. Rosevear and O. Thomas, have done an excellent job of making much sense out of these invaluable historical collections, and have published taxonomic and biogeographical treatises in books, checklists and journals.
In the early twentieth century, local mammalogists, notably G. C. Shortridge, W. L. Sclater, and A. Roberts, worked prodigiously to build up collections of southern African bats and other mammals (Smithers 1984). Their researches were part of broader faunal surveys and research, especially of the mammals of the region. Many benefited directly and in kind from the devotion of Sir John Ellerman, a most studious researcher of the world's rodents. A significant milestone was the publication of Southern African Mammals 1758–1951: A Reclassification (Ellerman et al. 1953), which remains a valuable reference for the mammalian taxonomist. This tradition was continued in the latter part of the twentieth century by several biologists – notably Reay H. N. Smithers, J. A. J. ‘Waldo’ Meester, R. C. Wood and W. Frank H. Ansell made invaluable collections of bats and many other mammals. In the 1960s and 1970s, the Smithsonian Institution initiated groundbreaking surveys of African mammals in at least 13 African countries; the vast collections obtained are a testimony to many dedicated field collectors, including Tim N. Liversedge and John Herbert in southern Africa (Schmidt et al. 2008). This led to the compilation of the authoritative The Mammals of Africa: An Identification Manual, under the editorship of Waldo Meester and H. W. Setzer between 1971 and 1977 – the chapter on Chiroptera by Robert W. Hayman and J. Edwards Hill still includes the most up-to-date identification keys available for much of Africa.
Suborder Pteropodiformes
- Ara Monadjem, Peter John Taylor, Fenton (Woody) Cotterill, M. Corrie Schoeman, University of KwaZulu-Natal, South Africa
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Echolocation
- Ara Monadjem, Peter John Taylor, Fenton (Woody) Cotterill, M. Corrie Schoeman, University of KwaZulu-Natal, South Africa
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Summary
The majority of vertebrates rely on vision to perceive their environment. Even nocturnal predators such as owls and lions principally use their eyes to navigate and hunt. In contrast, most bats find food and avoid obstacles at night with great ease using an alternative sensory mechanism, called echolocation. Although usually associated with bats, other animals such as toothed whales, porpoises, some species of shrews and tenrecs, oilbirds, and several species of swiftlets also use echolocation.
Echolocating bats emit sound pulses and analyse the returning echoes to detect, characterise, and localise objects that reflect the impinging pulse as an echo (Fenton 1990, Schnitzler and Kalko 2001, Fenton et al. 2016) (Figure 35). Sound pulses are generated in the larynx (except in Rousettus species, which produce echolocation pulses by repeatedly clicking their tongue against the palate), and emitted through the mouth (e.g. Vespertilionidae, Miniopteridae, Cistugidae, Emballonuridae, and Molossidae) or nose (e.g. Rhinolophidae, Hipposideridae, Rhinonycteridae and Nycteridae). The ears, or pinnae, receive returning echoes, which are then funnelled and processed into the rest of the bat's hearing system.
Not all vocalisations produced by bats are echolocation calls. For example, the audible squeaks bats make in their roosts, the calls that mother and young make to one another, or the calls flying bats make to defend their foraging territories, are not echolocation calls. Instead, these are usually referred to as social calls and are less well understood than echolocation calls.
The echolocation frequencies of most bat species are ultrasonic (i.e. above the range of human hearing), and peak echolocation frequencies (i.e. the frequencies with the highest intensity) usually fall within 20–60 kHz (Fenton 1990). This may be due to the frequency-dependent effects of atmospheric attenuation and target strength (Jones and Rydell 2004). In contrast, many social calls are audible to humans.
Echolocation coupled with flight enables bats to capture nocturnal flying insects in a variety of habitats. This ability probably explains how the radiation of bat species has manifested into the highest trophic diversity among mammals (Patterson et al. 2004, Roemer et al. 2019). Nevertheless, not all bats echolocate (e.g. fruit bats from the family Pteropodidae, except Rousettus species), nor do all echolocating bats use the same type of echolocation.
ECHOLOCATION SYSTEMS
Two different echolocation systems – high and low duty-cycle echolocation – evolved independently in the Chiroptera (Eick et al. 2005).
Family Vespertilionidae Plain-faced Bats (Vespers)
- Ara Monadjem, Peter John Taylor, Fenton (Woody) Cotterill, M. Corrie Schoeman, University of KwaZulu-Natal, South Africa
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Summary
This is the largest family of bats, with at least 17 genera recorded from Africa, including the newly described Parahypsugo (Hutterer et al. 2019), of which 34 species in 11 genera occur in southern Africa. Of the remaining African vesper genera not recorded in southern Africa, Barbastella and Nyctalus are restricted to Mediterranean North Africa; Plecotus occurs there, as well as in Ethiopia, while Otonycteris occurs widely in North Africa. The fourth genus is excluded from this book because it comprises a highly dubious record of Phoniscus from ‘South Africa, eastern coast’ – this Asian genus is known in Africa from only two specimens, but there is doubt whether these were collected from the continent (Happold 2013d). Since the records come from a well-collected region of the continent, we suspect that this represents an example of mislabelled geographic location and hence we do not include it on our list. Three of the four genera not found in southern Africa are European species that extend only marginally into Africa north of the Sahara (Simmons 2005).
Members of the Vespertilionidae, sometimes called ‘evening bats’ (owing to their time of activity), are characterised by the absence of noseleafs, a long tail fully enclosed within the tail membrane (Figure 263), and a conspicuous tragus of variable shape and size (Figure 265). With the Miniopteridae and Cistugidae elevated to full family status, there are now three subfamilies within the southern African vespertilionids: Vespertilioninae, Myotinae, and Kerivoulinae (Amador et al. 2018). Based on a molecular phylogeny of nine mitochondrial and nuclear genes, Amador et al. (2018) recognise nine tribes within Vespertilioninae (four of which occur in southern Africa).
The Myotinae includes one African genus, Myotis, which is identified by its soft orange fur and long, narrow tragus.
The Kerivoulinae comprises one genus, Kerivoula, which is identified by its woolly fur and funnel-shaped ears that have a long, narrow tragus (Figure 264). The two widespread southern African species possess a fringe of hair on the outer margin of the tail membrane, a feature that is unique among vespertilionids of this region; this fringe of hair is absent in the remaining African species, including K. cf. phalaena captured recently from Mount Mabu (Monadjem et al. 2010a). These are small, inconspicuous bats that have been poorly collected; hence, little is known about the details of their biology (Hill and Smith 1984).
Contents
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Glossary
- Ara Monadjem, Peter John Taylor, Fenton (Woody) Cotterill, M. Corrie Schoeman, University of KwaZulu-Natal, South Africa
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Introduction
- Ara Monadjem, Peter John Taylor, Fenton (Woody) Cotterill, M. Corrie Schoeman, University of KwaZulu-Natal, South Africa
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Summary
Bats show a remarkable variety of adaptations to their environment and consequently vary considerably in size, appearance, and morphology. For example, the world's smallest bat, the 2 g Craseonycteris thonglongyai (Figure 1) from Thailand, is 600 times smaller than Pteropus vampyrus, which weighs up to 1.2 kg and has a wingspan of 1.8 m (Jones 1996). Bats occur worldwide, except in extreme polar and desert habitats. The species diversity of bats is highest in equatorial regions, notably in tropical forests, with progressively fewer species encountered with increasing latitudes in temperate climates (Schoeman et al. 2013, Herkt et al. 2016).
The oldest known bat fossil, Onychonycteris finneyi, was discovered recently in fossil beds in Wyoming, USA, and dates to 52 million years ago. Its diagnostic characters are intermediate between bats and non-flying mammals (Simmons et al. 2008). The fossil bat species had claws on each finger and relatively long hind legs in relation to its forelimbs, similar in ratio to sloths and lemurs. Its short, broad wings indicate that it probably alternated between flapping flight and gliding, and that it was also capable of clambering in trees. The small cochlea (ear) bones show that it could not echolocate, suggesting that flight evolved before echolocation in bats. By 50 million years ago, during the Eocene, there were already at least three families and 13 species of bats. Eocene fossil bats are known from North America, Europe and Australia. All these ancestors were already fully developed as bats, although they did not closely resemble extant species. Recent discoveries in Egypt of extinct bats from the late Eocene and early Oligocene (37–27 million years ago), which appear more closely related to modern species, suggest that bats may have diversified in Africa following migration of primitive ancestors from Europe (Gunnell et al. 2008).
Taxonomic classifications are essential to the universal communication of verifiable scientific knowledge (Cotterill 1995a, Ghiselin 2005), in which descriptions of species, and any other taxon, should adhere to scientific conventions of nomenclature (Winston 1999, Gardner and Hayssen 2004). In this respect, the persisting confusion over the real species diversity of African house bats (genus Scotophilus) testifies to why precise and accurate taxonomy is so critical to all of biology and conservation. This example especially highlights the relevance of type material, preserved in museums, if we are to apply scientific evidence to clarify the distinctiveness of a population in a taxonomic revision.
Family Emballonuridae Sheath-tailed Bats
- Ara Monadjem, Peter John Taylor, Fenton (Woody) Cotterill, M. Corrie Schoeman, University of KwaZulu-Natal, South Africa
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Summary
Within Africa, this family comprises seven species in three genera; four species occur in southern Africa. Emballonurids are immediately recognisable by the particular configuration of the tail, which is only partly enclosed by the tail membrane proximally, but independent of the membrane distally (Figure 159a). Hence, the tail appears to protrude out from above the tail membrane. All members of this family also have large eyes and a plain face without noseleafs (Figure 159b).
Members of the genus Taphozous can be distinguished by the presence of a radio-metacarpal pouch (Figure 160), absent in Coleura and Saccolaimus. Coleura species are significantly smaller than Taphozous, while Saccolaimus species are much larger. Gular sacs are present in some species, but not others. In some species, such gular sacs are well developed in both sexes, while in other species they are only present in males. The development of gular sacs does not appear to be of taxonomical value. The wings of Emballonuridae are typically long and pointed – an adaptation for swift flight (Norberg and Rayner 1987). Echolocation calls emitted by this family are low duty-cycle constant frequency (LD-CF) and quasi-constant frequency (LD-QCF) with multi-harmonics (Hill and Smith 1984, Neuweiler 1990).
Description: Coleura afra is a small bat with a mass of around 11 g. The pelage is deep brown on the upper parts and paler on the underparts. The fur is bicoloured, with hairs paler at the base than at the tip (Dunlop 1997). The wings are translucent and light brown. The ears are long and narrow with a characteristically shaped tragus. The muzzle is narrow and naked with nostrils projecting beyond the lower jaw. The eyes are strikingly large for a microbat. A gular sac and radio-metacarpal pouches are absent. Females are slightly larger than males (McWilliam 1987a).
The skull is delicate with relatively weak zygomatic arches. The braincase is rounded and elevated above the level of the rostrum. A shallow frontal depression is present, flanked by inflations of bones on each side of the rostrum. The sagittal crest is well developed and the lambdoid crest is weak or absent. The dental formula is 1123/3123 = 32 (Rosevear 1965).
Species Accounts
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Summary
This section contains species accounts for 125 species of bats known to occur in the southern African region, accompanied by identification matrices, distribution maps, spectrograms (also called sonograms), photographs of bats and their skulls, and tables of measurements.
Rather than using dichotomous keys, which can be misleading, especially when characters are missing, we provide identification matrices throughout, providing important characters for all species. Table 4 presents an identification matrix for distinguishing between the 11 families of bats found in southern Africa.
The families are arranged phylogenetically within the two orders Pteropodiformes and Vespertilioniformes. Each family account starts with a general description of the family and genera, followed by one or more identification matrices to the genera and species. Within each identification matrix, taxa are arranged according to ascending size (FA length). Species accounts are arranged alphabetically within each family. Each species account covers the following aspects.
Name: Scientific and common names are given (following Simmons 2005, except as stated), together with the author and date of description of each genus and species.
Conservation status: Global Red List categories appear after the name in each account. These are sourced from the 2017 IUCN Red List of Threatened Species (www.iucnredlist.org); the original ratings were based on mutual consensus among bat biologists at the January 2004 Global Mammal Assessment meeting in London.
Description: External, cranial and dental characters are described. Where known, the dental formula of each species is presented as follows: 2132/2133 = 34, where the numbers before the slash refer to the teeth in half of the upper jaw and the ones after it to half the teeth in the lower jaw. In this example, the species has four upper incisors, two canines, six premolars and four molars in the upper jaw – a total of 34 teeth. Figure 40 illustrates the dental and osteological features referred to in this book.
Key identification features: Diagnostic species characteristics useful for identification are provided, as well as comparisons between closely related species that may be easily confused. These features are also summarised in the relevant identification matrices provided for each family and genus. In some cases, diagnostic traits are visible from the species photographs and skull photographs (dorsal, ventral and lateral views of the cranium and lateral view of the mandible).
Biogeography
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Summary
Biogeography is the study of the spatial patterns in the distribution of biodiversity and the causes of those patterns, especially differences in species’ distributions. At this level, species distribution patterns are explained through a combination of historical factors that have governed events of dispersal, speciation and extinction. Such factors include the influence of geological and climatic changes, notably continental drift, glaciation, variations in sea level, and drainage evolution, in combination with macro-ecological factors that determine the availability of ecological resources to species in an assemblage (Morrone 2009).
GEOLOGY
The geology of a continent has a profound influence on the living organisms that have evolved across its landscapes. Local and regional geological controls on the composition of Africa's bat faunas provide many textbook examples of this relationship. For example, the distributions of rupicolous and cavernicolous species are associated closely with particular formations of granites, sandstones and limestones that have crevices and caves eroded and weathered into these rocks (Figure 19). The geomorphology of the landscape plays an equally significant role, as such caves and crevices tend to be clustered along cliffs and scarps. In recent times, mining activity has influenced bat roost availability, especially in the gold-rich contact zones of the granite-greenstone belts. These artificial cavities are of great significance to many cavernicolous bat species. Numerous mining shafts and adits have been sunk over the past few centuries at many sites in Namibia, central Zimbabwe, South Africa, Zambia and the southern Democratic Republic of the Congo (DRC).
This overview of the formative events that forged the southern portion of the African continent singles out formations and events that bear directly on the composition of the regional bat fauna, especially rock formations that provide daylight roosts for cavernicolous and rupicolous species. Unless specifically referenced, the remainder of this section draws on syntheses of knowledge of the geology of South Africa, Lesotho and Eswatini (McCarthy and Rubidge 2005, Johnson et al. 2006), Namibia (Miller 2008) and Zimbabwe (Stagman 1978). The geology of Angola, the Congo basin, Botswana, Malawi, Mozambique and Zambia awaits deserving synthesis, with relevant knowledge scattered widely through the primary literature and in the less accessible in-house reports of mining companies.
Family Hipposideridae Leaf-Nosed Bats
- Ara Monadjem, Peter John Taylor, Fenton (Woody) Cotterill, M. Corrie Schoeman, University of KwaZulu-Natal, South Africa
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- Book:
- Bats of Southern and Central Africa
- Published by:
- Wits University Press
- Published online:
- 16 June 2021
- Print publication:
- 01 October 2020, pp 161-188
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Summary
The Hipposideridae previously included the genera Cloeotis and Triaenops, which are now placed in the separate family Rhinonycteridae (Foley et al. 2015). Furthermore, the speciose genus Hipposideros has recently been split up, adding the genera Doryrhina and Macronycteris to the family; all three genera occur in Africa (Foley et al. 2017). A fourth genus, Asellia, is widespread in the northern parts of the continent, but does not extend into southern Africa. Members of this family occurring in our region typically have a prominent noseleaf that is simple and elliptical (Figure 80a), in contrast to the threepronged noseleaf of the rhinonycterids (Figure 80b), and they lack the single, triangular erect process of the Rhinolophidae (Figure 97). Note that Asellia also has a three-pronged noseleaf, but does not occur in southern Africa.
Hipposideros is a large genus found throughout the Old World. It is well represented in tropical Africa with species richness declining away from the equator. In our region, no more than two species co-occur at any one site (and usually only one species). The noseleaf (half-moon-shaped; Figure 80a) is widely used to identify species within the genus.
Doryrhina cyclops (Temminck 1853), a forest species, occurs on the extreme northwest border of the region. This is based on two records: one from Lovanium, Republic of Congo (Congo–Brazzaville) (04.25S 15.03E, RMCA 31201), and a second mapped south of the Bas Congo in the DRC (which would fall within our region), but without reference to a museum specimen (Decher and Fahr 2005); however, this latter record is not mentioned by van Cakenberghe et al. (2017). For this reason, we do not provide an account for this species until a verified record is confirmed within our region. Doryrhina cyclops is a large bat (FA typically ~65–70 mm) with distinctive long woolly pelage and a noseleaf with two median club-shaped processes (Figure 81), making it unlikely to be confused with any other southern African hipposiderid (Decher and Fahr 2005).
The genus Macronycteris includes the largest bats in the family and is represented in the region by two species: M. vittatus is relatively widespread in the northern parts of southern Africa, whereas M. gigas is poorly known.
Echolocation calls in this family typically show a high duty-cycle, constant frequency (HD-CF) component. The CF component differs between species and may be used as a guide to their identification (Monadjem et al. 2007, 2013c, Webala et al. 2019).
Family Cistugidae Cistugo Bats
- Ara Monadjem, Peter John Taylor, Fenton (Woody) Cotterill, M. Corrie Schoeman, University of KwaZulu-Natal, South Africa
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- Book:
- Bats of Southern and Central Africa
- Published by:
- Wits University Press
- Published online:
- 16 June 2021
- Print publication:
- 01 October 2020, pp 467-476
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Summary
This is the smallest family of bats in Africa, with only two species in the genus Cistugo. This genus was previously listed in the family Vespertilionidae (Simmons 2005), but recent molecular analyses have demonstrated its ancient origins, having separated from the Vespertilionidae 34 million years ago (Lack et al. 2010). The family is endemic to southern Africa.
Members of the Cistugidae are small bats (mass typically 4–7 g) that share the following characteristics with the Vespertilionidae: the absence of noseleafs, a long tail fully enclosed within the tail membrane (Figure 263), and a conspicuous tragus of variable shape and size (Figure 265 illustrates different tragus designs in the Vespertilionidae). However, members of the Cistugidae can be distinguished from all African vespers on the presence of 2–4 glands in the plagiopatagium (wing membrane) situated posterior to the humerus (Seamark and Kearney 2006; Figure 259b). Cistugo has a similar dental formula to Myotis, both possessing a pair of tiny anterior premolars on each side of the upper jaw, and Cistugo was in fact considered a subgenus of Myotis until relatively recently (Bronner et al. 2003). However, these two genera are cytogenetically distinct, with Cistugo having the karyotype 2n = 50, whereas all Myotis have 2n = 44 (Lack et al. 2010). Practically nothing is known about the general biology of this genus.
Cistugids emit low duty-cycle, frequencymodulated (LD-FM) echolocation calls, at intermediate peak frequencies.
Description: Cistugo lesueuri is a small bat with a mass of around 6 g and very similar in appearance to the smaller C. seabrae. The individual hairs are long and stand away from the body, giving the fur a soft feel. It is dull yellow to yellow-beige above and paler yellow-cream below. The individual hairs are dark at their base and yellowish at their tips. The wings are dark brown with a conspicuous gland present in the membrane on each side. The face is plain, without any noseleafs. The ears are brown and moderately sized, with a long, narrow tragus. The sexes are alike.
The skull is delicate with weak zygomatic arches. In lateral view, the braincase rises slightly above the rostrum, with a smoothly concave forehead. There is a further concavity in the parietal region, owing to an occipital bulge. The sagittal and lambdoid crests are weak or absent. The mastoid processes are not evident in dorsal view.