Microbats | |
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Townsend's big-eared bats in a cave. | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Chiroptera |
Suborder: | Microchiroptera Dobson, 1875 |
Superfamilies | |
Microbats constitute the suborder Microchiroptera within the order Chiroptera (bats). Bats have long been differentiated into Megachiroptera (megabats) and Microchiroptera, based on their size, the use of echolocation by the Microchiroptera and other features; molecular evidence suggests a somewhat different subdivision, as the microbats have been shown to be a paraphyletic group. [1]
Microbats are 4 to 16 cm (1.6–6.3 in) long. [2] Most microbats feed on insects, but some of the larger species hunt birds, lizards, frogs, smaller bats or even fish. Only three species of microbat feed on the blood of large mammals or birds ("vampire bats"); these bats live in South and Central America.
Although most "Leaf-nose" microbats are fruit and nectar-eating, the name “leaf-nosed” isn't a designation meant to indicate the preferred diet among said variety. [3] Three species follow the bloom of columnar cacti in northwest Mexico and the Southwest United States northward in the northern spring and then the blooming agaves southward in the northern fall (autumn). [4] Other leaf-nosed bats, such as Vampyrum spectrum of South America, hunt a variety of prey such as lizards and birds. The horseshoe bats of Europe, as well as California leaf-nosed bats, have a very intricate leaf-nose for echolocation, and feed primarily on insects.
The form and function of microbat teeth differ as a result of the various diets these bats can have. Teeth are primarily designed to break down food; therefore, the shape of the teeth correlate to specific feeding behaviors. [5] In comparison to megabats which feed only on fruit and nectar, microbats illustrate a range of diets and have been classified as insectivores, carnivores, sanguinivores, frugivores, and nectarivores. [6] Differences seen between the size and function of the canines and molars among microbats in these groups vary as a result of this.
The diverse diets of microbats reflect having dentition, or cheek teeth, that display a morphology derived from dilambdodont teeth, which are characterized by a W-shaped ectoloph, or stylar shelf. [7] [ not specific enough to verify ] A W-shaped dilambdodont upper molar includes a metacone and paracone, which are located at the bottom of the “W”; while the rest of the “W” is formed by crests that run from the metacone and paracone to the cusps of the stylar self.
Microbats display differences between the size and shape of their canines and molars, in addition to having distinctive variations among their skull features that contribute to their ability to feed effectively. Frugivorous microbats have small stylar shelf areas, short molariform rows, and wide palates and faces. In addition to having wide faces, frugivorous microbats have short skulls, which place the teeth closer to the fulcrum of the jaw lever, allowing an increase in jaw strength. [8] Frugivorous microbats also possess a different pattern on their molars compared to carnivorous, insectivorous, nectarivorous, and sanguinivorous microbats. [6] In contrast, insectivorous microbats are characterized by having larger, but fewer teeth, long canines, and shortened third upper molars; while carnivorous microbats have large upper molars. Generally, microbats that are insectivores, carnivores, and frugivores have large teeth and small palates; however, the opposite is true for microbats that are nectarivores. Though differences exist between the palate and teeth sizes of microbats, the proportion of the sizes of these two structures are maintained among microbats of various sizes. [6]
Echolocation is the process where an animal produces a sound of certain wavelength, and then listens to and compares the reflected echoes to the original sound emitted. Bats use echolocation to form images of their surrounding environment and the organisms that inhabit it by eliciting ultrasonic waves via their larynx. [9] [10] The difference between the ultrasonic waves produced by the bat and what the bat hears provides the bat with information about its environment. Echolocation aids the bat in not only detecting prey, but also in orientation during flight. [11]
Most microbats generate ultrasound with their larynx and emit the sound through their nose or mouth. [12] Sound productions are generated from the vocal folds in mammals due to the elastic membranes that compose these folds. Vocalization requires these elastic membranes because they act as a source to transform airflow into acoustic pressure waves. Energy is supplied to the elastic membranes from the lungs, and results in the production of sound. The larynx houses the vocal cords and forms the passageway for the expiratory air that will produce sound. [13] Microbat range in frequency from 14,000 to over 100,000 hertz, well beyond the range of the human ear (typical human hearing range is considered to be from 20 to 20,000 Hz). The emitted vocalizations form a broad beam of sound used to probe the environment, as well as communicate with other bats. At the molecular level, it has been found that CPLX1 is involved in this ultrasonic wave production. [14]
Laryngeal echolocation is the dominant form of echolocation in microbats, however, it is not the only way in which microbats can produce ultrasonic waves. Excluding non-echolocating and laryngeally echolocating microbats, other species of microbats and megabats have been shown to produce ultrasonic waves by clapping their wings, clicking their tongues, or using their nose. [9] Laryngeally echolocating bats, in general, produce ultrasonic waves with their larynx that is specialized to produce sounds of short wavelength. The larynx is located at the cranial end of the trachea and is surrounded by cricothyroid muscles and thyroid cartilage. For reference, in humans, this is the area where the Adam's apple is located. Phonation of ultrasonic waves is produced through the vibrations of the vocal membranes in the expiratory air. The intensity that these vocal folds vibrate at varies with activity and between bat species. [15] A characteristic of laryngeally echolocating microbats that distinguishes them from other echolocating microbats is the articulation of their stylohyal bone with their tympanic bone. The stylohyal bones are part of the hyoid apparatus that help support the throat and larynx. The tympanic bone forms the floor of the middle ear. In addition to the connection between the stylohyal bone and the tympanic bone as being an indicator of laryngeally echolocating microbats, another definitive marker is the presence of a flattened and expanded stylohyal bone at the cranial end. [10]
Microbats that laryngeally echolocate must be able to distinguish between the differences of the pulse that they produce and the returning echo that follows by being able to process and understand the ultrasonic waves at a neuronal level, in order to accurately obtain information about their surrounding environment and orientation in it. [9] The connection between the stylohyal bone and the tympanic bone enables the bat to neurally register the outgoing and incoming ultrasonic waves produced by the larynx. [11] Furthermore, the stylohyal bones connect the larynx to the tympanic bones via a cartilaginous or fibrous connection (depending on the species of bat). Mechanically the importance of this connection is that it supports the larynx by anchoring it to the surrounding cricothyroid muscles, as well as draws it closer to the nasal cavity during phonation. The stylohyal bones are often reduced in many other mammals, however, they are more prominent in laryngeally echolocating bats and are part of the mammalian hyoid apparatus. The hyoid apparatus functions in breathing, swallowing, and phonation in microbats as well as other mammals. An important feature of the bony connection in laryngeally echolocating microbats is the extended articulation of the ventral portion of the tympanic bones and the proximal end of the stylohyal bone that bends around it to make this connection. [9]
While bats have been traditionally divided into megabats and microbats, recent molecular evidence has shown the superfamily Rhinolophoidea to be more genetically related to megabats than to microbats, indicating the microbats are paraphyletic. To resolve the paraphyly of microbats, the Chiroptera were redivided into suborders Yangochiroptera (which includes Nycteridae, vespertilionoids, noctilionoids, and emballonuroids) and Yinpterochiroptera, which includes megabats, rhinopomatids, Rhinolophidae, and Megadermatidae. [1]
This is the classification according to Simmons and Geisler (1998):
Superfamily Emballonuroidea
Superfamily Rhinopomatoidea
Superfamily Rhinolophoidea
Superfamily Vespertilionoidea
Superfamily Molossoidea
Superfamily Nataloidea
Superfamily Noctilionoidea
Echolocation, also called bio sonar, is a biological active sonar used by several animal groups, both in the air and underwater. Echolocating animals emit calls and listen to the echoes of those calls that return from various objects near them. They use these echoes to locate and identify the objects. Echolocation is used for navigation, foraging, and hunting prey.
Megabats constitute the family Pteropodidae of the order Chiroptera (bats). They are also called fruit bats, Old World fruit bats, or—especially the genera Acerodon and Pteropus—flying foxes. They are the only member of the superfamily Pteropodoidea, which is one of two superfamilies in the suborder Yinpterochiroptera. Internal divisions of Pteropodidae have varied since subfamilies were first proposed in 1917. From three subfamilies in the 1917 classification, six are now recognized, along with various tribes. As of 2018, 197 species of megabat had been described.
Vespertilionidae is a family of microbats, of the order Chiroptera, flying, insect-eating mammals variously described as the common, vesper, or simple nosed bats. The vespertilionid family is the most diverse and widely distributed of bat families, specialised in many forms to occupy a range of habitats and ecological circumstances, and it is frequently observed or the subject of research. The facial features of the species are often simple, as they mainly rely on vocally emitted echolocation. The tails of the species are enclosed by the lower flight membranes between the legs. Over 300 species are distributed all over the world, on every continent except Antarctica. It owes its name to the genus Vespertilio, which takes its name from a word for bat, vespertilio, derived from the Latin term vesper meaning 'evening'; they are termed "evening bats" and were once referred to as "evening birds".
Shrews are small mole-like mammals classified in the order Eulipotyphla. True shrews are not to be confused with treeshrews, otter shrews, elephant shrews, West Indies shrews, or marsupial shrews, which belong to different families or orders.
Horseshoe bats are bats in the family Rhinolophidae. In addition to the single living genus, Rhinolophus, which has about 106 species, the extinct genus Palaeonycteris has been recognized. Horseshoe bats are closely related to the Old World leaf-nosed bats, family Hipposideridae, which have sometimes been included in Rhinolophidae. The horseshoe bats are divided into six subgenera and many species groups. The most recent common ancestor of all horseshoe bats lived 34–40 million years ago, though it is unclear where the geographic roots of the family are, and attempts to determine its biogeography have been indecisive. Their taxonomy is complex, as genetic evidence shows the likely existence of many cryptic species, as well as species recognized as distinct that may have little genetic divergence from previously recognized taxa. They are found in the Old World, mostly in tropical or subtropical areas, including Africa, Asia, Europe, and Oceania.
The ghost bat is a species of bat found in northern Australia. The species is the only Australian bat that preys on large vertebrates – birds, reptiles and other mammals – which they detect using acute sight and hearing, combined with echolocation, while waiting in ambush at a perch. The wing membrane and bare skin is pale in colour, their fur is light or dark grey over the back and paler at the front. The species has a prominent and simple nose-leaf, their large ears are elongated and joined at lower half, and the eyes are also large and dark in colour. The first description of the species was published in 1880, its recorded range has significantly contracted since that time.
The short-palated fruit bat is a species of frugivorous megabat in the family Pteropodidae. It is found in Cameroon, Central African Republic, and Democratic Republic of the Congo. Its natural habitat is subtropical or tropical moist lowland forests. Births occur in May.
The diadem leaf-nosed bat or diadem roundleaf bat is one of the most widespread species of bat in the family Hipposideridae. It is probably most closely related to Hipposideros demissus from Makira and to Hipposideros inornatus from the Northern Territory in Australia. Hipposideros diadema is found in Australia, Indonesia, Malaysia, Myanmar, the Philippines, Thailand, and Vietnam.
The lesser mouse-tailed bat is a species of microbat in the family Rhinopomatidae. Also referred to as Hardwicke's lesser mouse-tailed bat and long-tailed bat, it is named after Major General Thomas Hardwicke (1755–1835), an English soldier and naturalist who served many years in India. It is found in North Africa, some parts of central and eastern Africa, West Asia and east to the Indian subcontinent.
In evolutionary biology, the flying primate hypothesis is that megabats, a subgroup of Chiroptera, form an evolutionary sister group of primates. The hypothesis began with Carl Linnaeus in 1758, and was again advanced by J.D. Smith in 1980. It was proposed in its modern form by Australian neuroscientist Jack Pettigrew in 1986 after he discovered that the connections between the retina and the superior colliculus in the megabat Pteropus were organized in the same way found in primates, and purportedly different from all other mammals. This was followed up by a longer study published in 1989, in which this was supported by the analysis of many other brain and body characteristics. Pettigrew suggested that flying foxes, colugos, and primates were all descendants of the same group of early arboreal mammals. The megabat flight and the colugo gliding could be both seen as locomotory adaptations to a life high above the ground.
The calcar, also known as the calcaneum, is the name given to a spur of cartilage arising from inner side of ankle and running along part of outer interfemoral membrane in bats, as well as to a similar spur on the legs of some arthropods.
Onychonycteris is the more primitive of the three oldest bats known from complete skeletons, having lived in the area that is current day Wyoming during the Eocene period, 52.5 million years ago.
The Yinpterochiroptera is a suborder of the Chiroptera, which includes taxa formerly known as megabats and five of the microbat families: Rhinopomatidae, Rhinolophidae, Hipposideridae, Craseonycteridae, and Megadermatidae. This suborder is primarily based on molecular genetics data. This proposal challenged the traditional view that megabats and microbats form monophyletic groups of bats. Further studies are being conducted, using both molecular and morphological cladistic methodology, to assess its merit.
Yangochiroptera, or Vespertilioniformes, is a suborder of Chiroptera that includes most of the microbat families, except the Rhinopomatidae, Rhinolophidae, Hipposideridae, and Megadermatidae. These other families, plus the megabats, are seen as part of another suborder, the Yinpterochiroptera. All bats in Yangochiroptera use laryngeal echolocation(LE), which involves the use of high-frequency sounds to detect prey and avoid obstacles.
Bats are flying mammals of the order Chiroptera. With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most birds, flying with their very long spread-out digits covered with a thin membrane or patagium. The smallest bat, and arguably the smallest extant mammal, is Kitti's hog-nosed bat, which is 29–34 mm (1.1–1.3 in) in length, 150 mm (5.9 in) across the wings and 2–2.6 g (0.071–0.092 oz) in mass. The largest bats are the flying foxes, with the giant golden-crowned flying fox reaching a weight of 1.6 kg (3.5 lb) and having a wingspan of 1.7 m.
Ultrasound avoidance is an escape or avoidance reflex displayed by certain animal species that are preyed upon by echolocating predators. Ultrasound avoidance is known for several groups of insects that have independently evolved mechanisms for ultrasonic hearing. Insects have evolved a variety of ultrasound-sensitive ears based upon a vibrating tympanic membrane tuned to sense the bat's echolocating calls. The ultrasonic hearing is coupled to a motor response that causes evasion of the bat during flight.
Echolocation systems of animals, like human radar systems, are susceptible to interference known as echolocation jamming or sonar jamming. Jamming occurs when non-target sounds interfere with target echoes. Jamming can be purposeful or inadvertent, and can be caused by the echolocation system itself, other echolocating animals, prey, or humans. Echolocating animals have evolved to minimize jamming, however; echolocation avoidance behaviors are not always successful.
Rhinonicteris tedfordi is an extinct species of microbat, of the order Chiroptera, known from fossil material found in Australia.
Brevipalatus mcculloughi is a species of bat that existed in the early Miocene. It was discovered at a fossil deposit of the Riversleigh World Heritage Area.