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Microbats constitute the
suborder Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and ...
Microchiroptera within the
order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of ...
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 In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and most of its descendants, excluding a few monophyletic subgroups. The group is said to be paraphyletic ''with respect to'' the excluded subgroups. In ...
group.


Characteristics

Microbats are long. Most microbats feed on insects, but some of the larger species hunt birds, lizards, frogs, smaller bats or even
fish Fish are Aquatic animal, aquatic, craniate, gill-bearing animals that lack Limb (anatomy), limbs with Digit (anatomy), digits. Included in this definition are the living hagfish, lampreys, and Chondrichthyes, cartilaginous and bony fish as we ...
. Only three species of microbat feed on the blood of large mammals or birds ("
vampire bat Vampire bats, species of the subfamily Desmodontinae, are leaf-nosed bats found in Central and South America. Their food source is blood of other animals, a dietary trait called hematophagy. Three extant bat species feed solely on blood: the ...
s"); 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. 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). Other leaf-nosed bats, such as ''
Vampyrum spectrum The spectral bat (''Vampyrum spectrum''), also called the great false vampire bat or Linnaeus's false vampire bat, is a large, carnivorous leaf-nosed bat found in Mexico, Central America, and South America. It is the only member of the genus '' ...
'' 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.


Differences from megabats

* Microbats use echolocation, whereas megabats do not typically. (The Egyptian fruit bat ''
Rousettus egyptiacus The Egyptian fruit bat or Egyptian rousette (''Rousettus aegyptiacus'') is a species of megabat that is found in Africa, the Middle East, the Mediterranean, and the Indian subcontinent. It is one of three ''Rousettus'' species with an African-M ...
'' is an exception, but does not use the larynx echolocation method of microbats, instead giving scientists the theory that it clicks using its nasal passages and back of its tongue.) * Microbats lack the claw at the second finger of the forelimb. This finger appears thinner and almost bonded by tissue with the third finger for extra support during flight. * Megabats lack tails, with the exception of a few genera such as ''Nyctimene'', whereas this trait only occurs in certain species of microbats. * The ears of microbats possess a tragus (thought to be crucial in echolocation) and are relatively larger than megabat ears, whereas megabat ears are comparatively small and lack a tragus. * Megabat eyes are quite large, whereas microbat eyes are comparatively smaller.


Dentition

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. In comparison to megabats which feed only on fruit and nectar, microbats illustrate a range of diets and have been classified as insectivores,
carnivores A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other so ...
, sanguinivores, frugivores, and nectarivores.Freeman, Patricia W., "Form, Function, and Evolution in Skulls and Teeth of Bats" (1998). Papers in Natural Resources. 9. 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. 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. Frugivorous microbats also possess a different pattern on their molars compared to carnivorous, insectivorous, nectarivorous, and sanguinivorous microbats. 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.


Echolocation

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 The larynx (), commonly called the voice box, is an organ in the top of the neck involved in breathing, producing sound and protecting the trachea against food aspiration. The opening of larynx into pharynx known as the laryngeal inlet is about ...
. 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.


Production of ultrasonic waves

Most microbats generate ultrasound with their larynx and emit the sound through their nose or mouth. Sound productions are generated from the
vocal folds In humans, vocal cords, also known as vocal folds or voice reeds, are folds of throat tissues that are key in creating sounds through vocalization. The size of vocal cords affects the pitch of voice. Open when breathing and vibrating for speec ...
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 In humans, vocal cords, also known as vocal folds or voice reeds, are folds of throat tissues that are key in creating sounds through vocalization. The size of vocal cords affects the pitch of voice. Open when breathing and vibrating for speec ...
and forms the passageway for the expiratory air that will produce sound. Microbat range in frequency from 14,000 to over 100,000
hertz The hertz (symbol: Hz) is the unit of frequency in the International System of Units (SI), equivalent to one event (or cycle) per second. The hertz is an SI derived unit whose expression in terms of SI base units is s−1, meaning that o ...
, 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.


Laryngeally echolocating microbats

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. Laryngeally echolocating bats, in general, produce ultrasonic waves with their larynx that is specialized to produce sounds of short
wavelength In physics, the wavelength is the spatial period of a periodic wave—the distance over which the wave's shape repeats. It is the distance between consecutive corresponding points of the same phase on the wave, such as two adjacent crests, tr ...
. The
larynx The larynx (), commonly called the voice box, is an organ in the top of the neck involved in breathing, producing sound and protecting the trachea against food aspiration. The opening of larynx into pharynx known as the laryngeal inlet is about ...
is located at the
cranial Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
end of the
trachea The trachea, also known as the windpipe, is a cartilaginous tube that connects the larynx to the bronchi of the lungs, allowing the passage of air, and so is present in almost all air- breathing animals with lungs. The trachea extends from t ...
and is surrounded by
cricothyroid muscle The cricothyroid muscle is the only tensor muscle of the larynx aiding with phonation. It is innervated by the superior laryngeal nerve. Its action tilts the thyroid forward to help tense the vocal cords. Structure The cricothyroid muscle ...
s and
thyroid cartilage The thyroid cartilage is the largest of the nine cartilages that make up the ''laryngeal skeleton'', the cartilage structure in and around the trachea that contains the larynx. It does not completely encircle the larynx (only the cricoid cartilage ...
. For reference, in
human Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, cultu ...
s, this is the area where the
Adam's apple The Adam's apple or laryngeal prominence is the protrusion in the human neck formed by the angle of the thyroid cartilage surrounding the larynx, typically visible in men, less frequently in women. Structure The topographic structure which is e ...
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. 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 The hyoid apparatus is the collective term used in veterinary anatomy for the bones which suspend the tongue and larynx. It consists of pairs of stylohyoid, thyrohyoid, epihyoid and ceratohyoid bones, and a single basihyoid bone. The hyoid appara ...
that help support the throat and larynx. The tympanic bone forms the floor of the
middle ear The middle ear is the portion of the ear medial to the eardrum, and distal to the oval window of the cochlea (of the inner ear). The mammalian middle ear contains three ossicles, which transfer the vibrations of the eardrum into waves in the ...
. 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. 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
neuron A neuron, neurone, or nerve cell is an electrically excitable cell that communicates with other cells via specialized connections called synapses. The neuron is the main component of nervous tissue in all animals except sponges and placozoa ...
al level, in order to accurately obtain information about their surrounding environment and orientation in it. 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 The larynx (), commonly called the voice box, is an organ in the top of the neck involved in breathing, producing sound and protecting the trachea against food aspiration. The opening of larynx into pharynx known as the laryngeal inlet is about ...
. Furthermore, the stylohyal bones connect the larynx to the tympanic bones via a
cartilaginous Cartilage is a resilient and smooth type of connective tissue. In tetrapods, it covers and protects the ends of long bones at the joints as articular cartilage, and is a structural component of many body parts including the rib cage, the neck a ...
or
fibrous Fiber or fibre (from la, fibra, links=no) is a natural or artificial substance that is significantly longer than it is wide. Fibers are often used in the manufacture of other materials. The strongest engineering materials often incorporat ...
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 cricothryroid muscles, as well as draws it closer to the
nasal cavity The nasal cavity is a large, air-filled space above and behind the nose in the middle of the face. The nasal septum divides the cavity into two cavities, also known as fossae. Each cavity is the continuation of one of the two nostrils. The nasal ...
during
phonation The term phonation has slightly different meanings depending on the subfield of phonetics. Among some phoneticians, ''phonation'' is the process by which the vocal folds produce certain sounds through quasi-periodic vibration. This is the defin ...
. 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.


Classification

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 In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and most of its descendants, excluding a few monophyletic subgroups. The group is said to be paraphyletic ''with respect to'' the excluded subgroups. In ...
of microbats, the Chiroptera were redivided into suborders
Yangochiroptera 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 ...
(which includes Nycteridae, vespertilionoids, noctilionoids, and emballonuroids) and
Yinpterochiroptera The Yinpterochiroptera (or Pteropodiformes) is a suborder of the Chiroptera, which includes taxa formerly known as megabats and five of the microbat families: Rhinopomatidae, Rhinolophidae, Hipposideridae, Craseonycteridae, and Megadermatid ...
, which includes megabats, rhinopomatids, Rhinolophidae, and Megadermatidae. This is the classification according to Simmons and Geisler (1998): Superfamily
Emballonuroidea Emballonuroidea is a superfamily of bats containing two families: Emballonuridae and Nycteridae. It is one of three superfamilies in the suborder Yangochiroptera, the others being Noctilionoidea Noctilionoidea is a superfamily of bats contai ...
* Family
Emballonuridae Emballonuridae is a family of microbats, many of which are referred to as sac-winged or sheath-tailed bats. They are widely distributed in tropical and subtropical regions around the world. The earliest fossil records are from the Eocene. Descr ...
(
sac-winged bat Emballonuridae is a family of microbats, many of which are referred to as sac-winged or sheath-tailed bats. They are widely distributed in tropical and subtropical regions around the world. The earliest fossil records are from the Eocene. Descr ...
s or
sheath-tailed bat Emballonuridae is a family of microbats, many of which are referred to as sac-winged or sheath-tailed bats. They are widely distributed in tropical and subtropical regions around the world. The earliest fossil records are from the Eocene. Descr ...
s) Superfamily Rhinopomatoidea * Family Rhinopomatidae ( mouse-tailed bats) * Family Craseonycteridae (bumblebee bat or Kitti's hog-nosed bat) Superfamily
Rhinolophoidea Rhinolophoidea is a superfamily of bats. It contains the following families: Craseonycteridae, Hipposideridae, Megadermatidae, Rhinolophidae, Rhinonycteridae, and Rhinopomatidae. It is one of two superfamilies that comprise the suborder Yinpt ...
* Family Rhinolophidae (
horseshoe bat 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 ...
s) * Family Nycteridae ( hollow-faced bats or
slit-faced bat ''Nycteris'' comprises a genus of bats commonly called slit-faced or hollow-faced bats. They are grouped in the family Nycteridae. The bats are found in East Malaysia, Indonesia, and many parts of Africa. Description They are small bats, from i ...
s) * Family
Megadermatidae Megadermatidae, or false vampire bats, are a family of bats found from central Africa, eastwards through southern Asia, and into Australia. They are relatively large bats, ranging from 6.5 cm to 14 cm in head-body length. They have la ...
(
false vampire The spectral bat (''Vampyrum spectrum''), also called the great false vampire bat or Linnaeus's false vampire bat, is a large, carnivorous leaf-nosed bat found in Mexico, Central America, and South America. It is the only member of the genus '' ...
s) Superfamily
Vespertilionoidea Vespertilionoidea is a superfamily of bats containing five families: Cistugidae, Miniopteridae, Molossidae, Natalidae, and Vespertilionidae. It is one of three superfamilies in the suborder Yangochiroptera, the others being Noctilionoidea and E ...
* Family
Vespertilionidae 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 famili ...
(
vesper bat 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 famili ...
s or
evening bat The evening bat (''Nycticeius humeralis'') is a species of bat in the vesper bat family that is native to North America. Hunting at night, they eat beetles, moths, and other flying insects. Description The evening bat is a small bat weigh ...
s) Superfamily Molossoidea * Family Molossidae (
free-tailed bat The Molossidae, or free-tailed bats, are a family of bats within the order Chiroptera. The Molossidae is the fourth-largest family of bats, containing about 110 species as of 2012. They are generally quite robust, and consist of many strong-flyi ...
s) * Family
Antrozoidae Antrozoini is a tribe of bats in the subfamily Vespertilioninae of the family Vespertilionidae. It contains the pallid bat (''Antrozous pallidus''), Van Gelder's bat (''Bauerus dubiaquercus''), the genus ''Rhogeessa'', and the fossil '' Anzanyc ...
( pallid bats) Superfamily Nataloidea * Family
Natalidae The family Natalidae, or funnel-eared bats, are found from Mexico to Brazil and the Caribbean islands. The family has three genera, '' Chilonatalus'', '' Natalus'' and '' Nyctiellus''. They are slender bats with unusually long tails and, as thei ...
(
funnel-eared bat The family Natalidae, or funnel-eared bats, are found from Mexico to Brazil and the Caribbean islands. The family has three genera, ''Chilonatalus'', ''Natalus'' and ''Nyctiellus''. They are slender bats with unusually long tails and, as their ...
s) * Family
Myzopodidae ''Myzopoda'', which has two described species, is the only genus in the bat family Myzopodidae. Myzopodidae is unique as the only family of bats presently endemic to Madagascar. However, fossil discoveries indicate that the family has an ancie ...
( sucker-footed bats) * Family Thyropteridae (
disk-winged bat Disk-winged bats are a small group of bats of the family Thyropteridae and genus Thyroptera. They are found in Central and South America, usually in moist tropical rain forests. It is a very small family, consisting of a single genus with five e ...
s) * Family
Furipteridae Furipteridae is family of bats, allying two genera of single species, '' Amorphochilus schnablii'' (smoky bat) and the type '' Furipterus horrens'' (thumbless bat). They are found in Central and South America and are closely related to the bats ...
(
smoky bat The smoky bat (''Amorphochilus schnablii'') is a species of bat in the family Furipteridae. It is the only species within the genus ''Amorphochilus''. Its natural habitat is rocky shores. It is also called the thumbless bat because its thumb i ...
s) Superfamily
Noctilionoidea Noctilionoidea is a superfamily of bats containing seven families: Thyropteridae, Furipteridae, Noctilionidae, Mormoopidae, Phyllostomidae, Myzopodidae, and Mystacinidae. It is one of three superfamilies in the suborder Yangochiroptera, the o ...
* Family Noctilionidae (
bulldog bat The bat family Noctilionidae, commonly known as bulldog bats or fishing bats, is represented by two extant species, the greater and the lesser bulldog bats, as well as at least one fossil species, '' Noctilio lacrimaelunaris'', from the Miocene ...
s or
fisherman bat The bat family Noctilionidae, commonly known as bulldog bats or fishing bats, is represented by two extant species, the greater and the lesser bulldog bats, as well as at least one fossil species, '' Noctilio lacrimaelunaris'', from the Miocene ...
s) * Family
Mystacinidae __NOTOC__ Mystacinidae is a family of unusual bats, the New Zealand short-tailed bats. There is one living genus, ''Mystacina'', with two species, one of which could have possibly become extinct in the 1960s. They are medium-sized bats, about in l ...
( New Zealand short-tailed bats) * Family
Mormoopidae The family Mormoopidae contains bats known generally as mustached bats, ghost-faced bats, and naked-backed bats. They are found in the Americas from the Southwestern United States to Southeastern Brazil. They are distinguished by the presenc ...
(
ghost-faced bat The ghost-faced bat (''Mormoops megalophylla'') is a bat in the genus ''Mormoops''. It is one of only two extant species within its genus, the other being the much smaller ''Mormoops blainvillii''. They are nocturnal and hunt using Animal echolo ...
s or
moustached bat ''Mormoops'' is a genus of bat in the family Mormoopidae. It contains the following species: * Antillean ghost-faced bat (''Mormoops blainvillii'') * Giant ghost-faced bat (†''Mormoops magna'') * Ghost-faced bat The ghost-faced bat (''Morm ...
s) * Family Phyllostomidae (
leaf-nosed bat The New World leaf-nosed bats (Phyllostomidae) are found from southern North America to South America, specifically from the Southwest United States to northern Argentina. They are ecologically the most varied and diverse family within the order ...
s)


References


External links


Bat World Sanctuary
(''see "Recent publications"'')
Bat Conservation International
* {{Authority control Bat taxonomy Animals that use echolocation Paraphyletic groups