Editing Microbat

{{short description|Suborder of bats}}
{{Automatic taxobox
| name = Microbats
| image = Townsends in music hall.jpg
| image_caption = [[Townsend's big-eared bat]]s in a [[cave]].
| image_alt = Townsend's big-eared bat
| taxon = Microchiroptera
| authority = [[George Edward Dobson|Dobson]], 1875
| synonyms = 
| synonyms_ref = 
| subdivision_ranks = Superfamilies
| subdivision = *[[Emballonuroidea]]
*[[Rhinopomatoidea]]
*[[Rhinolophoidea]]
*[[Vespertilionoidea]]
*[[Molossoidea]]
*[[Nataloidea]]
*[[Noctilionoidea]]
}}

'''Microbats''' constitute the [[suborder]] '''Microchiroptera''' within the [[Order (biology)|order]] Chiroptera ([[bat]]s). Bats have long been differentiated into [[Megachiroptera]] (megabats) and Microchiroptera, based on their size, the use of [[Animal echolocation|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.<ref name="microbat paraphyly" />

==Characteristics==
Microbats are {{convert|4|to(-)|16|cm|in|abbr=on}} long.<ref>{{cite journal | last1=Whitaker | first1=John O. | last2=Dannelly | first2=H. Kathleen | last3=Prentice | first3=David A. | title=Chitinase in Insectivorous Bats | journal=Journal of Mammalogy | volume=85 | issue=1 | date=2004 | issn=0022-2372 | doi=10.1644/1545-1542(2004)085<0015:CIIB>2.0.CO;2 | doi-access=free | pages=15–18 | url=https://academic.oup.com/jmammal/article-pdf/85/1/15/7023743/85-1-15.pdf}}</ref> 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 bat]]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.<ref>{{Cite book |last=Nowak |first=Ronald M. |url=https://books.google.com/books?id=OmkObkVIJ0EC&q=Walker%27s%2520Bats%2520of%2520the%2520World%252C%2520Ronald%2520M.%2520Nowak%2520%281994%29&pg=PA1 |title=Walker's Bats of the World |date=1994 |publisher=JHU Press |isbn=978-0-8018-4986-2 |language=en}}</ref> 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).<ref>{{Cite book |last1=Comus |first1=Patricia Wentworth |url=https://books.google.com/books?id=cbQwDwAAQBAJ&dq=A+Natural+History+of+the+Sonoran+Desert,+Edited+by+Steven+Phillips+and+Patricia+Comus,+University+of+California+Press,+Berkeley+p.+464&pg=PA3 |title=A Natural History of the Sonoran Desert |last2=Phillips |first2=Steven John |last3=Dimmitt |first3=Mark Alan |last4=Brewer |first4=Linda M. |date=2015-11-17 |publisher=Univ of California Press |isbn=978-0-520-28747-1 |pages=464 |language=en}}</ref> 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.

=== Differences from megabats ===
* Microbats use [[Animal echolocation|echolocation]], whereas megabats do not typically. (The Egyptian fruit bat ''[[Rousettus egyptiacus]]'' 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 (ear)|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 ===
[[File:Dilambdodont teeth pattern (microbat).png|thumb|alt=The ventral view of microbat teeth|Ventral view of a free-tailed microbat (genus ''Tadarida'') skull displaying a dilambdodont teeth pattern. Specimen from the Pacific Lutheran University Natural History collection.]]
[[File:Microbat canine teeth.png|thumb|alt=The frontal view of microbat teeth|Frontal view of a free-tailed microbat (genus ''Tadarida'') skull displaying the canine teeth. Specimen from the Pacific Lutheran University Natural History collection.]]

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.<ref>{{Cite journal |last=Evans |first=Alistair R. |date=2005-04-25 |title=Connecting morphology, function and tooth wear in microchiropterans: FUNCTIONAL DENTAL MORPHOLOGY IN MICROCHIROPTERANS |url=https://academic.oup.com/biolinnean/article-lookup/doi/10.1111/j.1095-8312.2005.00474.x |journal=Biological Journal of the Linnean Society |language=en |volume=85 |issue=1 |pages=81–96 |doi=10.1111/j.1095-8312.2005.00474.x}}</ref> In comparison to megabats which feed only on fruit and nectar, microbats illustrate a range of diets and have been classified as [[insectivore]]s, [[carnivore]]s, [[Hematophagy|sanguinivores]], [[frugivore]]s, and [[nectarivore]]s.<ref name=":3">{{cite book |last=Freeman |first=Patricia W. |chapter=Form, Function, and Evolution in Skulls and Teeth of Bats |year=1998 |title=Bat Biology and Conservation |editor1=Thomas H. Kunz |editor2=Paul A. Racey |location=Washington, DC. |publisher=Smithsonian Institution Press |pages= 140–156 |chapter-url=https://digitalcommons.unl.edu/cgi/viewcontent.cgi?article=1009&context=natrespapers |chapter-format=PDF}}</ref> 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.<ref>{{cite web | last1=Wund | first1=Matthew | last2=Myers | first2=Phil | title=Chiroptera (bats) | website=Animal Diversity Web | date=2005-08-16 | url=https://animaldiversity.org/accounts/Chiroptera/ }}</ref> 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.<ref>{{Cite journal |last=Freeman |first=Patricia W. |date=March 1988 |title=Frugivorous and animalivorous bats (Microchiroptera): dental and cranial adaptations |url=https://academic.oup.com/biolinnean/article-lookup/doi/10.1111/j.1095-8312.1988.tb00811.x |journal=Biological Journal of the Linnean Society |language=en |volume=33 |issue=3 |pages=249–272 |doi=10.1111/j.1095-8312.1988.tb00811.x|url-access=subscription }}</ref> Frugivorous microbats also possess a different pattern on their molars compared to carnivorous, insectivorous, nectarivorous, and sanguinivorous microbats.<ref name=":3" /> 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.<ref name=":3" />

== Echolocation ==
{{main|Animal echolocation}}
[[Animal 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 [[Ultrasound|ultrasonic waves]] via their [[larynx]].<ref name=":0">{{Cite journal|last1=Davies|first1=Kalina T. J.|last2=Maryanto|first2=Ibnu|last3=Rossiter|first3=Stephen J.|date=2013-01-01|title=Evolutionary origins of ultrasonic hearing and laryngeal echolocation in bats inferred from morphological analyses of the inner ear|journal=Frontiers in Zoology|volume=10|issue=1|pages=2|doi=10.1186/1742-9994-10-2|issn=1742-9994|pmc=3598973|pmid=23360746 |doi-access=free }}</ref><ref name=":1">{{Cite journal|last1=Reichard|first1=Jonathan D.|last2=Fellows|first2=Spenser R.|last3=Frank|first3=Alexander J.|last4=Kunz|first4=Thomas H.|date=2010-11-01|title=Thermoregulation during Flight: Body Temperature and Sensible Heat Transfer in Free-Ranging Brazilian Free-Tailed Bats (Tadarida brasiliensis)|journal=Physiological and Biochemical Zoology|volume=83|issue=6|pages=885–897|doi=10.1086/657253|pmid=21034204|s2cid=1028970|issn=1522-2152}}</ref> 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.<ref name=":2">{{Cite book|title=Handbook of Behavioral Neuroscience|last1=Berke|first1=Gerald S.|last2=Long|first2=Jennifer L.|date=2010-01-01|publisher=Elsevier|editor-last=Brudzynski|editor-first=Stefan M.|series=Handbook of Mammalian VocalizationAn Integrative Neuroscience Approach|volume=19|pages=419–426|doi=10.1016/B978-0-12-374593-4.00038-3|chapter=Functions of the larynx and production of sounds|isbn=9780123745934}}</ref>

=== Production of ultrasonic waves ===
Most microbats generate ultrasound with their larynx and emit the sound through their nose or mouth.<ref>{{Cite journal|last1=Hedenström|first1=Anders|last2=Johansson|first2=L. Christoffer|date=2015-03-01|title=Bat flight: aerodynamics, kinematics and flight morphology|journal=Journal of Experimental Biology|volume=218|issue=5|pages=653–663|doi=10.1242/jeb.031203|issn=0022-0949|pmid=25740899|doi-access=free|bibcode=2015JExpB.218..653H }}</ref> 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 folds|vocal cords]] and forms the passageway for the expiratory air that will produce sound.<ref>{{Cite journal|last1=Holbrook|first1=K A|last2=Odland|first2=G F|date=1978-05-01|title=A collagen and elastic network in the wing of the bat.|journal=Journal of Anatomy|volume=126|issue=Pt 1|pages=21–36|issn=0021-8782|pmc=1235709|pmid=649500}}</ref> Microbat {{Audio|Ultrasonic_bat_calls.ogg|calls}} 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&nbsp;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.<ref>{{Cite journal |last1=Liu |first1=Meiling |last2=Wang |first2=Changliang |last3=Huo |first3=Lifang |last4=Cao |first4=Jie |last5=Mao |first5=Xiuguang |last6=He |first6=Ziqing |last7=Hu |first7=Chuanxia |last8=Sun |first8=Haijian |last9=Deng |first9=Wenjun |last10=He |first10=Weiya |last11=Chen |first11=Yifu |last12=Gu |first12=Meifeng |last13=Liao |first13=Jiayu |last14=Guo |first14=Ning |last15=He |first15=Xiangyang |date=July 2024 |title=Complexin-1 enhances ultrasound neurotransmission in the mammalian auditory pathway |url=https://www.nature.com/articles/s41588-024-01781-z |journal=Nature Genetics |volume=56 |issue=7 |pages=1503–1515 |doi=10.1038/s41588-024-01781-z |pmid=38834904 |issn=1546-1718|url-access=subscription }}</ref>

=== Laryngeally echolocating microbats ===
[[File:BIOL352-WikiPic-2.jpg|thumb|alt=The skull of a microbat|Ventral view of a Florida Freetail bat (''Tadarida cyanocephala'') skull, highlighting both the stylohyal and tympanic bones. Specimen from the Pacific Lutheran University Natural History collection.]]

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 [[Ultrasound|ultrasonic waves]] by clapping their wings, clicking their tongues, or using their nose.<ref name=":0" /> 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 [[Cranium|cranial]] end of the [[trachea]] and is surrounded by [[cricothyroid muscle]]s and [[thyroid cartilage]]. For reference, in [[human]]s, 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.<ref>{{Cite journal|last1=Simmons|first1=Nancy B.|last2=Seymour|first2=Kevin L.|last3=Habersetzer|first3=Jörg|last4=Gunnell|first4=Gregg F.|date=2010-08-19|title=Inferring echolocation in ancient bats|journal=Nature|volume=466|issue=7309|pages=E8|doi=10.1038/nature09219|pmid=20724993|issn=0028-0836|bibcode=2010Natur.466E...8S|doi-access=free}}</ref> A characteristic of laryngeally echolocating microbats that distinguishes them from other echolocating microbats is the articulation of their [[stylohyal bone]] with their [[Tympanic part of the temporal bone|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.<ref name=":1" />

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 [[Ultrasound|ultrasonic waves]] at a [[neuron]]al level, in order to accurately obtain information about their surrounding environment and orientation in it.<ref name=":0" /> 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]].<ref name=":2" /> Furthermore, the stylohyal bones connect the larynx to the tympanic bones via a [[cartilaginous]] or [[Fibrous joint|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 muscle]]s, 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.<ref name=":0" />

== Classification ==
[[File:Bats Romania 2003.jpg|thumb|A series of Romanian postage stamps. The bats depicted are (from top to bottom and left to right): the [[greater mouse-eared bat]], the [[lesser horseshoe bat]], the [[brown long-eared bat]], the [[common pipistrelle]], the [[greater noctule bat]], and the [[barbastelle]], Romanian post miniature sheet, 2003]]

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.<ref name="microbat paraphyly">{{cite journal|last1=Teeling|first1=E. C.|last2=Madsen|first2=O.|last3=Van de Bussche|first3=R. A.|last4=de Jong|first4=W. W.|last5=Stanhope|first5=M. J.|last6=Springer|first6=M. S.|title=Microbat paraphyly and the convergent evolution of a key innovation in Old World rhinolophoid microbats|journal=Proceedings of the National Academy of Sciences of the United States of America|date=2002|volume=99|issue=3|pages=1431–1436|doi=10.1073/Pnas.022477199|pmid=11805285|pmc=122208|bibcode=2002PNAS...99.1431T}}</ref>

This is the classification according to Simmons and Geisler (1998):

Superfamily [[Emballonuroidea]]
* Family [[Emballonuridae]] ([[sac-winged bat]]s or [[sheath-tailed bat]]s)
Superfamily [[Rhinopomatoidea]]
* Family [[Rhinopomatidae]] ([[mouse-tailed bat]]s)
* Family Craseonycteridae (bumblebee bat or [[Kitti's hog-nosed bat]])
Superfamily [[Rhinolophoidea]]
* Family Rhinolophidae ([[horseshoe bat]]s)
* Family [[Nycteridae]] ([[hollow-faced bat]]s or [[slit-faced bat]]s)
* Family [[Megadermatidae]] ([[false vampire]]s)
Superfamily [[Vespertilionoidea]]
* Family [[Vespertilionidae]] ([[vesper bat]]s or [[evening bat]]s)
Superfamily [[Molossoidea]]
* Family [[Molossidae]] ([[free-tailed bat]]s)
* Family [[Antrozoidae]] ([[pallid bat]]s)
Superfamily [[Nataloidea]]
* Family [[Natalidae]] ([[funnel-eared bat]]s)
* Family [[Myzopodidae]] ([[sucker-footed bat]]s)
* Family [[Thyroptera|Thyropteridae]] ([[disk-winged bat]]s)
* Family [[Furipteridae]] ([[smoky bat]]s)
Superfamily [[Noctilionoidea]]
* Family [[Noctilionidae]] ([[bulldog bat]]s or [[fisherman bat]]s)
* Family [[Mystacinidae]] ([[New Zealand short-tailed bat]]s)
* Family [[Mormoopidae]] ([[ghost-faced bat]]s or [[moustached bat]]s)
* Family [[Phyllostomidae]] ([[leaf-nosed bat]]s)

==References==
{{Reflist}}

== External links ==
{{Commons category|Microchiroptera|Microbats}}
* [http://www.batworld.org Bat World Sanctuary]
* [https://web.archive.org/web/20051103015511/http://www.uni-tuebingen.de/tierphys/Kontakt/mitarbeiter_seiten/dietz.htm Illustrated Identification key to the bats of Europe]  (''see "Recent publications"'')
* [http://www.batcon.org Bat Conservation International]
* {{UCSC genomes|myoLuc2}}

{{Taxonbar|from=Q971343}}
{{Authority control}}

[[Category:Animals that use echolocation]]
[[Category:Bat taxonomy]]
[[Category:Paraphyletic groups]]
<!-- [[Category:Animal suborders]] already present in "microchiroptera" redirect -->