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Animal echolocation
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==== Calls and ecology ==== Echolocating bats occupy a diverse set of ecological conditions; they can be found living in environments as different as [[Europe]] and [[Madagascar]], and hunting for food sources as different as insects, frogs, nectar, fruit, and blood. The characteristics of an echolocation call are adapted to the particular environment, hunting behavior, and food source of the particular bat. The adaptation of echolocation calls to ecological factors is constrained by the phylogenetic relationship of the bats, leading to a process known as descent with modification, and resulting in the diversity of the Chiroptera today.<ref name="Jones_2006">{{cite journal |last1=Jones |first1=G. |last2=Teeling |first2=E. |title=The evolution of echolocation in bats |journal=Trends in Ecology & Evolution |volume=21 |issue=3 |pages=149β156 |date=March 2006 |pmid=16701491 |doi=10.1016/j.tree.2006.01.001 }}</ref><!--<ref name="Grinnell 1995"/>--><!--<ref name="Zupanc 2004">{{cite book |last=Zupanc |first=GΓΌnther K. H. |date=2004 |title=Behavioral Neurobiology: An Integrative Approach |publisher=Oxford University Press |location=Oxford, UK |isbn=978-0-1987-3872-5 |pages= }}</ref>--><ref name="Fenton_1995">{{cite book |last=Fenton |first=M. B. |date=1995 |chapter=Natural History and Biosonar Signals |title=Hearing in Bats |editor1=Popper, A. N. |editor2=Fay, R. R. |publisher=Springer Verlag |location=New York |pages=37β86}}</ref><!--<ref name="Neuweiler_2003"/>--><ref name="Simmons_1980">{{cite journal |last1=Simmons |first1=J. A. |last2=Stein |first2=R. A. |year=1980 |title=Acoustic Imaging in bat sonar: echolocation signals and the evolution of echolocation | journal=Journal of Comparative Physiology A |volume=135 |issue=1 |pages=61β84 |doi=10.1007/bf00660182 |s2cid=20515827 }}</ref> Bats can inadvertently jam each other, and in some situations they may stop calling to avoid jamming.<ref name="Chiu Xian 2008">{{cite journal |last1=Chiu |first1=Chen |last2=Xian |first2=Wei |last3=Moss |first3=Cynthia F. |title=Flying in silence: Echolocating bats cease vocalizing to avoid sonar jamming |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=105 |issue=35 |pages=13116β21 |date=September 2008 |pmid=18725624 |pmc=2529029 |doi=10.1073/pnas.0804408105 |bibcode=2008PNAS..10513116C |doi-access=free }}</ref> Flying insects are a common source of food for echolocating bats and some insects (moths in particular) can hear the calls of predatory bats. However the evolution of [[Tympanal organ|hearing organs]] in moths predates the origins of bats, so while many moths do listen for approaching bat echolocation their ears did not originally evolve in response to selective pressures from bats.<ref>{{Cite journal |last1=Kawahara |first1=Akito Y. |last2=Plotkin |first2=David |last3=Espeland |first3=Marianne |last4=Meusemann |first4=Karen |last5=Toussaint |first5=Emmanuel F. A. |last6=Donath |first6=Alexander |last7=Gimnich |first7=France |last8=Frandsen |first8=Paul B. |last9=Zwick |first9=Andreas |last10=dos Reis |first10=Mario |last11=Barber |first11=Jesse R. |last12=Peters |first12=Ralph S. |last13=Liu |first13=Shanlin |last14=Zhou |first14=Xin |last15=Mayer |first15=Christoph |date=2019-11-05 |title=Phylogenomics reveals the evolutionary timing and pattern of butterflies and moths |journal=Proceedings of the National Academy of Sciences |language=en |volume=116 |issue=45 |pages=22657β22663 |doi=10.1073/pnas.1907847116 |doi-access=free |issn=0027-8424 |pmc=6842621 |pmid=31636187|bibcode=2019PNAS..11622657K }}</ref> These moth adaptations provide [[Evolutionary pressure |selective pressure]] for bats to improve their insect-hunting systems and this cycle culminates in a moth-bat "[[evolutionary arms race]]".<ref>{{cite journal |last1=Goerlitz |first1=Holger R. |last2=ter Hofstede |first2=Hannah M. |last3=Zeale |first3=Matt R. K. |last4=Jones |first4=Gareth |last5=Holderied |first5=Marc W. |title=An aerial-hawking bat uses stealth echolocation to counter moth hearing |journal=Current Biology |volume=20 |issue=17 |pages=1568β1572 |date=September 2010 |pmid=20727755 |doi=10.1016/j.cub.2010.07.046 |doi-access=free |bibcode=2010CBio...20.1568G }}</ref><ref>{{cite journal |last1=Ratcliffe |first1=John M. |last2=Elemans |first2=Coen P. H. |last3=Jakobsen |first3=Lasse |last4=Surlykke |first4=Annemarie |title=How the bat got its buzz |journal=Biology Letters |volume=9 |issue=2 |pages=20121031 |date=April 2013 |pmid=23302868 |pmc=3639754 |doi=10.1098/rsbl.2012.1031 }}</ref>
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