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The '''aye-aye''' ('''''Daubentonia madagascariensis''''') is a long-fingered [[lemur]], a [[Strepsirrhini|strepsirrhine]] [[primate]] native to [[Madagascar]] with [[rodent]]-like teeth that perpetually grow<ref name=":0">{{Cite journal|title = Primates Prosimiens|last1 = Petter|first1 =J. |date = 1977|journal = ORSTOM|last2 = Albignac|first2 =R. |last3 = Rumpler|first3 =Y.}}</ref> and a special thin middle finger that they can use to catch grubs and larvae out of tree trunks.

It is the world's largest [[Nocturnality|nocturnal]] primate.<ref name="nationalgeographic" /> It is characterized by its unusual method of finding food: it taps on trees to find [[Larva|grubs]], then gnaws holes in the wood using its forward-slanting incisors to create a small hole into which it inserts its narrow middle finger to pull the grubs out. This foraging method is called ''percussive foraging'', and takes up 5–41% of foraging time.<ref name="Erickson1998">{{cite journal|last1= Erickson|first1=C.J. |last2= Nowicki |first2=S. |last3= Dollar|first3=L. |last4= Goehring|first4=N. |s2cid= 27737088|title= Percussive Foraging: Stimuli for Prey Location by Aye-Ayes (''Daubentonia madagascariensis'')|journal= International Journal of Primatology|year= 1998|volume= 19|issue= 1|page= 111|doi= 10.1023/A:1020363128240}}<!--|access-date=6 May 2013--></ref><ref>{{Cite journal |title = Adaptations in the ayeaye: A review|last1 = Sterling|first1 =E.J. |date = 2006|journal = Lemurs: Ecology and Adaptations|last2 = McCreless|first2 =E.E. |publisher = Gould L, Sauther ML|url=https://www.researchgate.net/publication/226739621}}</ref> The only other living mammal species known to find food in this way are the [[striped possum]] and trioks (genus ''[[Dactylopsila]]'') of northern Australia and New Guinea, which are marsupials.{{Sfn|Sterling|2003|p=1348}} From an ecological point of view, the aye-aye fills the niche of a [[woodpecker]], as it is capable of penetrating wood to extract the invertebrates within.{{Sfn|Piper|2007|p={{page needed|date=December 2020}}}}{{Sfn|Beck|2009}}

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Due to its [[Cladistics|derived]] [[morphology (biology)|morphological]] features, the classification of the aye-aye was debated following its discovery. The possession of continually growing [[incisor]]s (front teeth) parallels those of [[rodent]]s, leading early naturalists to mistakenly classify the aye-aye within the mammalian order [[Rodent]]ia{{Sfn|Ankel-Simons|2007|p=257}} and as a squirrel, due to its toes, hair coloring, and tail. However, the aye-aye is also similar to felines in its head shape, eyes, ears and nostrils.<ref name=Science>{{cite journal|title=The Aye-Ayes or Cheiromys of Madagascar |journal=Science |date=3 November 1881 |volume=2 |issue=75 |pages=574–576 |doi=10.1126/science.os-2.76.574 |pmid=17791380}}{{Dead link|date=July 2023}}</ref>

The aye-aye's classification with the order Primates has been just as uncertain. It has been considered a highly derived member of the family [[Indridae]], a [[basal (phylogenetics)|basalbasally diverging]] branch of the [[strepsirrhine]] suborder, and of indeterminate relation to all living primates.{{Sfn|Yoder|Vilgalys|Ruvolo|1996}} In 1931, Anthony and Coupin classified the aye-aye under infraorder '''Chiromyiformes''', a sister group to the other strepsirrhines. [[Colin Groves]] upheld this classification in 2005 because he was not entirely convinced the aye-aye formed a [[clade]] with the rest of the Malagasy lemurs.{{Sfn|Groves|2005|p=121}}

However, molecular results have consistently placed ''Daubentonia'' as the most [[basal (phylogenetics)|basal]]<nowiki/>ly diverging of [[lemurs]].{{Sfn|Yoder|Vilgalys|Ruvolo|1996}}<ref name="Del Pero1995">{{cite journal|last1= Del Pero|first1=M. |last2= Crovella|first2=S. |last3= Cervella|first3=P. |last4= Ardito|first4=G. |last5= Rumpler|first5=Y. |s2cid= 25558426|title= Phylogenetic relationships among Malagasy lemurs as revealed by mitochondrial DNA sequence analysis |journal= Primates|volume= 36|issue= 3|year= 1995|pages= 431–440|doi= 10.1007/BF02382865}}</ref><ref name="Porter1995">{{cite journal|display-authors=4 |last1= Porter|first1=C.A. |last2= Sampaio|first2=I. |last3= Schneider|first3=H. |last4= Schneider|first4=M.P.C. |last5= Czelusniak|first5=J. |last6= Goodman|first6=M. |s2cid= 9509799|title= Evidence on primate phylogeny from ε-globin gene sequences and flanking regions|journal= Journal of Molecular Evolution |volume= 40|issue= 1|year= 1995|pages= 30–55|doi= 10.1007/BF00166594|pmid= 7714911|bibcode= 1995JMolE..40...30P}}</ref><ref name="Roos2004">{{cite journal|last1= Roos|first1=C. |last2= Schmitz|first2=J. |last3= Zischler|first3=H. |title= Primate jumping genes elucidate strepsirrhine phylogeny |journal= Proceedings of the National Academy of Sciences |volume= 101|issue= 29|year= 2004|pages= 10650–10654|doi= 10.1073/pnas.0403852101|pmid= 15249661|pmc= 489989|bibcode= 2004PNAS..10110650R|doi-access=free }}</ref><ref name="Horvath2008">{{Cite journal |display-authors=4 |last1= Horvath |first1=J.E. |last2= Weisrock |first2=D.W. |last3= Embry |first3=S.L. |last4= Fiorentino |first4=I. |last5= Balhoff |first5=J.P. |last6= Kappeler |first6=P. |last7= Wray |first7=G.A. |last8= Willard |first8=H.F. |last9= Yoder |first9=A.D. |doi= 10.1101/gr.7265208 |title= Development and application of a phylogenomic toolkit: Resolving the evolutionary history of Madagascar's lemurs |journal= Genome Research |volume= 18 |issue= 3 |pages= 489–499 |year= 2008 |pmid= 18245770|pmc= 2259113}}</ref><ref name="Orlando2008">{{Cite journal |display-authors=4 |last1= Orlando |first1=L. |last2= Calvignac |first2=S. |last3= Schnebelen |first3=C. |last4= Douady |first4=C.J. |last5= Godfrey |first5=L.R. |last6= Hänni |first6=C. |doi= 10.1186/1471-2148-8-121 |title= DNA from extinct giant lemurs links archaeolemurids to extant indriids |journal= BMC Evolutionary Biology |volume= 8 |issue= 121 |pages= 121 |year= 2008 |pmid= 18442367|pmc= 2386821 |doi-access= free |bibcode= 2008BMCEE...8..121O }}</ref><ref name="Brosius2011">{{cite journal|display-authors=4 |last1= Brosius|first1=J. |last2= Perelman|first2=P. |last3= Johnson|first3=W.E. |last4= Roos|first4=C. |last5= Seuánez|first5=H.N. |last6= Horvath|first6=J.E. |last7=Moreira|first7=M.A.M. |last8= Kessing|first8=B. |last9= Pontius|first9=J. |last10= Roelke|first10=M. |last11= Rumpler|first11=Y. |last12= Schneider|first12=M.P.C. |last13= Silva|first13=A. |last14= O'Brien|first14=S.J. |last15= Pecon-Slattery|first15=J. |title=A Molecular Phylogeny of Living Primates |journal= PLOS Genetics|volume= 7|issue= 3|year= 2011|pages= e1001342|doi= 10.1371/journal.pgen.1001342|pmid= 21436896|pmc= 3060065|doi-access= free}}</ref><ref name="Perry2012">{{cite journal|display-authors=4 |last1= Perry|first1=G.H. |last2= Reeves|first2=D. |last3= Melsted|first3=P. |last4= Ratan|first4=A. |last5= Miller|first5=W. |last6= Michelini|first6=K. |last7= Louis|first7=E.E. |last8= Pritchard|first8=J.K. |last9= Mason|first9=C.E. |last10= Gilad|first10=Y. |title=A Genome Sequence Resource for the Aye-Aye (''Daubentonia madagascariensis''), a Nocturnal Lemur from Madagascar |journal= Genome Biology and Evolution |volume= 4|issue= 2|year= 2012|pages= 126–135|doi= 10.1093/gbe/evr132|pmid= 22155688|pmc= 3273163|doi-access= free}}</ref> The most parsimonious explanation for this is that all lemurs are derived from a single ancestor that [[Oceanic dispersal|rafted]] from Africa to Madagascar during the [[Paleogene]].<ref name="Roos2004"/><ref name=Sellers>{{cite web |title=Primate Evolution |url=http://homepage.mac.com/wis/Personal/lectures/human-origins/PrimateEvolution.pdf |access-date=2008-10-23 |last=Sellers |first=Bill |date=2000-10-20 |publisher=University of Edinburgh |pages=13–17| archive-url=https://web.archive.org/web/20090225011846/http://homepage.mac.com/wis/Personal/lectures/human-origins/PrimateEvolution.pdf |archive-date=2009-02-25}}</ref><ref name= "Ali&Huber">{{Cite journal |last=Ali |first=J.R. |author2=Huber, M. |s2cid=4333977 |title=Mammalian biodiversity on Madagascar controlled by ocean currents |journal=[[Nature (journal)|Nature]] |volume=463 |issue=4 |pages=653–656 |date= Feb 2010 |doi=10.1038/nature08706 |pmid=20090678 |bibcode=2010Natur.463..653A}}</ref> Similarities in dentition between aye-ayes and several African primate fossils (''[[Plesiopithecus]]'' and ''[[Propotto]]'') have led to the alternate theory that the ancestors of aye-ayes colonized Madagascar separately from other lemurs.<ref name="Gunnell2018">{{cite journal|display-authors=4 |last1= Gunnell|first1=G.F. |last2= Boyer|first2=D.M. |last3= Friscia|first3=A.R. |last4= Heritage|first4=S. |last5= Manthi|first5=F.K. |last6= Miller|first6=E.R. |last7= Sallam|first7=H.M. |last8= Simmons|first8=N.B. |last9= Stevens|first9=N.J. |last10= Seiffert|first10=E.R. |title= Fossil lemurs from Egypt and Kenya suggest an African origin for Madagascar's aye-aye|journal= Nature Communications|volume= 9|issue= 1|year= 2018|page= 3193|doi= 10.1038/s41467-018-05648-w|pmid= 30131571|pmc= 6104046|bibcode= 2018NatCo...9.3193G|doi-access= free}}</ref> In 2008, [[Russell Mittermeier]], Colin Groves, and others ignored addressing higher-level taxonomy by defining lemurs as monophyletic and containing five living families, including Daubentoniidae.<ref name="Mittermeier_et_al_2008"/>

Further evidence indicating that the aye-aye belongs in the superfamily [[Lemuroidea]] can be inferred from the presence of [[Petrous part of the temporal bone|petrosal]] bullae encasing the [[ossicles]] of the ear.{{Sfn|Ankel-Simons|2007|p=257}} The aye-ayes are also similar to lemurs in their shorter back legs.<ref name=Science/>

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[[File:Aye-Aye Daubentonia madagascariensis in Stockholm.jpg|thumb|upright|Skeleton]]

[[File:Aye-Aye Daubentonia madagascariensis in Copenhagen (Left Hand).jpg|thumb|upright|Closeup of the hand showing the elongated digits and the thinner third digit]]

[[File:AyeDaubentonia ayemadagascariensis 3d scan Natural History Museum University of Pisa C 1065.stl|thumb|3d scan of the skull of ''Daubentonia madagascariensis'']]

[[File:Daubentonia madagascariensis 3d scan Natural History Museum University of Pisa C 946.stl|thumb|3d scan of the left hand of ''Daubentonia madagascariensis'']]

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Young aye-ayes typically are silver colored on their front and have a stripe down their back. However, as the aye-ayes begin to reach maturity, their bodies will be completely covered in thick fur and are typically not one solid color. On the head and back, the ends of the hair are typically tipped with white while the rest of the body will ordinarily be a yellow and/or brown color.

Among the aye-aye's signature traits are its fingers.<ref name="Perry2012" /> The third finger, which is much thinner than the others, is used for extracting grubs and insects out of trees, using the hooked nail. The finger is unique in the animal kingdom in that it possesses a ball-and-socket [[metacarpophalangeal joint]],<ref>{{Cite journal |title= Anatomy of the hand and arm in Daubentonia madagascariensis: a functional and phylogenetic outlook |last= Soligo |first=C. |s2cid= 25535277 |date= 2005 |journal= Folia Primatol |volume=76 |issue=5 |pages=262–300 |doi=10.1159/000088034 |pmid=16230860 |url=https://www.karger.com/Article/Abstract/88034}}</ref> can reach the throat through a [[nostril]] and is used for [[Nose picking|picking one's nose]] and [[eating mucus]] ([[mucophagy]]) so harvested from inside the nose.<ref>{{cite news |last1=Davis |first1=Nicola |title=Researchers reveal secret of aye-ayes' long middle finger {{!}} Video shows captive Madagascan primates using elongated finger to pick nose and eat the mucus |url=https://www.theguardian.com/world/2022/oct/27/researchers-reveal-secret-aye-aye-long-middle-finger-pick-nose |access-date=30 October 2022 |work=The Guardian |date=26 October 2022 |language=en}}</ref><ref>{{cite journal |last1=Fabre |first1=A.-C. |last2=Portela Miguez |first2=R. |last3=Wall |first3=C. E. |last4=Peckre |first4=L. R. |last5=Ehmke |first5=E. |last6=Boistel |first6=R. |title=A review of nose picking in primates with new evidence of its occurrence in Daubentonia madagascariensis |journal=Journal of Zoology |date=26 October 2022 |volume=319 |issue=2 |pages=91–98 |doi=10.1111/jzo.13034 |s2cid=253169043 |language=en |issn=0952-8369|doi-access=free }}</ref> The aye-aye has also evolved a sixth digit, a [[pseudothumb]], to aid in gripping.<ref name= "Hartstone‐Rose2019Hartstone-Rose2019">{{cite journal |last1= Hartstone-Rose |first1=A. |last2= Dickinson |first2=E. |last3= Boettcher |first3=M.L. |last4= Herrel |first4=A. |title=A primate with a panda's thumb: The anatomy of the pseudothumb of ''Daubentonia madagascariensis'' |journal= American Journal of Physical Anthropology |volume=171 |issue=1 |pages=8–16 |year=2020 |doi= 10.1002/ajpa.23936 |pmid=31633197|doi-access=free }}</ref>

The complex geometry of ridges on the inner surface of aye-aye ears helps to sharply focus not only echolocation signals from the tapping of its finger, but also to passively listen for any other sound produced by the prey. These ridges can be regarded as the acoustic equivalent of a [[Fresnel lens]], and may be seen in a large variety of unrelated animals, such as [[lesser galago]], [[bat-eared fox]], [[mouse lemur]], and others.

Females have two nipples located in the region of the groin.<ref>{{citationCite book needed|datelast=FebruaryOstrom 2024|first=Homer |title=The Breast And its Surgical Disease |publisher=J.N. Stoddart & Co. |year=1877}}</ref> The male's genitalia are similar to those of [[canid]]s, with a large [[prostate]] and long [[baculum]].<ref>{{Cite book |last=Dixson |first=Alan F. |url=https://books.google.com/books?id=Z9N_BokMAMkC&pg=PA321 |title=Primate Sexuality: Comparative Studies of the Prosimians, Monkeys, Apes, and Humans |date=2012-01-26 |publisher=OUP Oxford |isbn=978-0-19-954464-6 |language=en}}</ref>

== Behaviour and lifestyle ==

The aye-aye is a nocturnal and arboreal animal meaning that it spends most of its life high in the trees. Although they are known to come down to the ground on occasion, aye-ayes sleep, eat, travel and mate in the trees and are most commonly found close to the canopy where there is plenty of cover from the dense foliage. During the day, aye-ayes sleep in spherical nests in the forks of tree branches that are constructed out of leaves, branches and vines before emerging after dark to begin their hunt for food. Aye-aye are solitary animals that mark their large home range with scent. The smaller territories of females often overlap those of at least a couple of males. Male aye-ayes tend to share their territories with other males and are even known to share the same nests (although not at the same time), and can seemingly tolerate each other until they hear the call of a female that is looking for a mate.{{cn|date=October 2023}}

Mating season extends throughout the year, with females typically starting to breed at the age of three or four. They give birth to one offspring every two to three years.<ref name=IUCN/> During the period of parenting, a female becomes the dominant figure over males, likely to secure better access to food while caring for her young. The infant remains in a nest for up to two months before venturing out, but it takes another seven months before the young aye-aye can maneuver the canopy as skillfully as an adult.{{cn|date<ref name=October":0" 2023}}/>

=== Diet and foraging ===

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=== Folk belief ===

The aye-aye is often viewed as a harbinger of evil and death and killed on sight. Others believe, if one points its narrowest finger at someone, they are marked for death. Some say that the appearance of an aye-aye in a village predicts the death of a villager, and the only way to prevent this is to kill it. The [[Sakalava weaver]] people go so far as to claim aye-ayes sneak into houses through the thatched roofs and murder the sleeping occupants by using their middle fingers to puncture their victims' [[aorta]].{{Sfn|Piper|2007 |p={{page needed|date=December 2020}}}}

The pig

-aye is often viewed as a harbinger of evil and death and killed on sight. Others believe, if one points its narrowest finger at someone, they are marked for death. Some say that the appearance of an aye-aye in a village predicts the death of a villager, and the only way to prevent this is to kill it. The [[Sakalava weaver]] people go so far as to claim aye-ayes sneak into houses through the thatched roofs and murder the sleeping occupants by using their middle fingers to puncture their victims' [[aorta]].{{Sfn|Piper|2007 |p={{page needed|date=December 2020}}}}

=== Captive breeding ===

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{{Reflist|colwidth=30em|refs=

<ref name=IUCN>{{cite iucn | pageauthor=Louis, eE.E.T6302A115560793 | titleauthor2=Sefczek, ''Daubentonia madagascariensis''T.M. | vauthorsauthor3= Louis EERandimbiharinirina, SefczekD.R. TM|author4=Raharivololona, Randimbiharinirina DR, Raharivololona B,. |author5=Rakotondrazandry JN, J.N. |author6=Manjary, D,. |author7=Aylward, M,. |author8=Ravelomandrato, F. | year= 2020 | access-datetitle=''Daubentonia 18madagascariensis'' July|page=e.T6302A115560793 |doi=10.2305/IUCN.UK.2020-2.RLTS.T6302A115560793.en}}</ref>

<ref name="2012Dunkel_et_al">{{cite journal | last1 = Dunkel | first1 =A.R. | last2 = Zijlstra | first2 =J.S. | last3 = Groves | first3 =C.P. | title = Giant rabbits, marmosets, and British comedies: etymology of lemur names, part 1 | url = http://www.aeecl.org/lemurnews/lemurnews2011_16.pdf | journal = Lemur News | volume = 16 | year = 2011–2012 | issn = 1608-1439 | pages = 64–70 | access-date = 2014-10-17 | archive-url = https://web.archive.org/web/20161106163717/http://www.aeecl.org/lemurnews/lemurnews2011_16.pdf | archive-date = 2016-11-06 | url-status = dead }}</ref>