View source for Myotragus

{{Short description|Extinct genus of mammals}}
{{Use dmy dates|date=October 2019}}
{{Automatic taxobox
| fossil_range = [[Pliocene]]–[[Holocene]] {{fossil range|5.3|0.0046}}
| image = Sóller. Museu Balear de Ciències Naturals. Myotragus balearicus (cropped).jpg
| image_upright = 1.2
| image_caption = Skeleton of ''Myotragus balearicus''
| taxon = Myotragus
| authority = [[Dorothea Bate|Bate]], 1909
| type_species = †'''''Myotragus balearicus'''''
| type_species_authority = Bate, 1909
| subdivision_ranks = Other species
| subdivision = *'''''M. palomboi''''' <small>Bover, Quintana & Alcover, 2010</small>
*'''''M. pepgonellae''''' <small>Moyà-Solà & Pons-Moyà, 1982</small>
*'''''M. antiquus''''' <small>Pons-Moyà, 1977</small>
*'''''M. kopperi''''' <small>Moya & Pons, 1980</small>
*'''''M. batei''''' <small>Crusafont & Angel, 1966</small>
}}

'''''Myotragus''''' ([[Neo-Latin]], derived from the [[Ancient Greek language|Greek]]: {{lang|grc|μῦς}}, {{lang|grc|τράγος}}  "mouse-goat") is an extinct [[genus]] of goat-antelope in the tribe [[Caprini]] which lived on the [[Balearic Islands]] of [[Mallorca]] and [[Menorca]] in the western Mediterranean until its [[extinction]] around 4,500 years ago.<ref name="Bate1909">{{cite book|author=Karolyn Shindler|title=Discovering Dorothea : The Life of the Pioneering Fossil-Hunter Dorothea Bate|year=2005|publisher=HarperCollins |isbn=978-0002571388}}</ref> The fossil record of ''Myotragus'' on the Balearic Islands extends over 5 million years back to the early [[Pliocene]] on Mallorca, where it presumably arrived after the evaporation of the Mediterranean Sea during the [[Messinian salinity crisis|Messinian Salinity Crisis]] (around 5.96-5.33 million years ago). 

''Myotragus'' is represented by six sequential [[chronospecies]] representing gradual change in morphology''.'' The youngest and best known species, ''M. balearicus,'' is noted for a number of unusual morphological adaptions, including forward facing eyes suggestive of [[binocular vision]], as well as a long lifespan, which developed in an unusual ecosystem where only a few other mammal species were present, terrestrial predators were absent, and ''Myotragus'' functioned as the only major herbivore. ''M.'' ''balearicus'' became extinct when humans arrived in the Balearic Islands during the [[3rd millennium BC]], along with the large shrew ''[[Nesiotites]]'' and the giant dormouse ''[[Hypnomys]]'', the only other terrestrial mammals native to the islands.

Early genetic research suggested that it was closely related to sheep of the genus ''[[Ovis]]'';<ref name="Lalueza2005">{{cite journal |author=Lalueza-Fox |display-authors=etal |date=6 December 2005 |title=Molecular dating of caprines using ancient DNA sequences of Myotragus balearicus, an extinct endemic Balearic mammal |journal=BMC Evolutionary Biology |volume=5 |issue=1 |pages=70 |doi=10.1186/1471-2148-5-70 |pmc=1325260 |pmid=16332256 |doi-access=free |bibcode=2005BMCEE...5...70L }}</ref> however, more recent research has indicated that its closest living relative is the [[takin]] (''Budorcas taxicolor'').<ref name=":0">{{Cite journal |last1=Bover |first1=Pere |last2=Llamas |first2=Bastien |last3=Mitchell |first3=Kieren J. |last4=Thomson |first4=Vicki A. |last5=Alcover |first5=Josep Antoni |last6=Lalueza-Fox |first6=Carles |last7=Cooper |first7=Alan |last8=Pons |first8=Joan |date=July 2019 |title=Unraveling the phylogenetic relationships of the extinct bovid Myotragus balearicus Bate 1909 from the Balearic Islands |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379118306929 |journal=Quaternary Science Reviews |language=en |volume=215 |pages=185–195 |bibcode=2019QSRv..215..185B |doi=10.1016/j.quascirev.2019.05.005 |s2cid=189965070}}</ref>

== History of discovery and taxonomy ==
The first remains of ''Myotragus'' were described by [[Dorothea Bate]] in 1909. Bate had been sent a letter by [[Robert Ashington Bullen]], who informed her about a bone-bearing [[breccia]] deposit on the east of [[Mallorca]], which prompted her to survey the island for [[Pleistocene]] aged cave deposits. Three such deposits were found, which yielded fragmentary remains of ''Myotragus'', including a mostly complete skull, associated with a mandible and [[Atlas (anatomy)|atlas vertebra]], which was designated the [[type specimen]] of the new species and genus ''Myotragus balearicus''.<ref>{{Cite journal|last=Bate|first=Dorothea M. A.|date=September 1909|title=I.—Preliminary Note on a New Artiodactyle from Majorca, Myotragus balearicus , gen. et sp. nov.|url=https://www.cambridge.org/core/product/identifier/S0016756800124665/type/journal_article|journal=Geological Magazine|language=en|volume=6|issue=9|pages=385–388|doi=10.1017/S0016756800124665|bibcode=1909GeoM....6..385B|s2cid=128708600 |issn=0016-7568}}</ref> In 1915, [[Charles William Andrews]] described more material discovered in the intervening years, including material that had been discovered on [[Menorca]].<ref>{{Cite journal|date=January 1915|title=VI. A description of the skull and skeleton of a peculiarly modified rupicaprine antelope (Myotragus balearicus , Bate), with a notice of a new variety, M. balearicus var. major|journal=Philosophical Transactions of the Royal Society of London. Series B, Containing Papers of a Biological Character|language=en|volume=206|issue=325–334|pages=281–305|doi=10.1098/rstb.1915.0006|bibcode=1915RSPTB.206..281A|issn=0264-3960|doi-access=free|last1=Andrews|first1=Chas. W.}}</ref>

=== Species ===
Six sequential [[chronospecies]] of ''Myotragus'' have been named,<ref name=":1">{{Cite journal |last1=Bover |first1=Pere |last2=Quintana |first2=Josep |last3=Alcover |first3=Josep Antoni |date=November 2010 |title=A new species of Myotragus Bate, 1909 (Artiodactyla, Caprinae) from the Early Pliocene of Mallorca (Balearic Islands, western Mediterranean) |url=https://www.cambridge.org/core/product/identifier/S0016756810000336/type/journal_article |journal=Geological Magazine |language=en |volume=147 |issue=6 |pages=871–885 |bibcode=2010GeoM..147..871B |doi=10.1017/S0016756810000336 |issn=0016-7568 |s2cid=35830707 |hdl-access=free |hdl=10261/49361}}</ref> representing 5 million years of gradual accumulated morphological change, including a reduction in body size and changes to the locomotor system, the teeth and the visual system.<ref name=":2">{{cite journal |author1=Köhler M. |author2=Moyà-Solà S. |year=2004 |title=Reduction of Brain and Sense Organs in the Fossil Insular Bovid Myotragus |journal=Brain, Behavior and Evolution |volume=63 |issue=3 |pages=125–140 |doi=10.1159/000076239 |pmid=14726622 |s2cid=42399219}}</ref>
*'''''M. palomboi''''' <small>Bover, Quintana & Alcover, 2010</small> Early Pliocene, Mallorca
*'''''M. pepgonellae''''' <small>Moyà-Solà & Pons-Moyà, 1982</small> Middle Pliocene, Mallorca
*'''''M. antiquus''''' <small>Pons-Moyà, 1977</small> Late Pliocene, Mallorca
*'''''M. kopperi''''' <small>Moya & Pons, 1980</small> Early Pleistocene, Mallorca
*'''''M. batei''''' <small>Crusafont & Angel, 1966</small> Middle Pleistocene, Mallorca, Menorca
*'''''M. balearicus''''' <small>Bate, 1909</small> Late Pleistocene-Holocene, Mallorca, Menorca
The species ''M. binigausensis'' <small>Moyà-Solà and Pons-Moyà</small>, <small>1980</small> from the Pleistocene of Menorca has been synonymised with ''M. batei.''<ref>{{Cite journal |last1=Palombo |first1=Maria Rita |last2=Rozzi |first2=Roberto |last3=Bover |first3=Pere |date=January 2013 |title=The endemic bovids from Sardinia and the Balearic Islands: State of the art |url=https://linkinghub.elsevier.com/retrieve/pii/S0016699513000120 |journal=Geobios |language=en |volume=46 |issue=1–2 |pages=127–142 |doi=10.1016/j.geobios.2012.10.011|bibcode=2013Geobi..46..127P }}</ref>

==Evolution ==
The closest fossil relatives of ''Myotragus'' are uncertain. A close relationship has been proposed to the genera ''[[Aragoral]]'' and ''[[Norbertia]]'' from the Late [[Miocene]] of mainland Europe,<ref name=":1" /> as well as the insular genera ''[[Ebusia]]'' from the Pliocene of Ibiza, and ''[[Nesogoral]]'' from the Early Pleistocene of Sardinia.<ref name=":7">{{Cite journal |last1=Moyà-Solà |first1=Salvador |last2=Quintana Cardona |first2=Josep |last3=Köhler |first3=Meike |date=2022-08-03 |title=Ebusia moralesi n. gen. nov. sp, a new endemic caprine (Bovidae, Mammalia) from the Neogene of Eivissa Island (Balearic Islands, Western Mediterranean): evolutionary implications |journal=Historical Biology |language=en |volume=34 |issue=8 |pages=1642–1659 |doi=10.1080/08912963.2022.2060099 |issn=0891-2963|doi-access=free |bibcode=2022HBio...34.1642M }}</ref> A 2005 study of a partial mitochondrial genome suggested that ''Myotragus'' was the [[sister group]] to the genus ''[[Ovis]]'', which includes sheep.<ref name="Lalueza2005" /> However, analysis of a complete mitochondrial genome of ''M. balearicus'' published in 2019 found that its closest living relative is the [[takin]] (''Budorcas taxicolor''), native to the eastern Himalayas with an estimated [[Speciation|divergence]] around 7.1 million years ago. A [[cladogram]] showing its position within Caprinae/Caprini is given below.<ref name=":0" />

{{clade|{{clade
   |1=''[[Bos]]''
   |label2=Caprini
   |2={{clade
      |1=''[[Pantholops]]'' (Tibetan antelope)
      |2={{clade
         |1={{clade
            |1={{clade
               |1=†''[[Bootherium]]'' (Helmeted muskox)
               |2=''[[Ovibos]]'' (Musk ox)
               }}
            |2={{clade
               |1=''[[Capricornis]]'' (Serow)
               |2=''[[Naemorhedus]]'' (Goral)
               }}
            }}
         |2={{clade
            |1={{clade
               |1=''[[Ovis]]'' (Sheep)
               }}
            |2={{clade
               |1={{clade
                  |1={{clade
                     |1={{clade
                        |1=''[[Oreamnos]]'' (Mountain goat)
                        }}
                     |2={{clade
                        |1={{clade
                           |1=''[[Budorcas]]'' (Takin)
                           }}
                        |2={{clade
                           |1={{clade
                              |1=†'''''Myotragus'''''
                              }}
                           }}
                        }}
                     }}
                  |2={{clade
                     |1={{clade
                        |1=''[[Rupicapra]]'' (Chamois)
                        }}
                     }}
                  }}
               |2={{clade 
                  |1={{clade
                     |1=''[[Ammotragus]]'' (Barbary sheep)
                     |2=''[[Arabitragus]]'' (Arabian tahr)
                     }}
                  |2={{clade
                     |1={{clade
                        |1=''[[Pseudois]]'' (Bharal)
                        }}
                     |2={{clade
                        |1=''[[Hemitragus]]'' (Himalayan tahr)
                        |2=''[[Capra (genus)|Capra]]'' (Markhor, ibexes, goats)
                        }}
                     }}
                  }}
               }}
            }}
         }}
      }}
   }}}}

The ancestor of ''Myotragus'' likely arrived in the Balearic Islands during the [[Messinian]] stage of the late Miocene at a time at which the [[Strait of Gibraltar]] closed and the Mediterranean Sea evaporated, reducing sea level within the basin by 800–1200 metres, in an event called the [[Messinian salinity crisis]] (spanning from 5.96 to 5.33 million years ago), allowing dispersal from the Iberian Peninsula to the Balearics.<ref>{{Cite journal |last1=Mas |first1=Guillem |last2=Maillard |first2=Agnès |last3=Alcover |first3=Josep A. |last4=Fornós |first4=Joan J. |last5=Bover |first5=Pere |last6=Torres-Roig |first6=Enric |date=2018-06-01 |title=Terrestrial colonization of the Balearic Islands: New evidence for the Mediterranean sea-level drawdown during the Messinian Salinity Crisis |url=https://pubs.geoscienceworld.org/gsa/geology/article/46/6/527/530693/Terrestrial-colonization-of-the-Balearic-Islands |journal=Geology |language=en |volume=46 |issue=6 |pages=527–530 |bibcode=2018Geo....46..527M |doi=10.1130/G40260.1 |issn=0091-7613}}</ref>

Later on, the re-opening of the straits and [[Zanclean flood|the refilling of the Mediterranean]] at the beginning of the [[Pliocene]] around 5.3 million years ago isolated the animal populations on the islands. The changes in morphology ''Myotragus'' developed over the course of its evolution were probably driven by resource limitation on the relatively resource poor Balearics, with the lack of competitors leading to increased [[intraspecific competition]], and the absence of effective predators meaning the population would periodically outstrip the [[carrying capacity]] of the islands, resulting in the denudation of most vegetation and consequently mass starvation,<ref name=":2" /><ref name=":4" /><ref name=":7" /> with only a small proportion of the population surviving a starvation episode, leading to strong [[selection pressure]].<ref name=":7" />

''Myotragus'' initially only colonized the island of Mallorca. Only a handful of mammal species aside from ''Myotragus'' were able to colonise the island, including shrews, [[hamsters]], [[dormice]], [[Murinae|murines]] and rabbits. By the Late Pliocene, ''Myotragus'' represented one of only three genera of mammal present on Mallorca, alongside the giant dormouse ''[[Hypnomys]]'' and the shrew ''[[Nesiotites]],'' all of which would continue to be present on the island until the [[Holocene]].<ref>{{Cite journal |last=Palombo |first=Maria Rita |date=January 2018 |title=Insular mammalian fauna dynamics and paleogeography: A lesson from the Western Mediterranean islands |journal=Integrative Zoology |language=en |volume=13 |issue=1 |pages=2–20 |doi=10.1111/1749-4877.12275 |issn=1749-4877 |pmc=5817236 |pmid=28688123}}</ref> On Menorca, a giant rabbit, ''[[Nuralagus|Nuralagus rex]]'' evolved that covered the same niche as ''Myotragus'' in Mallorca.<ref name="Quintana">{{cite journal |last1=Quintana |first1=Josep |last2=Kohler |first2=Mike |last3=Moya-Sola |first3=Salvador |year=2011 |title=''Nuralagus rex'', gen. et sp. nov., an endemic insular giant rabbit from the Neogene of Menorca |journal=Journal of Vertebrate Paleontology |volume=31 |issue=2 |pages=231–240 |doi=10.1080/02724634.2011.550367 |s2cid=83649921}}</ref> With the level of the sea falling due to [[Ice age|glacial cycles]] during the [[Pleistocene]], Mallorca and Menorca were periodically connected and the mammals of Mallorca, including ''Myotragus'' colonised Menorca, replacing the great Menorcan lagomorphs.<ref>{{cite journal |author1=Pere Bover |author2=Josep Quintanab |author3=Josep Antoni Alcover |year=2008 |title=Three islands, three worlds: Paleogeography and evolution of the vertebrate fauna from the Balearic Islands |journal=Quaternary International |volume=182 |issue=1 |pages=135–144 |bibcode=2008QuInt.182..135B |doi=10.1016/j.quaint.2007.06.039 |hdl-access=free |hdl=10261/85878}}</ref> Both islands separated again at the beginning of the [[Holocene]].

==Description==
[[File:Höhlenziege Museum Manacor 12.jpg|left|thumb|Skull in dorsal view]]
The size of ''Myotragus'' varied between species, generally reducing with time. The early species ''M. pepgonellae'' is estimated to have had a body mass of approximately {{Convert|60|kg|lb}}, while the later ''M. kopperi'' is estimated to have been approximately {{Convert|23|kg|lb}},<ref name=":2" /> representing an example of [[insular dwarfism]].<ref name=":4" /> ''M. balearicus'' is estimated to have been approximately {{Convert|50|cm|ft}} tall at the shoulder,<ref name=":3" /> with a 2004 study estimating an adult body mass of around {{Convert|23–32|kg|lb|abbr=on}}.<ref name=":2" /> The orbits of the skull of ''M. balearicus'' are roughly half the size those of other comparably sized caprines, and face-forward as opposed to the sides as in most ungulates and in earlier ''Myotragus'' species, allowing for [[binocular vision]].<ref name=":2" /> The skull had a pair of small posteriorly-directed horns.<ref name=":3" /> The species of ''Myotragus'' show a sequential reduction in the number of teeth through time, with ''M. balearicus'' having an adult dentition comprising a single evergrowing (hypselodont) [[incisor]] (a feature highly unusual among [[bovids]]), one [[premolar]], and three [[molars]] in each half of the lower jaw, and two premolars and three molars in each half of the upper jaw.<ref>{{Cite journal |last1=Bover |first1=Pere |last2=Alcover |first2=Josep Antoni |date=November 1999 |title=The evolution and ontogeny of the dentition of Myotragus balearicus Bate, 1909 (Artiodactyla, Caprinae): evidence from new fossil data |journal=Biological Journal of the Linnean Society |language=en |volume=68 |issue=3 |pages=401–428 |doi=10.1111/j.1095-8312.1999.tb01178.x|doi-access=free }}</ref><ref name=":1" /><ref name=":6">{{Cite journal |last1=Winkler |first1=Daniela Eileen |last2=Schulz |first2=Ellen |last3=Calandra |first3=Ivan |last4=Gailer |first4=Juan-Pablo |last5=Landwehr |first5=Christina |last6=Kaiser |first6=Thomas M. |date=January 2013 |title=Indications for a dietary change in the extinct Bovid genus Myotragus (Plio-Holocene, Mallorca, Spain) |url=https://linkinghub.elsevier.com/retrieve/pii/S0016699513000119 |journal=Geobios |language=en |volume=46 |issue=1–2 |pages=143–150 |doi=10.1016/j.geobios.2012.10.010|bibcode=2013Geobi..46..143W }}</ref><ref>{{Cite journal |last1=Jordana |first1=Xavier |last2=Marín-Moratalla |first2=Nekane |last3=Moncunill-Solé |first3=Blanca |last4=Bover |first4=Pere |last5=Alcover |first5=Josep Antoni |last6=Köhler |first6=Meike |date=2013-07-25 |editor-last=Laudet |editor-first=Vincent |title=First Fossil Evidence for the Advance of Replacement Teeth Coupled with Life History Evolution along an Anagenetic Mammalian Lineage |journal=PLOS ONE |language=en |volume=8 |issue=7 |pages=e70743 |doi=10.1371/journal.pone.0070743 |issn=1932-6203 |pmc=3723812 |pmid=23936247 |doi-access=free |bibcode=2013PLoSO...870743J }}</ref> By contrast, the earliest species had three incisors, one [[Canine tooth|canine]], and two premolars in each half of the lower jaw, as is typical of most [[Ruminant|ruminants]]<ref name=":3" /> with the incisors not being ever-growing, and a third premolar present in each half of the upper jaw.<ref name=":1" /> The teeth in the later species are much more [[hypsodont]] (high crowned) than in earlier species.<ref name=":3" /> The limbs of ''M. balearicus'' are relatively short in comparison to other caprines, with the [[Tarsal bone|tarsals]], [[Metatarsal bones|metatarsals]] and [[sesamoids]] being partially fused.<ref name=":5" />

== Paleobiology ==
[[Image:Myotragus balearicus.JPG|thumb|Restoration]]

=== Diet ===
While tooth morphology and tooth texture suggests that some earlier ''Myotragus'' species may have been [[Grazing (behaviour)|grazers]] or mixed feeders (consuming both grass and browse),<ref name=":6" /> preserved [[coprolite]]s of ''M. balearicus'' indicates that it was likely predominantly a [[Browsing (herbivory)|browser]], and heavily dependent on the native [[Buxus|boxwood]] species ''[[Buxus balearica]]'' for a large part of its diet.<ref>{{cite journal |author=Alcover|title=The diet of Myotragus balearicus Bate 1909 (Artiodactyla: Caprinae), an extinct bovid from the Balearic Islands: evidence from coprolites |journal=Biological Journal of the Linnean Society |year=1999 |volume=66 |issue=1 |pages=57–74 |doi=10.1111/j.1095-8312.1999.tb01917.x |display-authors=etal|doi-access=free }}</ref><ref>{{Cite journal|last1=Welker|first1=Frido|last2=Duijm|first2=Elza|last3=van der Gaag|first3=Kristiaan J.|last4=van Geel|first4=Bas|last5=de Knijff|first5=Peter|last6=van Leeuwen|first6=Jacqueline|last7=Mol|first7=Dick|last8=van der Plicht|first8=Johannes|last9=Raes|first9=Niels|last10=Reumer|first10=Jelle|last11=Gravendeel|first11=Barbara|date=January 2014|title=Analysis of coprolites from the extinct mountain goat Myotragus balearicus|url=https://www.cambridge.org/core/product/identifier/S0033589400002106/type/journal_article|journal=Quaternary Research|language=en|volume=81|issue=1|pages=106–116|doi=10.1016/j.yqres.2013.10.006|bibcode=2014QuRes..81..106W|s2cid=84266057|issn=0033-5894}}</ref> The increased hypsodonty over time of the teeth of ''Myotragus'' likely represents at least in part an adaptation to the increased consumption of abrasive food.<ref name=":6" />

=== Physiology and growth ===
The bone histology of ''M. balearicus'' shows [[Lamellar bone|lamellar-zonal tissue]] throughout the cortex, with lines of arrested growth indicating periods where growing ceased. Although this bone morphology has previously been asserted to be otherwise unique to reptiles and a sign that ''Myotragus'' was [[ectothermic]] like reptiles,<ref>{{cite journal |last1=Kohler |first1=Meike |last2=Moya-Sola |first2=Salvador |year=2009 |title=Physiological and life history strategies of a fossil large mammal in a resource-limited environment |journal=Proceedings of the National Academy of Sciences |volume=106 |issue=48 |pages=20354–8 |bibcode=2009PNAS..10620354K |doi=10.1073/pnas.0813385106 |jstor=25593378 |pmc=2777955 |pmid=19918076 |doi-access=free}}</ref> later research suggested that this bone morphology is common to all ruminants and is not unique to ''Myotragus''.<ref>{{Cite journal |last1=van der Geer |first1=Alexandra A. E. |last2=Lyras |first2=George A. |last3=MacPhee |first3=Ross D. E. |last4=Lomolino |first4=Mark |last5=Drinia |first5=Hara |date=2014-06-24 |title=Mortality in a Predator-free Insular Environment: the Dwarf Deer of Crete |url=http://www.bioone.org/doi/abs/10.1206/3807.1 |journal=American Museum Novitates |language=en |issue=3807 |pages=1–26 |doi=10.1206/3807.1 |hdl=10795/3278 |issn=0003-0082|hdl-access=free }}</ref><ref>{{Cite journal |last1=Köhler |first1=Meike |last2=Marín-Moratalla |first2=Nekane |last3=Jordana |first3=Xavier |last4=Aanes |first4=Ronny |date=2012-07-19 |title=Seasonal bone growth and physiology in endotherms shed light on dinosaur physiology |url=https://www.nature.com/articles/nature11264 |journal=Nature |language=en |volume=487 |issue=7407 |pages=358–361 |doi=10.1038/nature11264 |pmid=22763443 |bibcode=2012Natur.487..358K |issn=0028-0836}}</ref> Based on counting the lines of arrested growth, it has been estimated that ''M. balearicus'' reached maximum size (somatic maturity) and probably sexual maturity at 12 years of age.<ref>{{cite journal |last1=Kohler |first1=Meike |last2=Moya-Sola |first2=Salvador |year=2009 |title=Physiological and life history strategies of a fossil large mammal in a resource-limited environment |journal=Proceedings of the National Academy of Sciences |volume=106 |issue=48 |pages=20354–8 |bibcode=2009PNAS..10620354K |doi=10.1073/pnas.0813385106 |jstor=25593378 |pmc=2777955 |pmid=19918076 |doi-access=free}}</ref> Analysis of the high-crowned teeth of ''M. balearicus,'' shows that they grew more slowly than those of other caprines, with their last teeth erupting at approximately six years of age, likely as an adaption to their longevity,<ref name=":4">{{Cite journal |last1=Jordana |first1=Xavier |last2=Köhler |first2=Meike |date=February 2011 |title=Enamel microstructure in the fossil bovid Myotragus balearicus (Majorca, Spain): Implications for life-history evolution of dwarf mammals in insular ecosystems |url=https://linkinghub.elsevier.com/retrieve/pii/S0031018210007248 |journal=Palaeogeography, Palaeoclimatology, Palaeoecology |language=en |volume=300 |issue=1–4 |pages=59–66 |doi=10.1016/j.palaeo.2010.12.008|bibcode=2011PPP...300...59J }}</ref> though the rodent-like evergrowing lower incisor of ''M. balearicus'' erupted early, both relative to the posterior teeth and in absolute age compared to other bovids.<ref>{{Cite journal |last1=Jordana |first1=Xavier |last2=Marín-Moratalla |first2=Nekane |last3=Moncunill-Solé |first3=Blanca |last4=Bover |first4=Pere |last5=Alcover |first5=Josep Antoni |last6=Köhler |first6=Meike |date=2013-07-25 |editor-last=Laudet |editor-first=Vincent |title=First Fossil Evidence for the Advance of Replacement Teeth Coupled with Life History Evolution along an Anagenetic Mammalian Lineage |journal=PLOS ONE |language=en |volume=8 |issue=7 |pages=e70743 |doi=10.1371/journal.pone.0070743 |doi-access=free |issn=1932-6203 |pmc=3723812 |pmid=23936247|bibcode=2013PLoSO...870743J }}</ref> Based on skeletochronology and dental durability analysis, some individuals of ''M. balearicus'' are likely to have reached a lifespan of 27 years, which is exceptionally long relative to its body size. The estimated mortality rates are substantially lower than those found for other members of Bovidae, with a large proportion of individuals surviving into old age.<ref>{{Cite journal |last1=Jordana |first1=Xavier |last2=Marín-Moratalla |first2=Nekane |last3=DeMiguel |first3=Daniel |last4=Kaiser |first4=Thomas M. |last5=Köhler |first5=Meike |date=2012-08-22 |title=Evidence of correlated evolution of hypsodonty and exceptional longevity in endemic insular mammals |journal=Proceedings of the Royal Society B: Biological Sciences |language=en |volume=279 |issue=1741 |pages=3339–3346 |doi=10.1098/rspb.2012.0689 |issn=0962-8452 |pmc=3385741 |pmid=22535784}}</ref>

Newborn specimens of ''M. balearicus'' are estimated to have been approximately {{Convert|15–18|cm|in}} in height with a weight of about {{Convert|700-900|g|lb}}, approximately 2% the bodymass of a mature adult, much lower than that of a typical ruminant, in which newborns are usually over 4% the adult bodymass.<ref>P. Bover & J.A. Alcover. (1999). [https://natuurtijdschriften.nl/pub/538619 Estimating physical characteristics of neonate Myotragus balearicus (Artiodactyla, Caprinae).] ''Deinsea'', ''7''(1), 33–54.</ref>

=== Movement and senses ===
An analysis of the [[Phalanx bone|phalanges]] (digits) of ''M. balearicus'' found that the bones of the foot were tightly bound by [[ligament]]s and inelastic. This suggests that it was obligately a slow walker with a reduced step length, and lacked the ability to jump. The likely reason for this is as an energy saving measure, as the shock absorbing mechanism in the foot bones of other caprines requires large amounts of muscle energy, with the ability to move quickly unnecessary on the largely predator-free islands. The proximal (closest to the ankle) and medial (intermediate between the furthest and closest) phalanges were likely orientated vertically relative to the ground surface, which reduced bending stresses.<ref name=":5">{{Cite journal|last1=Köhler|first1=Meike|last2=Moyà-Solà|first2=Salvador|date=2001-08-22|title=Phalangeal adaptations in the fossil insular goat Myotragus|url=http://www.tandfonline.com/doi/abs/10.1671/0272-4634%282001%29021%5B0621%3APAITFI%5D2.0.CO%3B2|journal=Journal of Vertebrate Paleontology|language=en|volume=21|issue=3|pages=621–624|doi=10.1671/0272-4634(2001)021[0621:PAITFI]2.0.CO;2|s2cid=85654563 |issn=0272-4634}}</ref> 

The [[cranial endocast]] of ''M. balearicus'' indicates that the areas of the brain and structures associated with vision, sound and smell are small when compared with living caprines, with the brain only being half the size of other comparably sized modern caprines,<ref name=":2" /> though a 2024 study found when compared to its likely late Miocene ancestors, the size reduction is a more modest 10-17%, due to the much smaller size of the brains of late Miocene bovids than contemporary bovid species, with the majority of the brain reduction in ''M. balearicus'' compared to late Miocene taxa being from the vision-associated [[occipital lobe]] including the [[visual cortex]].<ref name=":8">{{Cite journal |last=Liakopoulou |first=Dionysia |last2=Roussiakis |first2=Socrates |last3=Lyras |first3=George |date=2024-10-11 |title=The brain of Myotragus balearicus , an insular bovid from the Balearics |url=https://www.tandfonline.com/doi/full/10.1080/08912963.2024.2406962 |journal=Historical Biology |language=en |pages=1–8 |doi=10.1080/08912963.2024.2406962 |issn=0891-2963|doi-access=free }}</ref> The smaller brain likely represents  an optimisation  to the animals energy budget, as neural tissue is energetically expensive to maintain, and better developed senses were  unnecessary in a resource limited environment where there was little need to detect predators. The binocular vision of ''M. balearicus'' likely enhanced [[depth perception]] at the expense of the [[field of view]]. The outwards-facing eyes giving a large field of view typical of ungulates is thought to be an adaptation for vigilance against predators.<ref name=":2" /> ''M. balearicus'' likely relied heavily on tactile senses rather than sight when selecting food.<ref name=":8" />

==Extinction==
Diverse datings indicate that the three native terrestrial mammals of Mallorca and Menorca (''Myotragus balearicus'', the giant dormouse ''[[Hypnomys]]'' and the large shrew ''[[Nesiotites hidalgo]]'') disappeared all in the same very short period of time, during the third millennium BC.<ref>{{cite journal |author1=Pere Bover |author2=Josep Antoni Alcover |title=Understanding Late Quaternary extinctions: the case of Myotragus balearicus (Bate, 1909) |journal=Journal of Biogeography |volume=30 |issue=5 |pages=771–781 |year=2003 |doi=10.1046/j.1365-2699.2003.00872.x |bibcode=2003JBiog..30..771B |hdl=10261/85881 |s2cid=84386172 }}</ref> Historically there was debate as to whether the extinctions were caused by climate change, or whether they were exterminated by the first human settlers of the Balearic Islands.

The dominant theory is the one that postulates an extinction by human causes. Traditional methods had dated the first human colonization of the Balearic Islands towards 5000 BC or even before, but subsequent tests with modern methods of dating clearly indicate that there was no human presence before 3000 BC. This date agrees very closely with the fast decline of the three forms. The youngest remains of ''Myotragus'' date to around 2632 calibrated years BC, while the minimum date of human arrival on the Balearic Islands is currently 2282 BC. Extinction was likely rapid within less than 100 years of human arrival on the islands.<ref>{{Cite journal|last1=Bover|first1=Pere|last2=Valenzuela|first2=Alejandro|last3=Torres|first3=Enric|last4=Cooper|first4=Alan|last5=Pons|first5=Joan|last6=Alcover|first6=Josep A|date=November 2016|title=Closing the gap: New data on the last documented Myotragus and the first human evidence on Mallorca (Balearic Islands, Western Mediterranean Sea)|url=http://journals.sagepub.com/doi/10.1177/0959683616645945|journal=The Holocene|language=en|volume=26|issue=11|pages=1887–1891|doi=10.1177/0959683616645945|bibcode=2016Holoc..26.1887B |s2cid=131641995 |issn=0959-6836}}</ref><ref>{{Cite journal|last1=Valenzuela|first1=Alejandro|last2=Torres-Roig|first2=Enric|last3=Zoboli|first3=Daniel|last4=Pillola|first4=Gian Luigi|last5=Alcover|first5=Josep Antoni|date=2021-11-29|title=Asynchronous ecological upheavals on the Western Mediterranean islands: New insights on the extinction of their autochthonous small mammals|url=http://journals.sagepub.com/doi/10.1177/09596836211060491|journal=The Holocene|volume=32 |issue=3 |language=en|pages=137–146|doi=10.1177/09596836211060491|s2cid=244763779 |issn=0959-6836}}</ref>

In 1969 it was suggested that ''Myotragus'' shows signs of [[domestication]]<ref name=":3">{{cite book |author1=Alexandra Van Der Geer |title=Evolution of Island Mammals: Adaptation and Extinction of Placental Mammals on Islands |author2=George Lyras |author3=John De Vos |publisher=Wiley-Blackwell |year=2010 |isbn=978-1405190091 |pages=132–146}}</ref> but a 2001 study found that there was no empirical evidence supporting the idea, with marks on ''Myotragus'' horns that were suggested to be made by humans were probably actually the result of gnawing on bones by other ''Myotragus'', likely for their mineral content.<ref>{{cite journal|author1=Damià Ramis|author2=Pere Bover|year=2001|title=A Review of the Evidence for Domestication of Myotragus balearicus Bate 1909 (Artiodactyla, Caprinae) in the Balearic Islands|journal=Journal of Archaeological Science|volume=28|issue=3|pages=265–282|doi=10.1006/jasc.2000.0548|bibcode=2001JArSc..28..265R |hdl=10261/54367}}</ref>

==See also==
*[[List of extinct animals of Europe]]
*[[Schultz's rule]]

==References==
{{Reflist}}
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{{Taxonbar|from1=Q3011985|from2=Q1643902}}

{{DEFAULTSORT:Myotragus Balearicus}}
[[Category:Prehistoric caprids]]
[[Category:Prehistoric mammals of Europe]]
[[Category:Endemic fauna of the Balearic Islands]]
[[Category:Holocene extinctions]]
[[Category:Fossil taxa described in 1909]]
[[Category:Prehistoric Artiodactyla genera]]