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=== Ecological aspects of historical distribution === [[File:Maclura pomifera in Kansas winter.jpg|thumb|Evidence of a seed predator (February in Kansas).]][[File:Maclura pomifera fruit sprouting seeds.jpg|thumb|Mound of a single fallen fruit sprouting seeds (April in Illinois).]] Because of the limited original range and lack of obvious effective means of propagation, the Osage orange has been the subject of controversial claims by some authors to be an [[evolutionary anachronism]], whereby one or more now extinct [[Pleistocene megafauna]], such as [[Ground sloth|ground sloths]], [[mammoth|mammoths]], [[mastodon|mastodons]] or [[gomphothere|gomphotheres]], fed on the fruit and aided in seed dispersal.<ref name=":0">Connie Barlow. [http://www.thegreatstory.org/anachronistic_fruits/index.html Anachronistic Fruits and the Ghosts Who Haunt Them] {{Webarchive|url=https://web.archive.org/web/20070106121158/http://www.thegreatstory.org/anachronistic_fruits/index.html |date=2007-01-06 }}. ''[[Arnoldia]]'', vol. 61, no. 2 (2001)</ref><ref>{{cite journal|url=http://www.americanforests.org/magazine/article/trees-that-miss-the-mammoths/|title=The Trees That Miss The Mammoths|first= Whit |last=Bronaugh|year=2010|journal= American Forests|volume=115|issue=Winter|pages=38β43}}</ref> An [[equine]] species that became extinct at the same time also has been suggested as the plant's original dispersal agent because modern horses and other livestock will sometimes eat the fruit.<ref name=Ghosts_Evo/> This hypothesis is controversial. For example, a 2015 study indicated that Osage orange seeds are not effectively spread by extant horse or elephant species,<ref name="Anachronistic_fruits">{{cite journal|last1=Boone|first1=Madison J.|last2=Davis|first2=Charli N.|last3=Klasek|first3=Laura|last4=del Sol |first4=Jillian F. |last5=Roehm|first5=Katherine|last6=Moran|first6=Matthew D.|title=A Test of Potential Pleistocene Mammal Seed Dispersal in Anachronistic Fruits using Extant Ecological and Physiological Analogs|journal=Southeastern Naturalist|date=11 March 2015 |volume=14 |issue=1|pages=22β32|doi=10.1656/058.014.0109|s2cid=86809830}}</ref> while a 2018 study concludes that squirrels are ineffective, short-distance seed dispersers.<ref name="murphy" /> The claim has been criticised as a "[[just-so story]]" that lacks any empirical evidence.<ref name=":1" /> The fruit is not poisonous to humans or livestock, but is not preferred by them,<ref name=IPM_IowaU_2>{{cite web |url=http://www.ipm.iastate.edu/ipm/hortnews/1997/10-10-1997/hedgeapple.html |title=Facts and Myths Associated with "Hedge Apples" |date=October 10, 1997 |last=Jauron |first=Richard |work=Horticulture and Home Pest News |publisher=Iowa State University |access-date=October 22, 2014}}</ref> because it is mostly inedible due to a large size (about the diameter of a [[Softball (ball)|softball]]) and hard, dry texture.<ref name=Ghosts_Evo/> The edible seeds of the fruit are used by [[squirrel]]s as food.<ref>Murphy, Serena, Virginia Mitchell, Jessa Thurman, Charli N. Davis, Mattew D. Moran, Jessica Bonumwezi, Sophie Katz, Jennifer L. Penner, and Matthew D. Moran. "Seed Dispersal in Osage Orange (Maclura pomifera) by Squirrels (Sciurus spp.)." The American Midland Naturalist 180, no. 2 (2018): 312-317. Harvard</ref> Large animals such as [[livestock]], which typically would consume fruits and disperse seeds, mainly ignore the fruit.<ref name=Ghosts_Evo/>
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