Episodic memory and mental time travel have been viewed as uniquely human traits.1Tulving E. Elements of Episodic Memory. Oxford University Press, 1983Google Scholar,2Suddendorf T. Corballis M.C. Mental time travel and the evolution of the human mind.Genet. Psychol. Monogr. 1997; 123: 133-167Google Scholar,3Raby C.R. Alexis D.M. Dickinson A. Clayton N.S. Planning for the future by western scrub-jays.Nature. 2007; 445: 919-921Crossref PubMed Scopus (433) Google Scholar This view began to shift with the development of behavioral criteria to assess what is referred to as "episodic-like memory" in animals.4Clayton N.S. Dickinson A. Episodic-like memory during cache recovery by scrub jays.Nature. 1998; 395: 272-274Crossref PubMed Scopus (1064) Google Scholar,5Clayton N.S. Bussey T.J. Dickinson A. Can animals recall the past and plan for the future?.Nat. Rev. Neurosci. 2003; 4: 685-691Crossref PubMed Scopus (508) Google Scholar Key findings have ranged from evidence of what-where-when memory in scrub-jays, rats, and bees; through decision-making that impacts future foraging in frugivorous primates; to evidence of planning based on future needs in scrub-jays and tool use planning in great apes.4Clayton N.S. Dickinson A. Episodic-like memory during cache recovery by scrub jays.Nature. 1998; 395: 272-274Crossref PubMed Scopus (1064) Google Scholar,6Roberts W.A. Feeney M.C. MacPherson K. Petter M. McMillan N. Musolino E. Episodic-like memory in rats: is it based on when or how long ago?.Science. 2008; 320: 113-115Crossref PubMed Scopus (124) Google Scholar,7Pahl M. Zhu H. Pix W. Tautz J. Zhang S. Circadian timed episodic-like memory–a bee knows what to do when, and also where.J. Exp. Biol. 2007; 210: 3559-3567Crossref PubMed Scopus (0) Google Scholar,8Noser R. Byrne R.W. Travel routes and planning of visits to out-of-sight resources in wild chacma baboons, Papio ursinus.Anim. Behav. 2007; 73: 257-266Crossref Scopus (89) Google Scholar,9Ban S.D. Boesch C. Janmaat K.R.L. Taï chimpanzees anticipate revisiting high-valued fruit trees from further distances.Anim. Cogn. 2014; 17: 1353-1364Crossref PubMed Scopus (24) Google Scholar,10Janmaat K.R.L. Polansky L. Ban S.D. Boesch C. Wild chimpanzees plan their breakfast time, type, and location.Proc. Natl. Acad. Sci. USA. 2014; 111: 16343-16348Crossref PubMed Scopus (113) Google Scholar,11Correia S.P.C. Dickinson A. Clayton N.S. Western scrub-jays anticipate future needs independently of their current motivational state.Curr. Biol. 2007; 17: 856-861Abstract Full Text Full Text PDF PubMed Scopus (193) Google Scholar,12Dufour V. Pelé M. Sterck E.H.M. Thierry B. Chimpanzee (Pan troglodytes) anticipation of food return: coping with waiting time in an exchange task.J. Comp. Psychol. 2007; 121: 145-155Crossref PubMed Scopus (0) Google Scholar,13Mulcahy N.J. Call J. Apes save tools for future use.Science. 2006; 312: 1038-1040Crossref PubMed Scopus (381) Google Scholar Field studies of these issues have been rare, though there is field-based evidence for future-oriented behaviors in primates.8Noser R. Byrne R.W. Travel routes and planning of visits to out-of-sight resources in wild chacma baboons, Papio ursinus.Anim. Behav. 2007; 73: 257-266Crossref Scopus (89) Google Scholar,10Janmaat K.R.L. Polansky L. Ban S.D. Boesch C. Wild chimpanzees plan their breakfast time, type, and location.Proc. Natl. Acad. Sci. USA. 2014; 111: 16343-16348Crossref PubMed Scopus (113) Google Scholar,14van Schaik C.P. Damerius L. Isler K. Wild orangutan males plan and communicate their travel direction one day in advance.PLoS One. 2013; 8e74896Crossref Scopus (39) Google Scholar,15Janson C.H. Experimental evidence for route integration and strategic planning in wild capuchin monkeys.Anim. Cogn. 2007; 10: 341-356Crossref PubMed Scopus (93) Google Scholar We report evidence that free-ranging wild fruit bats rely on mental temporal maps and exhibit future-oriented behaviors when foraging. We tracked young bats as they navigated and foraged, documenting every tree they visited over many months. We prevented the bats from foraging outside for different time periods and monitored their foraging decisions, revealing that the bats map the spatiotemporal patterns of resources in their environment. Following a long period in captivity, the bats did not visit those trees that were no longer providing fruit. We show that this time-mapping ability requires experience and is lacking in inexperienced bats. Careful analysis of the bats' movement and foraging choices indicated that they plan which tree to visit while still in the colony, thus exhibiting future-oriented behavior and delayed gratification on a nightly basis. Our findings demonstrate how the need for spatiotemporal mental mapping can drive the evolution of high cognitive abilities that were previously considered exclusive to humans.
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