We trace the history of the endangered Northern Bald Ibis through different epochs to the present. A particular focus is placed on its life in and disappearance from ancient Egypt, where the bird attained great cultural and religious significance, and on the modern endeavour to re-wild the species. Due to the characteristic appearance, behaviour and habitat of the species as well as its need for open foraging areas, a close mutualistic relationship between humans and the birds was formed in ancient Egypt, as in other cultures. A clear benefit for the Northern Bald Ibis was the availability of feeding habitats, which were cleared by humans for farming or grazing. The benefit to people was rather cultural because the bird attracted religious veneration or symbolic meanings from ancient Egypt to medieval Europe. The proximity to humans, however, carried a high risk as well. We discuss various types of impact (including human impacts as well as climate change) as triggers for the extinction of the species. The evidence for a triple disappearance of the Northern Bald Ibis (around 2000 BCE, around 1600 CE and in modern time) represents a unique basis for studying both the bird's habitat preferences and its vulnerability. This is because different, mainly anthropogenic, causes stood behind these three historical disappearances, although the disappearances in all three epochs occurred during a period of climate change.

Czech Institute of Egyptology Charles University 11000 Prague Czech Republic

Department of Behavioral and Cognitive Biology University of Vienna 1090 Vienna Austria

Waldrappteam Conservation and Research 6162 Mutters Austria

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Bennett N.J., Roth R., Klain S.C., Chan K.M.A., Clark D.A., Cullman G., Epstein G., Nelson M.P., Stedman R., Teel T.L., et al. Mainstreaming the social sciences in conservation. Conserv. Biol. 2017;31:56–66. doi: 10.1111/cobi.12788. PubMed DOI

Campbell L.M. Overcoming obstacles to interdisciplinary research. Conserv. Biol. 2005;19:574–577. doi: 10.1111/j.1523-1739.2005.00058.x. DOI

Janák J. Running with images: Ritualized script in the Vogellauf, Rudderlauf and Vasenlauf. In: Bárta J.J.M., editor. Profane Landscapes, Sacred Spaces. Equinox; Sheffield, UK: 2020. pp. 89–96.

Janák J. Spotting the Akh. The Presence of the Northern Bald Ibis in Ancient Egypt and Its Early Decline. J. Am. Res. Cent. Egypt. 2011;46:17–32.

Fritz J., Unsoeld M., Voelkl B. Back into European Wildlife: The Reintroduction of the Northern Bald Ibis (Geronticus eremita) In: Kaufman A., Bashaw M., Maple T., editors. Scientific Foundations of Zoos and Aquariums: Their Role in Conservation and Research. Cambridge University Press; Cambridge, UK: Cambridge University Press; Cambridge, UK: 2019. 2019. pp. 339–366.

Fritz J., Unsoeld M. Artenschutz für einen histrosichen Schweizer Vogel: Der Waldrapp im Aufwind. Wildbiol. Int. 2011;5:121–138.

Pegoraro K. Der Waldrapp. AULA-Verlag; Wiesbaden, Germany: 1996.

Hirsch U. Die Rettung der heiligen Vögel. Tierpark. 1976;9:4–11.

Fritz J. The European LIFE+ northern bald ibis reintroduction project. Oryx. 2021;55:809–810. doi: 10.1017/S003060532100123X. DOI

Fritz J., Kramer R., Hoffmann W., Trobe D., Unsöld M. Back into the wild: Establishing a migratory Northern bald ibis Geronticus eremita population in Europe. Int. Zoo Yearb. 2017;51:107–123. doi: 10.1111/izy.12163. DOI

Fritz J., Eberhard B., Esterer C., Goenner B., Trobe D., Unsoeld M., Voelkl B., Wehner H., Scope A. Biologging is suspect to cause corneal opacity in two populations of wild living Northern Bald Ibises (Geronticus eremita) Avian Res. 2020;11:38. doi: 10.1186/s40657-020-00223-8. DOI

Wehner H., Huchler K., Fritz J. Quantification of Foraging Areas for the Northern Bald Ibis (Geronticus eremita) in the Northern Alpine Foothills: A Random Forest Model Fitted with Optical and Actively Sensed Earth Observation Data. Remote Sens. 2022;14:1015. doi: 10.3390/rs14041015. DOI

Fritz J., Wirtz S., Unsoeld M. Aspekte der Nahrungsökologie und Genetik des Waldrapps: Reply zu Bauer et al. (2016) Vogelneozoen in Deutschland—Revision der nationalen Statuseinstufungen. Vogelwarte. 2017;55:141–145.

Sperger C., Heller A., Voelkl B., Fritz J. Flight Strategies of Migrating Northern Bald Ibises—Analysis of GPS Data during Human-led Migration Flights. Agit J. Für Angew. Geoinf. 2017;3:62–72. doi: 10.14627/537633007. DOI

Portugal S.J., Hubel T.Y., Fritz J., Heese S., Trobe D., Voelkl B., Hailes S., Wilson A.M., Usherwood J.R. Upwash exploitation and downwash avoidance by flap phasing in ibis formation flight. Nature. 2014;505:399–402. doi: 10.1038/nature12939. PubMed DOI

Voelkl B., Portugal S.J., Unsoeld M., Usherwood J.R., Wilson A.M., Fritz J. Matching times of leading and following suggest cooperation through direct reciprocity during V-formation flight in ibis. Proc. Natl. Acad. Sci. USA. 2015;112:2115–2120. doi: 10.1073/pnas.1413589112. PubMed DOI PMC

Drenske S., Radchuk V., Scherer C., Esterer C., Kowarik I., Fritz J., Kramer-Schadt S. On the road to selfsustainability: Reintroduced migratory European Northern Bald Ibises (Geronticus eremita) still need management interventions for population viability. bioRxiv. 2021;14 doi: 10.1101/2021.04.03.438331. DOI

Wirtz S., Boehm C., Fritz J., Kotrschal K., Veith M., Hochkirch A. Optimizing the genetic management of reintroduction projects: Genetic population structure of the captive Northern Bald Ibis population Sarah Wirtz. Conserv. Genet. 2018;19:853–864. doi: 10.1007/s10592-018-1059-6. DOI

Boehm C., Bowden C., Seddon P., Hatipoglu T., Oubrou W., el Bekkay M., Quevedo M., Fritz J., Yeniyurt C., Lopez J., et al. Northern Bald Ibis: History, current status, and future perspectives. Oryx. 2021;55:934–946. doi: 10.1017/S0030605320000198. DOI

Bowden C.G.R., Aghnaj A., Smith K.W., Ribi M. The status and recent breeding performance of the critically endangered Northern Bald Ibis Geronticus eremita population on the Atlantic coast of Morocco. Ibis. 2003;145:419–431. doi: 10.1046/j.1474-919X.2003.00181.x. DOI

Gesner C. Historiae Animalium Liber III, qui est de Avium Natura. Christoffel Froschouer; Zurich, Switzerland: 1557. Erste Deutsche Übersetzung.

Schenker A. Das ehemalige Verbreitungsgebiet des Waldrapps Geronticus eremita in Europa. Der Ornithol. Beob. 1977;74:13–30.

Unsoeld M., Fritz J. Der Waldrapp-ein Vogel zwischen Ausrottung und Wiederkehr. Wildbiologie. 2011;2:1–16.

Kumerloeve H. Waldrapp, Gerontocus eremita (LINNAEUS 1758) und Glattnackenrapp, Geronticus calvus (BODDAERT 1783): Zur Geschichte ihrer Erforschung und zur gegenwärtigen Bestandssituation. Ann. Naturhistor. Mus. Wien. 1978;81:319–349.

Sánchez I. Evidence of the historic presence of the Northern Bald Ibis (Geronticus eremita) in Spain. 2006. [(accessed on 11 March 2022)]. Available online: https://www.researchgate.net/publication/235994501.

Perco F., Tout P. Notes on recent discoveries regarding the presence of the Northern Bald Ibis Geronticus eremita in the Upper Adriatic Region. Acrocephalus. 2001;22:81–87.

Boev Z. Presence of Bald Ibises (Geronticus Wagler 1832) (Threskionitidae (sic)—Aves) in the Late Pliocene of Bulgaria. Geol. Balc. 1998;28:45–52.

Hölzinger J. Waldrapp (Geronticus eremita): Knochenfunde aus der spätrömischen Befestigung Sponeck am Kaiserstuhl. Orn. Jh. Baden Württemberg. 1988;4:57–67.

Mourer-Chauviré C., Philippe M., Guillard S., Meyssonnier M. Presence of the Northern Bald Ibis Geronticus eremita (L.) during the Holocene in the Ardèche valley, southern France. Ibis. 2006;148:820–823. doi: 10.1111/j.1474-919X.2006.00563.x. DOI

Schenker A. Der Waldrapp—ein historisches Wildbret. Wildbiologie. 1981;4:1–12.

Kinzelbach R.K. The distribution of the serin (Serinus serinus L.; 1766) in the 16th century. J. Ornithol. 2004;145:177–187. doi: 10.1007/s10336-004-0038-5. DOI

Bauer K.M., Blutz von Blotzheim U.N. Handbuch der Vögel Mitteleuropas. Ak. Verlagsgesellschaft; Frankfurt, Germany: 1966.

Peter H. Waldrappdämmerung am Euphrat. Max Kasparek Verlag; Heidelberg, Germany: 1990.

Hirsch U. Der Waldrapp Geronticus eremita, ein Beitrag zur Situation in seinem östlichen Verbreitungsgebiet. Vogelwelt. 1980;101:219–236.

Hatipoglu T. Report of 4th IAGNBI Meeting Seekirchen. In: Boehm C., Bowden C., editors. Alpenzoo; Innbruck, Austria: 2016. pp. 40–46.

Yeniyurt C., Oppel S., Isfendiyaroglu S., Özkinaci G., Erkol I.L., Bowden C. Influence of feeding ecology on breeding success of a semi-wild population of the critically endangered Northern Bald Ibis Geronticus eremita in southern Turkey. Bird Conserv. Int. 2016;27:1–13. doi: 10.1017/S0959270916000253. DOI

Serra G., Abdallah M.S., Assaed A., Abdallah A., al Qaim G., Fayad T., Willliamson D. Discovery of a relict breeding colony of northern bald ibis Geronticus eremita in Syria. Oryx. 2004;38:1–7. doi: 10.1017/S003060530400016X. DOI

Serra G., Lindsell J.A., Peske L., Fritz J., Bowden C.G.R., Bruschini C., Welch G., Tavares J.P., Wondafrash M. Accounting for the low survival of the Critically Endangered northern bald ibis Geronticus eremita on a major migratory flyway. Oryx. 2014;49:312–320. doi: 10.1017/S0030605313000665. DOI

Lindsell J.A., Serra G., Peške L., Abdullah M.S., Al Qaim G., Kanani A., Wondafrash M. Satellite tracking reveals the migration route and wintering area of the Middle East population of Critically Endangered northern bald ibis Geronticus eremita. Oryx. 2009;43:329–335. doi: 10.1017/S0030605309001963. DOI

Fritz J., Riedler B. Neue Hoffnung für das Überleben einer hoch bedrohtesten Zugvogelart im Mittleren Osten: Freisetzung von Jungvögeln bei den letzten migrierenden Waldrappen in Syrien. Vogelwarte. 2010;48:417–418.

Serra G. The Northern Bald Ibis is extinct in the Middle East – but we can’t blame it on IS. The Ecologist. 2015. [(accessed on 11 March 2022)]. Available online: https://theecologist.org/2015/may/29/northern-bald-ibis-extinct-middle-east-we-cant-blame-it.

Friedman F. Ph.D. Thesis. Brown University; Providence, RI, USA: 1981. On the Meaning of Akh in Egyptian Mortuary Texts.

Friedman F. A Review of Akh: Une Notion Religieuse Dans L’Égypte Pharaonique by Gertie Englund. J. Am. Res. Cent. Egypt. 1982;19:145–148. doi: 10.2307/40000440. DOI

Friedman F. In: Oxford Encyclopedia of Ancient Egypt, I. Redford D.B., editor. Oxford University Press; Oxford, UK: 2001.

Smith M. Traversing Eternity: Texts for the Afterlife from Ptolemaic and Roman Egypt. Oxford University Press; Oxford, UK: 2009.

Janák J. Staroegyptské Náboženství II. Život a Úděl Člověka. Oikoymenh; Prague, Czech Republic: 2012.

Assmann J. Tod und Jenseits im alten Ägypten. C.H. Beck; München, Germany: 2001.

Janák J. UCLA Encyclopedia of Egyptology. University of California in Los Angeles; Los Angeles, CA, USA: 2013. Northern Bald Ibis (Akh-Bird)

Leprohon R.J. Gatekeepers of This and the Other World. J. Soc. Study Egypt. Antiq. 1994;24:77–91.

Jansen-Winkler K. Horizont und Verklärheit: Zur Bedeutung der Wurzel Ax. Stud. Altägyptischen Kult. 1996;23:201–215.

Allen J.P. Reading a Pyramid. In: Berger C., editor. Homages à Jean Leclant. Volume 1: Études Pharaoniques. Orientale, Institut Français d’Archéologie; Cairo, Egypt: 1994.

Hays H.M. Unreading the Pyramids. Le Bull. L’inst. Franç. D’archéol. Orient. 2009;109:195–220.

Fritz J., Unsoeld M. Aufwind für den Waldrapp: Von der Wiederansiedlung eines europäischen Zugvogels. Jahrb. Ver. Zum Schutz Bergwelt. 2013;78:121–138.

Zoufal K., Fritz J., Bichler M., Kirbauer M., Markut T., Meran I., Wolf A., Kotrschal K. Feeding Ecology of the Northern Bald Ibis in Different Habitat Types: An Experimental Field Study with Handraised Individuals. In: Böhm C., Bowden C., editors. Report of the 2nd IAGNBI Meeting, Vejer 2006. Alpenzoo; Innsbruck, Austria: 2007. pp. 77–84.

Serra G., Abdallah M.S., al Qaim G. Feeding ecology and behaviour of the last known surviving Northern Bald Ibises, Gerontius eremita, at theier breeing quarter in Syria. Zool. Middle East. 2008;43:55–68. doi: 10.1080/09397140.2008.10638269. DOI

Serra G., Bruschini C., Peske L., Kubsa A., Wondafrash M., Lindsell J.A. An assessment of ecological conditions and threats at the Ethiopian wintering site of the last known eastern colony of Critically Endangered Northern Bald Ibis Geronticus eremita. Bird Conserv. Int. 2013;23:399–413. doi: 10.1017/S0959270913000397. DOI

Bowden C.G.R., Smith K.W., El xBekkay M., Oubrou W., Aghnaj A., Jimenez-Armesto M. Contribution of research to conservation action for the Northern Bald Ibis Geronticus eremita in Morocco. Bird Conserv. Int. 2008;18:74–90. doi: 10.1017/S0959270908000403. DOI

Serra G., Peske L., Abdallah M.S., al Qaim G., Kanani A. Breeding ecology and behaviour of the last wild oriental Northern Bald Ibises (Geronticus eremita) in Syria. J. Ornithol. 2009;150:769–782. doi: 10.1007/s10336-009-0398-y. DOI

Schenker A., Cahenzli F., Gutbrod K.G., Thevenot M., Erhardt A. The Northern Bald Ibis Geronticus eremita in Morocco since 1900: Analysis of ecological requirements. Bird Conserv. Int. 2019;30:117–138. doi: 10.1017/S0959270919000170. DOI

Yeniyurt C. Conservation of the Northern Bald Ibis, Birecik, Turkey 2013–2016. In: Böhm C., Bowden C., editors. Report of 4th IAGNBI Meeting Seekirchen. Alpenzoo; Innsbruck, Austria: 2016. pp. 34–39.

Smith K.W., Aghnaj A., El Bekkay M., Oubrou W., Ribi M., Armesto M.J., Bowden C.G.R. The provision of supplementary fresh water improves the breeding success of the globally threatened Northern Bald Ibis Geronticus eremita. Ibis. 2008;150:728–734. doi: 10.1111/j.1474-919X.2008.00844.x. DOI

Sorato E., Kotrschal K. Hormonal and behavioural symmetries between the sexes in the Northern bald ibis. Gen. Comp. Endocrinol. 2006;146:265–274. doi: 10.1016/j.ygcen.2005.11.020. PubMed DOI

Pegoraro K., Föger M. Die Chrup-Rufe des Waldrapps Geronticus eremita: Ihre verschiedenen Funktionen in einem komplexen Sozialsystem. J. Ornithol. 1995;136:243–252. doi: 10.1007/BF01651291. DOI

Szipl G., Boeckle M., Werner S.A.B., Kotrschal K. Mate recognition and expression of affective state in croop calls of Northern Bald Ibis (Geronticus eremita) PLoS ONE. 2014;9:e88265. doi: 10.1371/journal.pone.0088265. PubMed DOI PMC

Pegoraro K., Föger M. Individuality in the Northern Bald Ibis or Waldrapp Ibis Geronticus eremita—Key features for a complex social system. Acrocephalus. 2001;22:73–79.

Sorato E., Kotrschal K. Skin Ornaments Reflect Social Status and Immunocompetence in Male and Female Northern Bald Ibises (Geronticus eremita) bioRxiv. 2020 doi: 10.1101/2020.11.25.397570. DOI

Galván I., Palacios D., Negro J.J. The bare head of the Northern bald ibis (Geronticus eremita) fulfills a thermoregulatory function. Front. Zool. 2017;14:1–9. doi: 10.1186/s12983-017-0201-5. PubMed DOI PMC

Unsoeld M., Melzer R. Sunning behaviour in Ibises (Threskiornithidae): Observations on four species at Tierpark Hellabrunn, Munich Markus. Der Zooloigsche Gart. 2010;79:89–104. doi: 10.1016/j.zoolgart.2010.07.001. DOI

Hegemann A., Fudickar A.M., Nilsson J.-Å. A physiological perspective on the ecology and evolution of partial migration. J. Ornithol. 2019;160:893–905. doi: 10.1007/s10336-019-01648-9. DOI

Fritz J., Feurle A., Kotrschal K. Physiological regulation of bird migration: A study with northern bald ibises undergoing human-led autumnal migration. J. Ornithol. 2016;147:168.

Bairlein F., Fritz J., Scope A., Schwendenwein I., Stanclova G., Van Dijk G., Meijer H.A.J., Verhulst S., Dittami J. Energy expenditure and metabolic changes of free-flying migrating northern bald ibis. PLoS ONE. 2015;10:e0134433. doi: 10.1371/journal.pone.0134433. PubMed DOI PMC

Fritz J. Ultraleichtflieger weisen den Weg—Der Waldrapp in den Alpen. Der Falke. 2010;57:95–105.

Aourir M., Bousadik H., El Bekkay M., Oubrou W., Znari M., Qninba A. New Breeding Sites of the Critically Endangered Northern Bald Ibis Geronticus Eremita on the Moroccan Atlantic Coast. Int. J. Avian Wildl. Biol. 2017;2:1–4. doi: 10.15406/ijawb.2017.02.00021. DOI

Touti J., Oumellouk F., Bowden C.G.R., Kirkwood J.K., Smith K.W. Mortality incident in northern bald ibis Geronticus eremita in Morocco in May 1996. Oryx. 1999;33:160–167. doi: 10.1046/j.1365-3008.1999.00050.x. DOI

Schilling J., Freier K.P., Hertig E., Scheffran J. Climate change, vulnerability and adaptation in North Africa with focus on Morocco. Agric. Ecosyst. Environ. 2012;156:12–26. doi: 10.1016/j.agee.2012.04.021. DOI

Visser M.E., Perdeck A.C., van Balen J.H., Both C. Climate change leads to decreasing bird migration distances. Glob. Chang. Biol. 2009;15:1859–1865. doi: 10.1111/j.1365-2486.2009.01865.x. DOI

Radchuk V., Reed T., Teplitsky C., van de Pol M., Charmantier A., Hassall C., Adamík P., Adriaensen F., Ahola M.P., Arcese P., et al. Adaptive responses of animals to climate change are most likely insufficient. Nat. Commun. 2019;10:3109. doi: 10.1038/s41467-019-10924-4. PubMed DOI PMC

Boessneck J. Die Tierwelt des Alten Ägypten. C.H. Beck; Munich, Germany: 1988.

Ikram S. Divine Creatures: Animal Mummies in Ancient Egypt. American University in Cairo Press; Cairo, Egypt: 2005.

Houlihan P. The Birds of Ancient Egypt. American University in Cairo Press; Cairo, Egypt: 1988.

Bárta M. Kolaps Staré říše, éry stavitelů pyramid. In: Pokorná P., Bárta M., editors. Něco Překrásného se Končí. Kolapsy v Přírodě a Společnosti. Dokořán; Prague, Czech Republic: 2008. pp. 121–144.

Dalfes H.N., Kukla G., Weiss H. Third Millennium BC Climate Change and Old World Collapse. Springer; Berlin, Germany: New York, NY, USA: 1997.

Issar A.S., Zohar M. Climate Change: Environment and Civilization in the Middle East. Springer; Berlin, Germany: New York, NY, USA: 2004.

Janák J. UCLA Encyclopedia of Egyptology. University of California in Los Angeles; Los Angeles, CA, USA: 2014. Saddle-billed Stork (ba-bird)

Janák J. Running with Images: Ritualised script in the Vogellauf, Rudderlauf and Vasenlauf. In: Levy T.E., editor. New Directions in Antrophology. University of California, San Diego; La Jolla, CA, USA: 2020. pp. 89–96.

Schenker A. Replik zum Beitrag von Armin Landmann betreffend den Waldrapp Geronticus eremita. Vogelwarte. 2017;55:129–138.

Bowden C.G.R. (Compiler) International Single Species Action Plan for the Conservation of the Northern Bald Ibis (Geronticus eremita) Volume 55 The Royal Society for the Protection of Birds; Bonn, Germany: 2015.

Lelieveld J., Hadjinicolaou P., Kostopoulou E., Chenoweth J., El Maayar M., Giannakopoulos C., Hannides C., Lange M.A., Tanarhte M., Tyrlis E., et al. Climate change and impacts in the Eastern Mediterranean and the Middle East. Clim. Change. 2012;114:667–687. doi: 10.1007/s10584-012-0418-4. PubMed DOI PMC

Demarée R.J. The Ax iqr n Ra-stelae on Ancestor Worship in Ancient Egypt. Leiden, The Netherlands: 1983.

Allen J.P. Religion and Philosophy in Ancient Egypt, Yale Egyptological Studies 3. Yale Universitxy Press; New Haven, CT, USA: 1989. The Cosmology of the Pyramid Texts; pp. 1–29.

Dunham D. An Egyptian diadem of the Old Kingdom. Bull. Museum Fine Arts. 1946:23–29.

Staehelin E. Ph.D. Thesis. Verlag Bruno Hessling; Berlin, Germany: 1966. Untersuchungen zur Agyptischen Tracht im Alten Reich.

Janák J. A question of size: A remark on early attestations of the ba hieroglyph. Stud. zur Altägyptischen Kult. 2011:143–153.

Kees H. Hinrichs J.C.; München, Germany: 1912.

Decker M., Herb W. Bildatlas zum Sport im alten Ägypten. Corpus der bildlichen Quellen zu Leibesübungen, Spiel, Jagd, Tanz und verwandten Themen. Brill; Leiden, The Netherlands: 1994.