{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,4,13]],"date-time":"2026-04-13T07:30:03Z","timestamp":1776065403488,"version":"3.50.1"},"reference-count":23,"publisher":"MDPI AG","issue":"3","license":[{"start":{"date-parts":[[2023,1,19]],"date-time":"2023-01-19T00:00:00Z","timestamp":1674086400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"University Research Expeditions Program (UREP), University of California, Berkeley"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Animals"],"abstract":"<jats:p>The size of adult male giraffes (Giraffa giraffa angolensis) far exceeds the size of the females. At the Namutoni waterholes in Etosha National Park, bulls were seen many times each day screening adult females for their pending sexual receptivity by provoking them to urinate; this mainly involved sniffing their genitalia. If the female accedes to the male\u2019s invitation, she widens her hindleg stance, braces her body, and then urinates, usually for at least five seconds. The male places his muzzle and tongue in the urine stream, and then performs flehmen, often raising his head high in the air. Males never investigated urine on the ground. The bilateral papillae on the giraffe\u2019s hard palate connect with the nasopalatine ducts, which enter the bilateral vomeronasal organ (VNO). Unlike many mammals, the giraffe\u2019s VNO lacks a prominent connection to the nasal cavity and its connections to the oral cavity are primarily via the incisive papillae. Most days, some giraffes were observed searching for bones for extended periods, chewing them, and sometimes being troubled by a bone stuck in their mouth. A giraffe killed by lions was frequented for several days by a procession of giraffes investigating the carcass. A very dark-colored bull giraffe emitted loud pulsed growls that drove off most of the surrounding giraffes.<\/jats:p>","DOI":"10.3390\/ani13030354","type":"journal-article","created":{"date-parts":[[2023,1,19]],"date-time":"2023-01-19T07:42:27Z","timestamp":1674114147000},"page":"354","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":7,"title":["Flehmen, Osteophagia, and Other Behaviors of Giraffes (Giraffa giraffa angolensis): Vomeronasal Organ Adaptation"],"prefix":"10.3390","volume":"13","author":[{"given":"Lynette A.","family":"Hart","sequence":"first","affiliation":[{"name":"Department of Population Health and Reproduction, School of Veterinary Medicine, University of California, Davis, CA 95616, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Benjamin L.","family":"Hart","sequence":"additional","affiliation":[{"name":"Department of Anatomy, Physiology and Cell Biology, School of Veterinary Medicine, University of California, Davis, CA 95616, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1968","published-online":{"date-parts":[[2023,1,19]]},"reference":[{"key":"ref_1","unstructured":"Moss, C. (1976). Portraits in the Wild: Animal Behaviour in East Africa, Hamish Hamilton."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"245","DOI":"10.1111\/j.1096-3642.1958.tb00687.x","article-title":"The behaviour of the giraffe, Giraffa camelopardalis, in eastern Transvaal","volume":"131","author":"Innis","year":"1958","journal-title":"Proc. Zool. Soc. Lond."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"686","DOI":"10.1515\/mamm.1970.34.4.686","article-title":"Flehmen","volume":"34","author":"Dagg","year":"1970","journal-title":"Mammalia"},{"key":"ref_4","unstructured":"Dagg, A.I., and Foster, J.B. (1976). The Giraffe: Its Biology, Behavior, and Ecology, Van Nostrand Reinholt."},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Dagg, A.I. (2014). Giraffe: Biology, Behaviour and Conservation, Cambridge University Press.","DOI":"10.1017\/CBO9781139542302"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"314","DOI":"10.1016\/j.yhbeh.2006.04.004","article-title":"Sociosexual behavior, male mating tactics, and the reproductive cycle of giraffe Giraffa camelopardalis","volume":"50","author":"Bercovitch","year":"2006","journal-title":"Horm. Behav."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"771","DOI":"10.1080\/00222938500770471","article-title":"Giraffe social behavior","volume":"19","author":"Pratt","year":"1985","journal-title":"J. Nat. Hist."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"771","DOI":"10.1086\/285955","article-title":"Winning by a neck: Sexual selection in the evolution of giraffe","volume":"148","author":"Simmons","year":"1996","journal-title":"Am. Nat."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"257","DOI":"10.1111\/j.1469-7998.1999.tb00989.x","article-title":"The remarkable anatomy of the giraffe\u2019s neck","volume":"247","author":"Solounias","year":"1999","journal-title":"J. Zool. Lond."},{"key":"ref_10","unstructured":"Brand, R. (2007). Evolutionary Ecology of Giraffes (Giraffa camelopardalis) in Etosha National Park, Namibia. [Ph.D. Thesis, Newcastle University]."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1111\/mam.12268","article-title":"A review of the social behaviour of the giraffe Giraffa camelopardalis: A misunderstood but socially complex species","volume":"52","author":"Muller","year":"2021","journal-title":"Mammal Rev."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"421","DOI":"10.1093\/icb\/7.3.421","article-title":"Olfaction in mammals","volume":"7","author":"Moulton","year":"1967","journal-title":"Am. Zool."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"315","DOI":"10.1515\/mamm.1972.36.3.315","article-title":"The role of the vomeronasal organ in mammalian reproduction","volume":"36","author":"Estes","year":"1972","journal-title":"Mammalia"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"353","DOI":"10.1016\/0093-691X(80)90047-3","article-title":"Tongue manipulation of the palate assists estrous detection in the bovine","volume":"13","author":"Jacobs","year":"1980","journal-title":"Theriogenology"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"2913","DOI":"10.1242\/jeb.201.21.2913","article-title":"Structure and function of the vomeronasal organ","volume":"201","author":"Doving","year":"1998","journal-title":"J. Exp. Biol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"1067","DOI":"10.1016\/0031-9384(80)90049-9","article-title":"Flehmen and vomeronasal function in male goats","volume":"24","author":"Ladewig","year":"1980","journal-title":"Physiol. Behav."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"652","DOI":"10.1002\/jemt.22843","article-title":"Histological features of the vomeronasal organ in the giraffe, Giraffa camelopardalis","volume":"80","author":"Kondoh","year":"2017","journal-title":"Microsc. Res. Tech."},{"key":"ref_18","first-page":"197","article-title":"Chemosensory investigation, flehmen behaviour and vomeronasal organ function in antelope","volume":"Volume 61","author":"Jewell","year":"1989","journal-title":"Symposia of the Zoological Society of London: The Biology of Large African Mammals in Their Environment"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"299","DOI":"10.1037\/0735-7036.101.4.299","article-title":"Species-specific patterns of urine investigation and flehmen in Grant\u2019s Gazelle (Gazella granti), Thomson\u2019s Gazelle (G. thomsoni), Impala (Aepyceros melampus), and Eland (Taurotragus oryx)","volume":"101","author":"Hart","year":"1987","journal-title":"J. Comp. Psychol."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"155","DOI":"10.1016\/0031-9384(88)90291-0","article-title":"Alteration in vomeronasal system anatomy in alcelaphine antelopes: Correlation with alteration in chemosensory investigation","volume":"42","author":"Hart","year":"1988","journal-title":"Physiol. Behav."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"4139","DOI":"10.1016\/j.jas.2013.06.004","article-title":"Osteophagia and bone modifications by giraffe and other large ungulates","volume":"40","author":"Hutson","year":"2013","journal-title":"J. Archaeol. Sci."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1007\/s00265-018-2504-8","article-title":"The bone collector: Temporal patterns of bone-gnawing behaviour define osteophagia as a female prerogative in a large rodent","volume":"72","author":"Mori","year":"2018","journal-title":"Behav. Ecol. Sociobiol."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"271","DOI":"10.1016\/j.ygcen.2005.01.011","article-title":"Fecal steroid analysis of female giraffe (Giraffa camelopardalis) reproductive condition and the impact of endocrine status on daily time budgets","volume":"141","author":"Bashaw","year":"2005","journal-title":"Gen. Comp. Endocrinol."}],"container-title":["Animals"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2076-2615\/13\/3\/354\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,10]],"date-time":"2025-10-10T18:10:44Z","timestamp":1760119844000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2076-2615\/13\/3\/354"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2023,1,19]]},"references-count":23,"journal-issue":{"issue":"3","published-online":{"date-parts":[[2023,2]]}},"alternative-id":["ani13030354"],"URL":"https:\/\/doi.org\/10.3390\/ani13030354","relation":{},"ISSN":["2076-2615"],"issn-type":[{"value":"2076-2615","type":"electronic"}],"subject":[],"published":{"date-parts":[[2023,1,19]]}}}