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{{short description|Tall African hoofed mammal}} {{other uses|Giraffe (disambiguation)}} {{featured article}} {{pp-semi-indef}} {{Use dmy dates|date=December 2023}} {{Automatic taxobox | name = Giraffes | fossil_range = {{Fossilrange|11.61|0}} [[Miocene]] to [[Holocene|Recent]] | image = Giraffe Mikumi National Park.jpg | image_caption = [[Masai giraffe]] (''Giraffa tippelskirchi'') in [[Mikumi National Park]], [[Tanzania]] | status = VU | status_system = IUCN3.1 | status_ref = <ref name=iucn>{{cite iucn |author=Muller, Z. |author2=Bercovitch, F. |author3=Brand, R. |author4=Brown, D. |author5=Brown, M. |author6=Bolger, D. |author7=Carter, K. |author8=Deacon, F. |author9=Doherty, J.B. |author10=Fennessy, J. |author11=Fennessy, S. |author12=Hussein, A.A. |author13=Lee, D. |author14=Marais, A. |author15=Strauss, M. |author16=Tutchings, A. |author17=Wube, T. |date=2018 |amends=2016 |title=''Giraffa camelopardalis'' |volume=2018 |page=e.T9194A136266699 |doi=10.2305/IUCN.UK.2016-3.RLTS.T9194A136266699.en |access-date=12 November 2021}}</ref> (As the species complex) <!-- refers to species | status = VU | status_system = IUCN3.1 | status_ref = <ref name=":3">{{cite iucn |author=Muller, Z. |author2=Bercovitch, F. |author3=Brand, R. |author4=Brown, D. |author5=Brown, M. |author6=Bolger, D. |author7=Carter, K. |author8=Deacon, F. |author9=Doherty, J.B. |author10=Fennessy, J. |author11=Fennessy, S. |author12=Hussein, A.A. |author13=Lee, D. |author14=Marais, A. |author15=Strauss, M. |author16=Tutchings, A. |author17=Wube, T. |date=2018 |title=''Giraffa camelopardalis'' |volume=2018 |page=e.T9194A136266699 |doi=10.2305/IUCN.UK.2016-3.RLTS.T9194A136266699.en |access-date=19 November 2021}}</ref> --> | taxon = Giraffa | authority = Brisson, 1762 | subdivision_ranks = Species | subdivision = [[#Species and subspecies|See taxonomy]] | range_map = Giraffa_camelopardis_distribution_2018.png | range_map_caption = Distribution of the giraffe }} The '''giraffe''' is a large [[Fauna of Africa|African]] [[even-toed ungulate|hoofed mammal]] belonging to the [[genus]] '''''Giraffa.''''' It is the [[Largest mammals#Even-toed Ungulates (Artiodactyla)|tallest]] living [[terrestrial animal]] and the largest [[ruminant]] on [[Earth]]. Traditionally, giraffes have been thought of as one [[species]], ''[[Giraffa camelopardalis]]'', with nine [[subspecies]]. Most recently, researchers proposed dividing them into four [[Neontology#Extant taxa versus extinct taxa|extant]] species due to new research into their [[Mitochondrial DNA|mitochondrial]] and [[nuclear DNA]], and individual species can be distinguished by their fur [[Animal coat|coat]] patterns. Six valid [[Lists of extinct species|extinct species]] of ''Giraffa'' from Africa and Pakistan are known from the [[fossil]] record. The giraffe's distinguishing characteristics are its extremely long neck and legs, horn-like [[ossicone]]s, and spotted coat patterns. It is classified under the [[Family (biology)|family]] [[Giraffidae]], along with its closest extant relative, the [[okapi]]. Its scattered range extends from [[Chad]] in the north to [[South Africa]] in the south and from [[Niger]] in the west to [[Somalia]] in the east. Giraffes usually inhabit [[savannah]]s and [[woodland]]s. Their food source is leaves, fruits, and flowers of woody plants, primarily [[Acacieae|acacia]] species, which they browse at heights most other ground-based [[herbivore]]s cannot reach. [[Lion]]s, [[leopard]]s, [[spotted hyena]]s, and [[African wild dog]]s may [[prey]] upon giraffes. Giraffes live in herds of related females and their offspring or bachelor herds of unrelated adult males but are [[gregarious]] and may gather in large groups. Males establish social hierarchies through "necking", combat bouts where the neck is used as a weapon. [[dominance (ethology)|Dominant]] males gain mating access to females, which bear sole responsibility for rearing the young. The giraffe has intrigued various ancient and modern cultures for its peculiar appearance and has often been featured in paintings, books, and cartoons. It is classified by the [[International Union for Conservation of Nature]] (IUCN) as [[Vulnerable species|vulnerable]] to extinction. It has been [[Local extinction|extirpated]] from many parts of its former range. Giraffes are still found in many [[national park]]s and [[game reserve]]s, but estimates as of 2016 indicate there are approximately 97,500 members of ''Giraffa'' in the wild. More than 1,600 were kept in zoos in 2010. ==Etymology== The name "giraffe" has its earliest known origins in the [[Arabic language|Arabic]] word {{transliteration|ar|zirāfah}} ({{lang|ar|زِرَافَةْ}}), of an ultimately unclear [[Languages of Africa|Sub-Saharan African language]] origin.<ref name=OED/> The [[Middle English]] and [[early Modern English]] spellings, {{lang|enm|jarraf}} and {{lang|enm|ziraph}}, derive from the Arabic form-based [[Spanish language|Spanish]] and [[Portuguese language|Portuguese]] ''girafa''.<ref>{{cite book |last=Őrsi |first=Tibor |author-link= |date=2006 |title=French Linguistic Influence in the Cotton Version of Mandeville's Travels |url=https://books.google.com/books?id=qQUSAQAAIAAJ&q=%22jarraf%22 |location= |publisher=Tinta Könyvkiadó |page=113 |isbn=9789637094545 |access-date=20 April 2021 |archive-date=22 September 2023 |archive-url=https://web.archive.org/web/20230922073613/https://books.google.com/books?id=qQUSAQAAIAAJ&q=%22jarraf%22 |url-status=live }}</ref> The modern English form developed around 1600 from the [[French language|French]] {{lang|fr|girafe}}.<ref name=OED>{{cite encyclopedia|url= http://etymonline.com/?term=giraffe|title= Giraffe|dictionary= Online Etymology Dictionary|access-date= 1 November 2011|archive-date= 19 March 2015|archive-url= https://web.archive.org/web/20150319133340/http://etymonline.com/?term=giraffe|url-status= live}}</ref> "Camelopard" ({{IPAc-en|k|ə|ˈ|m|ɛ|l|ə|ˌ|p|ɑr|d}}) is an [[archaism|archaic]] English name for the giraffe; it derives from the [[Ancient Greek]] {{lang|grc|καμηλοπάρδαλις}} ({{transliteration|grc|kamēlopárdalis}}), from {{lang|grc|κάμηλος}} ({{transliteration|grc|kámēlos}}), "[[camel]]", and {{lang|grc|πάρδαλις}} ({{transliteration|grc|párdalis}}), "[[leopard]]", referring to its camel-like shape and leopard-like colouration.<ref>{{cite web|title=Definition of CAMELOPARD|url=http://www.merriam-webster.com/dictionary/camelopard|website=m-w.com|publisher=Encyclopædia Britannica: Merriam-Webster|access-date=3 September 2014|archive-date=25 April 2009|archive-url=https://web.archive.org/web/20090425003508/http://www.merriam-webster.com/dictionary/camelopard|url-status=live}}</ref><ref>{{cite web|title=Definition of camelopard|url=http://medieval_terms.enacademic.com/615/Camelopard|work=Dictionary of Medieval Terms and Phrases|access-date=3 September 2014|archive-date=4 September 2014|archive-url=https://web.archive.org/web/20140904044941/http://medieval_terms.enacademic.com/615/Camelopard|url-status=dead}}</ref> ==Taxonomy== ===Evolution=== The giraffe is one of only two living genera of the family Giraffidae in the order [[Artiodactyla]], the other being the [[okapi]].<ref name="Mitchell20003"/> They are [[ruminant]]s of the clade [[Pecora]], along with [[Antilocapridae]] ([[pronghorn]]s), [[Cervidae]] (deer), [[Bovidae]] (cattle, antelope, goats and sheep) and [[Moschidae]] (musk deer). A 2019 [[genome]] study (cladogram below) finds that Giraffidae are a [[sister taxon]] to Antilocapridae, with an estimated [[Split (phylogenetics)|split]] of over 20 million years ago.<ref name=RumiantPhylo2020>{{Cite journal |last1=Chen |first1=L. |last2=Qiu |first2=Q. |last3=Jiang |first3=Y. |last4=Wang |first4=K. |title=Large-scale ruminant genome sequencing provides insights into their evolution and distinct traits |doi=10.1126/science.aav6202 |journal=Science |volume=364 |issue=6446 |page=eaav6202 |year=2019 |pmid=31221828 |bibcode=2019Sci...364.6202C |doi-access=free}}</ref> {{Clade | style=font-size: 100%; line-height:100% |label1=[[Ruminantia]] |1={{clade |label1=[[Tragulina]] |1=[[Tragulidae]] [[File:Tragulus napu - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - (white background).jpg|50 px]] |label2=[[Pecora]] |2={{clade |1={{clade |1=[[Antilocapridae]] [[File:Antilocapra white background.jpg|50 px]] |2=[[Giraffidae]] [[File:Giraffa camelopardalis Brockhaus white background.jpg|50 px]] }} |2={{clade |1=[[Cervidae]] [[File:The deer of all lands (1898) Hangul white background.png|50 px]] |2={{clade |1=[[Bovidae]] [[File:Birds and nature (1901) (14562088237) white background.jpg|50px]] |2=[[Moschidae]] [[File:Moschus chrysogaster white background.jpg|50 px]] }} }} }} }} }} The family Giraffidae was once much more extensive, with over 10 fossil [[Genus|genera]] described.<ref name=Mitchell20003>{{cite journal |author1=Mitchell, G. |author2=Skinner, J. D. |year=2003 |title=On the origin, evolution, and phylogeny of giraffes ''Giraffa camelopardalis'' |journal=[[Transactions of the Royal Society of South Africa]] |volume=58 |issue=1|pages=51–73 |doi=10.1080/00359190309519935 |bibcode=2003TRSSA..58...51M |s2cid=6522531}}</ref> The elongation of the neck appears to have started early in the giraffe [[Lineage (evolution)|lineage]]. Comparisons between giraffes and their ancient relatives suggest vertebrae close to the skull lengthened earlier, followed by lengthening of vertebrae further down.<ref name="Danowitz2015">{{cite journal|author1=Danowitz, M. |author2=Vasilyev, A. |author3=Kortlandt, V. |author4=Solounias, V. |year=2015|title=Fossil evidence and stages of elongation of the ''Giraffa camelopardalis'' neck |journal=[[Royal Society Open Science]]|volume=2|issue=10 |page=150393 |doi=10.1098/rsos.150393 |pmid=26587249 |pmc=4632521 |bibcode=2015RSOS....250393D}}</ref> One early giraffid ancestor was ''[[Canthumeryx]]'', which has been dated variously to have lived {{mya|25|20}}, 17–15 mya or 18–14.3 mya and whose deposits have been found in Libya. This animal resembled an [[antelope]] and had a medium-sized, lightly built body. ''[[Giraffokeryx]]'' appeared 15–12 mya on the [[Indian subcontinent]] and resembled an okapi or a small giraffe, and had a longer neck and similar [[ossicone]]s.<ref name=Mitchell20003/> ''Giraffokeryx'' may have shared a clade with more massively built giraffids like ''[[Sivatherium]]'' and ''[[Bramatherium]]''.<ref name=Danowitz2015/> [[File:Giraffidcomparison.jpg|thumb|upright|The extinct giraffid ''[[Samotherium]]'' (middle) in comparison with the [[okapi]] (below) and giraffe. The anatomy of ''Samotherium'' appears to have shown a transition to a giraffe-like neck.<ref name=Danowitz/>]] Giraffids like ''[[Palaeotragus]]'', ''[[Shansitherium]]'' and ''[[Samotherium]]'' appeared 14 mya and lived throughout Africa and Eurasia. These animals had broader skulls with reduced frontal cavities.<ref name=Mitchell20003/><ref name=Danowitz2015/> ''Paleotragus'' resembled the okapi and may have been its ancestor.<ref name=Mitchell20003/> Others find that the okapi lineage diverged earlier, before ''Giraffokeryx''.<ref name=Danowitz2015/> ''Samotherium'' was a particularly important [[transitional fossil]] in the giraffe lineage, as the length and structure of its [[cervical vertebrae]] were between those of a modern giraffe and an okapi, and its neck posture was likely similar to the former's.<ref name=Danowitz>{{cite journal|author1=Danowitz, M. |author2=Domalski, R. |author3=Solounias, N. |year=2015 |title=The cervical anatomy of ''Samotherium'', an intermediate-necked giraffid |journal=Royal Society Open Science |volume=2|issue=11 |page=150521|doi=10.1098/rsos.150521|pmid=26716010 |pmc=4680625 |bibcode=2015RSOS....250521D}}</ref> ''[[Bohlinia]]'', which first appeared in southeastern Europe and lived 9–7 mya, was likely a direct ancestor of the giraffe. ''Bohlinia'' closely resembled modern giraffes, having a long neck and legs and similar ossicones and dentition.<ref name=Mitchell20003/> ''Bohlinia'' colonised China and northern India and produced the ''Giraffa'', which, around {{mya|7}}, reached Africa. [[Climate change]]s led to the extinction of the Asian giraffes, while the African giraffes survived and radiated into new species. Living giraffes appear to have arisen around {{mya|1}} in eastern Africa during the [[Pleistocene]].<ref name=Mitchell20003/> Some biologists suggest the modern giraffes descended from ''[[Giraffa jumae|G. jumae]]'';<ref name=sim1996/> others find ''[[Giraffa gracilis|G. gracilis]]'' a more likely candidate. ''G. jumae'' was larger and more robust, while ''G. gracilis'' was smaller and more slender.<ref name=Mitchell20003/> The changes from extensive forests to more open [[habitat]]s, which began 8 mya, are believed to be the main driver for the evolution of giraffes.<ref name=Mitchell20003/> During this time, tropical plants disappeared and were replaced by arid [[C4 plant]]s, and a dry [[savannah]] emerged across eastern and northern Africa and western India.<ref>{{Cite journal|last=Janis|first=C. M. |date=1993 |title=Tertiary Mammal Evolution in the Context of Changing Climates, Vegetation, and Tectonic Events |journal=[[Annual Review of Ecology and Systematics]] |volume=24|pages=467–500|doi=10.1146/annurev.ecolsys.24.1.467|jstor=2097187}}</ref><ref>{{Cite journal |last1=Ramstein |first1=G. |last2=Fluteau|first2=F. |last3=Besse|first3=J. |last4=Joussaume|first4=S. |date=1997 |title=Effect of orogeny, plate motion and land–sea distribution on Eurasian climate change over the past 30 million years|journal=[[Nature (journal)|Nature]] |volume=386 |issue=6627 |pages=788–795 |doi=10.1038/386788a0 |bibcode=1997Natur.386..788R |s2cid=4335003}}</ref> Some researchers have hypothesised that this new habitat, coupled with a different diet, including acacia species, may have exposed giraffe ancestors to toxins that caused higher mutation rates and a higher rate of evolution.<ref name=bada/> The coat patterns of modern giraffes may also have coincided with these habitat changes. Asian giraffes are hypothesised to have had more okapi-like colourations.<ref name=Mitchell20003/> The giraffe genome is around 2.9 billion [[base pair]]s in length, compared to the 3.3 billion base pairs of the okapi. Of the proteins in giraffe and okapi genes, 19.4% are identical. The [[Divergent evolution|divergence]] of giraffe and okapi lineages dates to around 11.5 mya. A small group of [[regulatory gene]]s in the giraffe appears responsible for the animal's height and associated circulatory adaptations.<ref name=Holmes/><ref name=Genome>{{cite journal|author1=Agaba, M. |author2=Ishengoma, E. |author3=Miller, W. C. |author4=McGrath, B. C. |author5=Hudson, C. N. |author6=Bedoya, R. O. C. |author7=Ratan, A. |author8=Burhans, R. |author9=Chikhi, R. |author10=Medvedev, P. |author11=Praul C. A. |author12=Wu-Cavener, L. |author13=Wood, B. |author14=Robertson, H. |author15=Penfold, L. |author16=Cavener, D. R. |year=2016|title=Giraffe genome sequence reveals clues to its unique morphology and physiology |journal=[[Nature Communications]]|volume=7 |page=11519 |doi=10.1038/ncomms11519 |pmid=27187213 |pmc=4873664 |bibcode=2016NatCo...711519A}}</ref> ===Species and subspecies=== [[File:Genetic subdivision in the giraffe based on mitochondrial DNA sequences.png|thumb|right|alt=Approximate geographic ranges, fur patterns, and phylogenetic relationships between some giraffe subspecies|Map showing "Approximate geographic ranges, fur patterns, and [[phylogenetics|phylogenetic]] relationships between some giraffe subspecies based on [[mitochondrial DNA]] sequences. Colored dots on the map represent sampling localities. The phylogenetic tree is a [[maximum-likelihood]] [[Phylogenetic tree#Special tree types|phylogram]] based on samples from 266 giraffes. Asterisks along branches correspond to [[Clade#Definitions|node]] values of more than 90% [[Resampling (statistics)|bootstrap]] support. Stars at branch tips identify [[paraphyletic]] [[haplotype]]s found in Maasai and reticulated giraffes".<ref name=GeneticStructure/>]] The [[International Union for Conservation of Nature]] (IUCN) currently recognises only one species of giraffe with nine subspecies.<ref name=iucn/> [[Carl Linnaeus]] originally classified living giraffes as one species in 1758. He gave it the [[Binomial nomenclature|binomial]] name ''[[Cervus]] camelopardalis''. [[Mathurin Jacques Brisson]] coined the generic name ''Giraffa'' in 1762.<ref name=Dagg1971>{{Cite journal|author=Dagg, A. I.|author-link=Anne Innis Dagg |year=1971 |title=''Giraffa camelopardalis''|journal=[[Mammalian Species]]|pages=1–8 |issue=5 |jstor=3503830 |url=http://www.science.smith.edu/msi/pdf/i0076-3519-005-01-0001.pdf |doi=10.2307/3503830 |access-date=25 October 2011 |archive-date=21 April 2017|archive-url=https://web.archive.org/web/20170421052751/http://www.science.smith.edu/msi/pdf/i0076-3519-005-01-0001.pdf |url-status=dead}}</ref> During the 1900s, various taxonomies with two or three species were proposed.<ref>{{Cite journal |last=Gippoliti |first=S. |date=2018|title=Impacts of taxonomic inertia for the conservation of African ungulate diversity: an overview |url=https://onlinelibrary.wiley.com/doi/abs/10.1111/brv.12335 |journal=[[Biological Reviews]]|volume=93|issue=1|pages=115–130 |doi=10.1111/brv.12335 |pmid=28429851 |s2cid=5189968|access-date=21 April 2021|archive-date=3 April 2021|archive-url=https://web.archive.org/web/20210403050548/https://onlinelibrary.wiley.com/doi/abs/10.1111/brv.12335|url-status=live}}</ref> A 2007 study on the genetics of giraffes using mitochondrial DNA suggested at least six lineages could be recognised as species.<ref name=GeneticStructure>{{cite journal|title=Extensive population genetic structure in the giraffe|author1=Brown, D. M. |author2=Brenneman R. A. |author3=Koepfli, K.-P. |author4=Pollinger, J. P. |author5=Milá, B. |author6=Georgiadis, N. J. |author7=Louis Jr., E. E. |author8=Grether, G. F. |author9=Jacobs, D. K. |author10=Wayne R. K. |journal= [[BMC Biology]] |year=2007 |volume=5 |issue=1 |page=57 |doi=10.1186/1741-7007-5-57 |pmc=2254591|pmid=18154651 |doi-access=free}}</ref> A 2011 study using detailed analyses of the morphology of giraffes, and application of the [[Species|phylogenetic species concept]], described eight species of living giraffes.<ref name=":2">{{Cite book|url=https://books.google.com/books?id=v3uZtA1ZpTAC&q=ungulate+taxonomy&pg=PP2|title=Ungulate Taxonomy |last1=Groves |first1=C. |last2=Grubb |first2=P. |year=2011 |pages=68–70|publisher=[[JHU Press]] |isbn=9781421400938 |access-date=17 October 2020|archive-date=22 September 2023 |archive-url=https://web.archive.org/web/20230922073613/https://books.google.com/books?id=v3uZtA1ZpTAC&q=ungulate+taxonomy&pg=PP2 |url-status=live}}</ref> A 2016 study also concluded that living giraffes consist of multiple species. The researchers suggested the existence of four species, which have not exchanged genetic information between each other for one to two million years.<ref name="cur.biol 2016">{{cite journal |last1=Fennessy |first1=J. |last2=Bidon |first2=T. |last3=Reuss |first3=F. |last4=Kumar |first4=V. |last5=Elkan |first5=P. |last6=Nilsson |first6=M. A. |last7=Vamberger|first7=M. |last8=Fritz|first8=U. |last9=Janke |first9=A. |year=2016 |title=Multi-locus Analyses reveal four giraffe species instead of one |journal=[[Current Biology]] |volume=26 |issue=18 |pages=2543–2549 |doi=10.1016/j.cub.2016.07.036 |pmid=27618261 |s2cid=3991170 |doi-access=free|bibcode=2016CBio...26.2543F }}</ref> A 2020 study showed that depending on the method chosen, different taxonomic hypotheses recognizing from two to six species can be considered for the genus ''Giraffa''. That study also found that multi-species coalescent methods can lead to taxonomic over-splitting, as those methods delimit geographic structures rather than species. The three-species hypothesis, which recognises ''G''. ''camelopardalis'', ''G''. ''giraffa'', and ''G''. ''tippelskirchi'', is highly supported by [[Phylogenetics|phylogenetic]] analyses and also corroborated by most population genetic and multi-species coalescent analyses.<ref name=":6">{{Cite journal |last1=Petzold|first1=A. |last2=Hassanin |first2=A. |date=2020 |title=A comparative approach for species delimitation based on multiple methods of multi-locus DNA sequence analysis: A case study of the genus ''Giraffa'' (Mammalia, Cetartiodactyla) |journal=[[PLOS One]] |volume=15 |issue=2 |pages=e0217956 |doi=10.1371/journal.pone.0217956 |pmid=32053589 |pmc=7018015 |bibcode=2020PLoSO..1517956P |doi-access=free}}</ref> A 2021 [[whole genome sequencing]] study suggests the existence of four distinct species and seven subspecies,<ref name=Coimbra2021>{{cite journal|last1=Coimbra|first1=R.T.F. |last2=Winter|first2=S. |last3=Kumar |first3=V. |last4=Koepfli|first4=K.-P. |last5=Gooley|first5=R. M. |last6=Dobrynin |first6=P. |last7=Fennessy|first7=J. |last8=Janke|first8=A. |title=Whole-genome analysis of giraffe supports four distinct species |journal=Current Biology |volume=31 |issue=13 |year=2021 |pages=2929–2938.e5 |doi=10.1016/j.cub.2021.04.033 |pmid=33957077 |s2cid=|doi-access=free|bibcode=2021CBio...31E2929C }}</ref> which was supported by a 2024 study of cranial morphology.<ref>{{cite journal |last1=Kargopoulos |first1=Nikolaos |last2=Marugán-Lobón |first2=Jesús |last3=Chinsamy |first3=Anusuya |last4=Agwanda |first4=Bernard R. |last5=Brown |first5=Michael Butler |last6=Fennessy |first6=Stephanie |last7=Ferguson |first7=Sara |last8=Hoffman |first8=Rigardt |last9=Lala |first9=Fredrick |last10=Muneza |first10=Arthur |last11=Mwebi |first11=Ogeto |last12=Otiende |first12=Moses |last13=Petzold |first13=Alice |last14=Winter |first14=Sven |last15=Zabeirou |first15=Abdoul Razack Moussa |last16=Fennessy |first16=Julian |title=Heads up–Four Giraffa species have distinct cranial morphology |journal=PLOS ONE |date=19 December 2024 |volume=19 |issue=12 |pages=e0315043 |doi=10.1371/journal.pone.0315043 |doi-access=free |pmid=39700177 |issn=1932-6203 |pmc=11658530}}</ref> A 2024 study found a higher amount of ancient [[gene flow]] than expected between populations.<ref name=Bertola/> The [[cladogram]] below shows the phylogenetic relationship between the four proposed species and seven subspecies based on a 2021 genome analysis.<ref name=Coimbra2021/> The eight lineages correspond to eight traditional subspecies in the one-species hypothesis. The Rothschild giraffe is subsumed into ''G. camelopardalis camelopardalis''. {{clade gallery |align=center |cladogram1= {{clade |label1='''''Giraffa''''' |1={{clade |1={{clade |label1=''Giraffa camelopardalis'' |sublabel1=(northern giraffe) |1={{clade |1={{clade |1=''G. camelopardalis antiquorum'' (Kordofan giraffe) |2=''G. c. camelopardalis'' (Nubian giraffe) }} |2={{clade |1=''G. c. peralta'' (West African giraffe) }} }} |label2=''G. reticulata'' |sublabel2=(reticulated giraffe) |2={{clade |state=none |1={{clade |state=none |1=(no subspecies) }}}} }} |2={{clade |label1=''G. tippelskirchi'' |sublabel1=(Masai giraffe ''sensu lato'') |1={{clade |1=''G. tippelskirchi tippelskirchi'' (Masai giraffe ''sensu stricto'') |2=''G. t. thornicrofti'' (Luangwa or Thornicroft's giraffe) }} |label2=''G. giraffa'' |sublabel2=(southern giraffe) |2={{clade |1=''G. giraffa angolensis'' (Angolan giraffe) |2=''G. g. giraffa'' (South African giraffe) }} }} }} }} }} The following table compares the different hypotheses for giraffe species. The description column shows the traditional nine subspecies in the one-species hypothesis.<ref name=iucn/><ref name=":1">{{Cite book|title=Systema Naturæ|last=Linnaeus|first=C. |year=1758}}</ref> {| class="wikitable" |+ Species and subspecies of giraffe ! Description ! Image ! Eight species taxonomy<ref name=":2" /> ! Four species taxonomy<ref name="cur.biol 2016" /><ref name="Coimbra2021"/> ! Three species taxonomy<ref name=":6" /> |- | The '''[[Kordofan giraffe]]''' (''G. c. antiquorum'') has a distribution which includes southern [[Chad]], the [[Central African Republic]], northern [[Cameroon]], and the northeastern [[Democratic Republic of the Congo]].<ref name=iucn/> Populations in Cameroon were formerly included in ''G. c. peralta'', but this was incorrect.<ref name=WestAfricaGiraffe>{{Cite journal|author1=Hassanin, A. |author2=Ropiquet, A. |author3=Gourmand, B.-L. |author4=Chardonnet, B. |author5=Rigoulet, J. |year=2007 |title=Mitochondrial DNA variability in ''Giraffa camelopardalis'': consequences for taxonomy, phylogeography and conservation of giraffes in West and Central Africa |journal=[[Comptes Rendus Biologies]] |volume=330 |issue=3 |pages=173–183 |pmid=17434121|doi=10.1016/j.crvi.2007.02.008}}</ref> Compared to the Nubian giraffe, this subspecies has smaller and more irregular spotting patterns. Its spots are present on insides of the legs, sometimes below the [[Hock (anatomy)|hock]]s. A median lump is present in males.<ref name=Seymour/>{{rp|51–52}} Some 2,000 are believed to remain in the wild.<ref name=iucn/> Considerable confusion has existed over the status of this subspecies and ''G. c. peralta'' in zoos. In 2007, all alleged ''G. c. peralta'' in [[List of zoos#Europe|European zoos]] were shown to be, in fact, ''G. c. antiquorum''.<ref name=WestAfricaGiraffe/> With this correction, about 65 are living in zoos.<ref name=ISIS/> |[[File:Zoo de Vincennes, Paris, France April 2014 (7), crop.jpg|150px]] |'''[[Kordofan giraffe]]''' <br />(''G. antiquorum'')<ref>Swaison 1835. Camelopardalis antiquorum. Bagger el Homer, Kordofan, about 10° N, 28° E (as fixed by Harper, 1940)</ref> | rowspan="4" colspan="2" style="border-bottom:0px none| '''[[Northern giraffe]]''' <br />(''G. camelopardalis'') {{smalldiv|1= Three or four subspecies: * ''G. c. antiquorum'' * ''G. c. camelopardalis'' * ''G. c. peralta'' * ''G. c. reticulata'' (only in three-species hypothesis) }} |- | The '''[[Nubian giraffe]]''' (''G. c. camelopardalis''), is found in eastern [[South Sudan]] and southwestern [[Ethiopia]], in addition to [[Kenya]] and [[Uganda]].<ref name=iucn/> It has sharp-edged [[chestnut (color)|chestnut]]-coloured spots surrounded by mostly white lines, while undersides lack spotting. A lump is prominent in the middle of the male's head.<ref name=Seymour>{{cite thesis |author=Seymour, R. |year=2001 |title=Patterns of subspecies diversity in the giraffe, Giraffa camelopardalis (L. 1758): comparison of systematic methods and their implications for conservation policy |type=Ph.D. thesis|url=https://kar.kent.ac.uk/86053/}}</ref>{{rp|51}} Around 2,150 are thought to remain in the wild, with another 1,500 individuals belonging to the [[Rothschild's giraffe|Rothschild's]] [[ecotype]].<ref name=iucn/> With the addition of Rothschild's giraffe to the Nubian subspecies, the Nubian giraffe is very common in captivity, although the original phenotype is rare — a group is kept at [[Al Ain Zoo]] in the [[United Arab Emirates]].<ref name="Al Ain exhibits">{{cite web |title=Exhibits|url=http://www.awpr.ae/en/Visit/Pages/AfricanMixedExhibit.aspx|archive-url=https://web.archive.org/web/20111129023015/http://www.awpr.ae/en/Visit/Pages/AfricanMixedExhibit.aspx|archive-date=29 November 2011 |publisher=[[Al Ain Zoo]] |date=25 February 2003|access-date=21 November 2011}}</ref> In 2003, this group numbered 14.<ref>{{cite web |url=http://www.uaeinteract.com/docs/Nubian_giraffe_born_in_Al_Ain_zoo/6971.htm |title=Nubian giraffe born in Al Ain zoo |publisher=UAE Interact |access-date=21 December 2010|url-status=dead |archive-url=https://web.archive.org/web/20120320090832/http://www.uaeinteract.com/docs/Nubian_giraffe_born_in_Al_Ain_zoo/6971.htm|archive-date=20 March 2012}}</ref> |[[File:Giraffa camelopardalis camelopardalis (Al Ain Zoo, UAE), crop & flip.jpg|150px]] | rowspan="2" |'''[[Nubian giraffe]]''' <br />(''G. camelopardalis'')<ref name=":1" /> Also known as '''[[Lake Baringo|Baringo]] giraffe''' or '''[[Uganda]]n giraffe''' {{smalldiv|1= Two subspecies: * ''G. c. camelopardalis'' * ''G. c. rothschildi'' (Rothschild's giraffe) }} |- |'''[[Rothschild's giraffe]]''' (''G. c. rothschildi'') may be an ecotype of ''G. camelopardalis''. Its range includes parts of Uganda and [[Kenya]].<ref name=iucn/> Its presence in South Sudan is uncertain.<ref name=IUCNrothschildi>{{cite iucn |author=Fennessy, S. |author2=Fennessy, J. |author3=Muller, Z. |author4=Brown, M. |author5=Marais, A. |date=2018 |title=''Giraffa camelopardalis'' ssp. ''rothschildi'' |volume=2018 |page=e.T174469A51140829 |doi=10.2305/IUCN.UK.2018-2.RLTS.T174469A51140829.en |access-date=19 November 2021}}</ref> This giraffe has large dark patches with normally well-defined edges but sometimes split. The dark spots may also have swirls of pale colour within them. Spotting rarely reaches below the hocks and rarely to the hooves. This ecotype may also develop five "horns".<ref name=Seymour/>{{rp|53}} Around 1,500 individuals are believed to remain in the wild,<ref name=iucn/> and more than 450 are living in zoos.<ref name=ISIS/> According to genetic analysis circa September 2016, it is [[Biological specificity|conspecific]] with the [[Nubian giraffe]] (''G. c. camelopardalis'').<ref name="cur.biol 2016" /> |[[File:Rothschild's Giraffe (Giraffa camelopardalis rothschildi) male (7068054987), crop & edit.jpg|150px]] |- | The '''[[West African giraffe]]''' (''G. c. peralta'') is [[endemic]] to southwestern Niger.<ref name=iucn/> This animal has a lighter [[pelage]] (fur) than other subspecies,<ref name=Kingdon1988>{{Cite book|author=Kingdon, J. |year=1988|title=East African Mammals: An Atlas of Evolution in Africa |volume=((3, Part B: Large Mammals)) |pages=313–337 |publisher=[[University of Chicago Press]] |isbn=978-0-226-43722-4}}</ref>{{rp|322}} with red lobe-shaped blotches that reach under the hocks. The ossicones are more erect than in other subspecies, and males have well-developed median lumps.<ref name=Seymour/>{{rp|52–53}} It is the most endangered subspecies within ''Giraffa'', with 400 individuals remaining in the wild.<ref name=iucn/> Giraffes in Cameroon were formerly believed to belong to this species, but are actually ''G. c. antiquorum''. This error resulted in some confusion over its status in zoos, but in 2007 it was established that all "''G. c. peralta''" kept in European zoos are actually ''G. c. antiquorum''. The same 2007 study found that the West African giraffe was more closely related to Rothschild's giraffe than the Kordofan, and its ancestor may have migrated from eastern to northern Africa and then west as the [[Sahara]] Desert spread. At its largest, [[Lake Chad]] may have acted as a boundary between the West African and Kordofan giraffes during the [[Holocene]] (before 5000 BC).<ref name=WestAfricaGiraffe/> |[[File:Giraffe-solo Koure-NIGER.jpg|150px]] |'''[[West African giraffe]]''' <br />(''G. peralta''),<ref name=IUCNperalta>{{cite iucn|author=Fennessy, J. |author2=Marais, A. |author3=Tutchings, A. |year=2018 |title=''Giraffa camelopardalis'' ssp. ''peralta'' |page=e.T136913A51140803}}</ref> Also known as '''[[Niger]] giraffe''' or '''[[Nigeria]]n giraffe''' |- | The '''[[reticulated giraffe]]''' (''G. c. reticulata'') is native to northeastern Kenya, southern Ethiopia, and [[Somalia]].<ref name=iucn/> Its distinctive coat pattern consists of sharp-edged, reddish-brown polygonal patches surrounded by thin white lines. Spots may or may not extend under the hocks, and a median lump is present in males.<ref name=Seymour/>{{rp|53}} An estimated 8,660 individuals remain in the wild,<ref name=iucn/> and based on [[International Species Information System]] records, more than 450 are living in zoos.<ref name=ISIS>{{cite web|url=https://app.isis.org/abstracts/Abs77545.asp |title=Giraffa |publisher=[[International Species Information System|ISIS]] |year=2010|access-date= 4 November 2010|archive-url=https://web.archive.org/web/20100706081917/https://app.isis.org/abstracts/Abs77545.asp |archive-date=6 July 2010 |url-status=dead}}</ref> A 2024 study found that the reticulated giraffe is the result of [[Hybrid (biology)|hybridisation]] between northern and southern giraffe lineages.<ref name=Bertola>{{cite journal|last1=Bertola|first1=L. D.|last2=Quinn|first2=L|last3=Hanghøj|first3=K|last4=Garcia-Erill|first4=E|last5=Rasmussen|first5=S. M.|last6=Meisner|first6=J|last7=Bøggild|first7=T|last8=Wang|first8=X|last9=Lin|first9=L|last10=Nursyifa|first10=C|last11=Liu|first11=X|last12=Li|first12=Z|last13=Chege|first13=M|last14=Moodley|first14=Y|last15=Brüniche-Olsen|first15=A|last16=Kuja|first16=J|last17=Schubert|first17=M|last18=Agaba|first18=M|last19=Santander|first19=C. G.|last20=Sinding|first20=M-H. S.|last21=Muwanika|first21=V|last22=Masembe|first22=C|last23=Siegismund|first23=H. R.|last24=Moltke|first24=I|last25=Albrechtsen|first25=A|last26=Heller|first26=R.|year=2024|title=Giraffe lineages are shaped by major ancient admixture events|journal=Current Biology|volume=34 |issue=7 |pages=1576–1586.e5 |doi=10.1016/j.cub.2024.02.051|doi-access=free|pmid=38479386 |bibcode=2024CBio...34E1576B }}</ref> |[[File:Giraffa camelopardalis reticulata 01, flip.jpg|150px]] | colspan="2" | '''[[Reticulated giraffe]]''' <br />(''G. reticulata''),<ref>{{Cite journal |last=Winton|first=W. E. de|date=1899 |title=XXXVIII.—On mammals collected by Lieut.-Colonel W. Giffard in the northern territory of the Gold Coast |journal=[[Annals and Magazine of Natural History]] |volume=4 |issue=2 3|pages=353–359 |doi=10.1080/00222939908678212|issn=0374-5481 |url=https://zenodo.org/record/1430367|access-date=16 September 2019|archive-date=1 August 2020|archive-url=https://web.archive.org/web/20200801031317/https://zenodo.org/record/1430367|url-status=live}}</ref> Also known as '''Somali giraffe''' |style="border-top:0px none;" | <!-- empty cell joined to northern giraffe by hiding border --> |- | The '''[[Angolan giraffe]]''' (''G. c. angolensis'')<ref name=MacDonald/> occurs in northern [[Namibia]], southwestern [[Zambia]], central [[Botswana]], western [[Zimbabwe]], southern Zimbabwe and, since mid-2023, again in Angola.<ref name="2023-07-11_NPR">{{cite web |date=2023 |url=https://www.npr.org/2023/07/11/1187035189/for-the-first-time-in-decades-angolan-giraffes-now-populate-a-park-in-angola |title=For the first time in decades, Angolan giraffes now populate a park in Angola |website=[[NPR]] |access-date=12 July 2023 |archive-date=12 July 2023 |archive-url=https://web.archive.org/web/20230712085939/https://www.npr.org/2023/07/11/1187035189/for-the-first-time-in-decades-angolan-giraffes-now-populate-a-park-in-angola |url-status=dead }}, Megan Lim, [[NPR]], 11 July 2023</ref><ref name=iucn/><ref name=":3">{{Cite journal |last1=Bock |first1=F. |last2=Fennessy |first2=J. |last3=Bidon |first3=T. |last4=Tutchings |first4=A. |last5=Marais |first5=A. |last6=Deacon |first6=F. |last7=Janke |first7=A. |date=2014 |title=Mitochondrial sequences reveal a clear separation between Angolan and South African giraffe along a cryptic rift valley |journal=BMC Evolutionary Biology |volume=14 |issue=1 |page=219 |doi=10.1186/s12862-014-0219-7 |issn=1471-2148 |pmc=4207324 |pmid=25927851 |doi-access=free|bibcode=2014BMCEE..14..219B }}</ref><ref name=":10">{{Cite journal |last1=Winter |first1=S. |last2=Fennessy |first2=J. |last3=Fennessy |first3=S. |last4=Janke |first4=A. |date=2018 |title=Matrilineal population structure and distribution of the Angolan giraffe in the Namib desert and beyond |journal=Ecological Genetics and Genomics |url=https://www.sciencedirect.com/science/article/pii/S2405985417300344 |volume=7-8 |pages=1–5 |doi=10.1016/j.egg.2018.03.003 |bibcode=2018EcoGG...7....1W |s2cid=90395544}}</ref> A 2009 genetic study on this subspecies suggested the northern [[Namib Desert]] and [[Etosha National Park]] populations form a separate subspecies.<ref>{{cite journal |doi=10.1111/j.1365-2028.2009.01078.x |author1=Brenneman, R. A. |author2=Louis, E. E. Jr. |author3=Fennessy, J. |year=2009 |title=Genetic structure of two populations of the Namibian giraffe, ''Giraffa camelopardalis angolensis''|journal=[[African Journal of Ecology]] |volume=47 |issue=4|pages=720–728|bibcode=2009AfJEc..47..720B }}</ref> This subspecies is white with large brown blotches with pointed or cut edges. The spotting pattern extends throughout the legs but not the upper part of the face. The neck and rump patches tend to be fairly small. The subspecies also has a white ear mark.<ref name=Seymour/>{{rp|51}} About 13,000 animals are estimated to remain in the wild,<ref name=iucn/> and about 20 are living in zoos.<ref name=ISIS/> |[[File:Giraffa camelopardalis angolensis, flip.jpg|150px]] |'''[[Angolan giraffe]]''' <br />(''G. angolensis'') Also known as '''Namibian giraffe''' | rowspan="2" colspan="2" |'''[[Southern giraffe]]''' (''G. giraffa'') {{smalldiv|1= Two subspecies: * ''G. g. angolensis'' * ''G. g. giraffa'' }} |- | The '''[[South African giraffe]]''' (''G. c. giraffa'') is found in northern [[South Africa]], southern Botswana, northern Botswana and southwestern [[Mozambique]].<ref name=iucn/><ref name=":3" /><ref name=":10" /> It has a tawny background colour marked with dark, somewhat rounded patches "with some fine projections". The spots extend down the legs, growing smaller as they do. The median lump of males is relatively small.<ref name=Seymour/>{{rp|52}} A maximum of 31,500 are estimated to remain in the wild,<ref name=iucn/> and around 45 are living in zoos.<ref name=ISIS/> |[[File:Giraffe standing.jpg|150px]] |'''[[South African giraffe]]''' <br />(''G. giraffa'')<ref>{{Cite book |url=https://books.google.com/books?id=IdyuQaviYjIC&q=Rookmaaker,+1989&pg=PR9 |title=The Zoological Exploration of Southern Africa 1650–1790 |last=Rookmaaker |first=L. C. |date=1989 |publisher=[[CRC Press]] |isbn=9789061918677 |access-date=17 October 2020|archive-date=22 September 2023 |archive-url=https://web.archive.org/web/20230922073614/https://books.google.com/books?id=IdyuQaviYjIC&q=Rookmaaker,+1989&pg=PR9 |url-status=live}}</ref> Also known as '''Cape giraffe''' |- | The '''[[Masai giraffe]]''' (''G. c. tippelskirchi'') can be found in central and southern Kenya and in [[Tanzania]].<ref name=iucn/> Its coat patterns are highly diverse, with spots ranging from mostly rounded and smooth-edged to oval-shaped and incised or loped-edged.<ref name="Lee2018"/> A median lump is usually present in males.<ref name=Seymour/>{{rp|54}}<ref name=estes/> A total of 32,550 are thought to remain in the wild,<ref name=iucn/> and about 100 are living in zoos.<ref name=ISIS/> |[[File:GiraffaCamelopardalisTippelskirchi-Masaai-Mara.JPG|150px]] |'''[[Masai giraffe]]''' <br />(''G. tippelskirchi'')<ref name="MacDonald"/> Also known as '''[[Kilimanjaro]] giraffe''' | rowspan="2" colspan="2" |'''[[Masai giraffe]]''' (''G. tippelskirchi'') {{smalldiv|1= Two subspecies: * ''G. t. tippelskirchi'' * ''G. t. thornicrofti'' }} |- | [[Rhodesian giraffe|'''Thornicroft's giraffe''']] (''G. c. thornicrofti'') is restricted to the [[Luangwa River|Luangwa Valley]] in eastern Zambia.<ref name=iucn/> It has notched and somewhat [[Star polygon|star-shaped]] patches which and may or may not extend across the legs. The median lump of males is modestly sized.<ref name=Seymour/>{{rp|54}} No more than 550 remain in the wild,<ref name=iucn/> with none in zoos.<ref name=ISIS/> It was named after [[Harry Scott Thornicroft]].<ref name=MacDonald>{{Cite book|title=The Encyclopedia of Mammals |publisher=[[Oxford University Press]] |year=2001|isbn=978-0-7607-1969-5|edition=2nd|pages=520–527 |author=Pellow, R. A. |contribution=Giraffe and Okapi |editor=MacDonald, D.}}</ref> |[[File:Giraffe Walking Square, flip.jpg|150px]] |'''[[Rhodesian giraffe|Thornicroft's giraffe]]''' <br />("''G. thornicrofti''") Also known as '''Luangwa giraffe''' or '''Rhodesian giraffe''' |} The first extinct species to be described was ''[[Giraffa sivalensis]]'' from Pakistan, the holotype of which was reevaluated as a vertebra of separate species within the genus that was initially described as a fossil of the living giraffe.<ref name=Sittert2015>{{cite journal |last1=van Sittert |first1=S. J. |last2=Mitchell |first2=G. |title=On reconstructing ''Giraffa sivalensis'', an extinct giraffid from the Siwalik Hills, India |journal=PeerJ |date=2015 |volume=3 |page=e1135 |doi=10.7717/peerj.1135 |pmid=26290791 |pmc=4540016 |doi-access=free}}</ref> Another extinct species ''Giraffa punjabiensis'' is known from Pakistan.<ref>{{cite journal|last1=Barry|first1=John C.|last2=Morgan|first2=Michèle E.|last3=Flynn|first3=Lawrence J.|last4=Pilbeam|first4=David|last5=Behrensmeyer|first5=Anna K.|last6=Raza|first6=S. Mahmood|last7= A. Khan|first7=Imran|last8=Badgley|first8=Catherine|last9=Hicks|first9=Jason|last10=Kelley|first10=Jay|year=2002|title=Faunal and Environmental Change in the Late Miocene Siwaliks of Northern Pakistan|journal=Paleobiology|volume=28|issue=2|pages=1–71|doi=10.1666/0094-8373(2002)28[1:FAECIT]2.0.CO;2|s2cid=18408360 }}</ref> Four other valid extinct species of ''Giraffa'' known from Africa are ''Giraffa gracilis'', ''[[Giraffa jumae]]'', ''[[Giraffa pygmaea]]'' and ''[[Giraffa stillei]]''.<ref name="Danowitz2015"/> "''G.''" ''pomeli'' from Algeria and Tunisia is not a species of Giraffinae, but a species of Palaeotraginae related to ''Mitilanotherium''.<ref>{{cite book|author1=Harris, J.M.|author2=Solounias, N.|author3=Geraads, D.|year=2010|chapter=Giraffoidea|title=Cenozoic Mammals of Africa|editor1=Werdelin, L.|editor2=Sanders, W.J.|publisher=University of California Press|location=Berkeley, Los Angeles & London|pages=797-811|isbn=9780520257214}}</ref> ==Anatomy== [[File:Giraffe skeleton.jpg|thumb|upright|alt=Photograph of a Giraffe skeleton|Giraffe skeleton on display at the [[Museum of Osteology]], [[Oklahoma City]]]] Fully grown giraffes stand {{cvt|4.3|–|5.7|m}} tall, with [[sexual dimorphism|males taller]] than females.<ref name=Nowak1999>{{cite book|author=Nowak, R.M. |date=1999 |url=https://books.google.com/books?id=7W-DGRILSBoC&pg=PA1086 |pages=1086–1089|title=Walker's Mammals of the World |volume=1 |publisher=The [[Johns Hopkins University Press]] |isbn=978-0801857898|access-date=30 August 2021|archive-date=22 September 2023|archive-url=https://web.archive.org/web/20230922073614/https://books.google.com/books?id=7W-DGRILSBoC&pg=PA1086|url-status=live}}</ref> The average weight is {{cvt|1192|kg}} for an adult male and {{cvt|828|kg}} for an adult female.<ref name=Skinner1990>{{Cite book|author1=Skinner, J. D. |author2=Smithers, R. H. M. |year=1990|title=The mammals of the southern African subregion |pages=616–20|publisher=[[University of Pretoria]]|isbn=978-0-521-84418-5}}</ref> Despite its long neck and legs, its body is relatively short.<ref name=anatomy/>{{rp|66}} The skin is mostly gray<ref name=Skinner1990/> or tan,<ref>{{cite web |author=Langley, L. |date=2017 |title=Do zebras have stripes on their skin?|publisher=[[National Geographic]]|access-date=2 June 2020 |url=https://www.nationalgeographic.com/news/2017/03/animals-skin-colors-zebras-big-cats/|archive-date=1 April 2019|archive-url=https://web.archive.org/web/20190401134219/https://news.nationalgeographic.com/2017/03/animals-skin-colors-zebras-big-cats/|url-status=dead}}</ref> and can reach a thickness of {{cvt|20|mm}}.<ref name=Dagg2014/>{{rp|87}} The {{cvt|80|–|100|cm}} long<ref name=MacDonald/> tail ends in a long, dark tuft of hair and is used as a defense against insects.<ref name=Dagg2014/>{{rp|94}} The [[Animal coat|coat]] has dark blotches or patches, which can be orange, [[Chestnut (color)|chestnut]], brown, or nearly black, surrounded by light hair, usually white or [[Cream (colour)|cream]] coloured.<ref name="Prothero 2003"/> Male giraffes become darker as they grow old.<ref name=estes>{{Cite book|title=The Behavior Guide to African Mammals: including Hoofed Mammals, Carnivores, Primates |author=Estes, R. |publisher=[[University of California Press]] |pages=202–207 |year=1992 |isbn=978-0-520-08085-0 |url=https://archive.org/details/isbn_0520080858/page/202}}</ref> The coat pattern [[Disruptive coloration#A disputed case: the giraffe|has been claimed]] to serve as [[camouflage]] in the light and shade patterns of savannah woodlands.<ref name=MacDonald/> When standing among trees and bushes, they are hard to see at even a few metres distance. However, adult giraffes move about to gain the best view of an approaching predator, relying on their size and ability to defend themselves rather than on camouflage, which may be more important for calves.<ref name=Mitchell20003/> Each giraffe has a unique coat pattern.<ref>{{Cite journal |last=Foster |first=J. B. |title=The Giraffe of Nairobi National Park: Home range, sex ratios, the herd, and food |date=1966 |journal=African Journal of Ecology |volume=4 |issue=1 |pages=139–148 |doi=10.1111/j.1365-2028.1966.tb00889.x |bibcode=1966AfJEc...4..139F |archive-date=14 December 2022 |url-status=live |url=https://onlinelibrary.wiley.com/doi/10.1111/j.1365-2028.1966.tb00889.x |archive-url=https://web.archive.org/web/20221214102948/https://onlinelibrary.wiley.com/doi/10.1111/j.1365-2028.1966.tb00889.x}}</ref><ref>{{Cite journal |last1=Lee |first1=D. E. |last2=Lohay |first2=G. G. |last3=Cavener |first3=D. R. |last4=Bond |first4=M. L. |date=2022 |title=Using spot pattern recognition to examine population biology, evolutionary ecology, sociality, and movements of giraffes: a 70-year retrospective |url=https://doi.org/10.1007/s42991-022-00261-3 |journal=[[Mammalian Biology]] |volume=102 |issue=4 |pages=1055–1071 |doi=10.1007/s42991-022-00261-3 |s2cid=252149865 |access-date=14 December 2022 |archive-date=22 September 2023 |archive-url=https://web.archive.org/web/20230922073617/https://link.springer.com/article/10.1007/s42991-022-00261-3 |url-status=live}}</ref> Calves inherit some coat pattern traits from their mothers, and variation in some spot traits is correlated with calf survival.<ref name=Lee2018>{{Cite journal|last1=Lee|first1=D. E.|last2=Cavener|first2=D. R.|last3=Bond|first3=M. L.|date=2018 |title=Seeing spots: quantifying mother-offspring similarity and assessing fitness consequences of coat pattern traits in a wild population of giraffes (''Giraffa camelopardalis'') |journal=[[PeerJ]] |volume=6 |pages=e5690|doi=10.7717/peerj.5690|pmid=30310743|pmc=6173159 |doi-access=free}}</ref> The skin under the blotches may regulate the animal's body temperature, being sites for complex blood vessel systems and large [[sweat gland]]s.<ref name=Mitchell2004>{{Cite journal |author1=Mitchell, G. |author2=Skinner, J.D. |year=2004 |title=Giraffe thermoregulation: a review |journal=[[Transactions of the Royal Society of South Africa]] |volume=59 |issue=2 |pages=49–57 |url=http://www.sabinet.co.za/abstracts/royalsa/royalsa_v59_n2_a13.html |doi=10.1080/00359190409519170 |bibcode=2004TRSSA..59..109M |s2cid=87321176 |access-date=19 October 2011 |archive-date=20 November 2018 |archive-url=https://web.archive.org/web/20181120033724/https://www.sabinet.co.za/abstracts/royalsa/royalsa_v59_n2_a13.html |url-status=live}}</ref> Spotless or solid-color giraffes are very rare, but have been observed.<ref name="National Geographic 12 September 2023">{{cite news |last1=Fine Maron |first1=Dina |title=Another rare spotless giraffe found—the first ever seen in the wild |url=https://www.nationalgeographic.com/animals/article/spotless-giraffe-found-in-the-wild-for-the-first-time |access-date=6 March 2024 |work=[[National Geographic]] |date=12 September 2023 |language=en}}</ref><ref name="NPR 6 September 2023">{{cite news |last1=Romo |first1=Vanessa |last2=Jones |first2=Dustin |title=A rare spotless giraffe gets a name to match |url=https://www.npr.org/2023/09/06/1197789671/a-rare-spotless-giraffe-gets-a-name-to-match |access-date=6 March 2024 |work=[[NPR]] |date=6 September 2023}}</ref> The fur may give the animal chemical defense, as its parasite repellents give it a characteristic scent. At least 11 main [[aromaticity|aromatic]] chemicals are in the fur, although [[indole]] and [[3-methylindole]] are responsible for most of the smell. Because males have a stronger odour than females, it may also have a [[Animal sexual behaviour|sexual function]].<ref>{{Cite journal|author1=Wood, W. F. |author2=Weldon, P. J. |year=2002 |title=The scent of the reticulated giraffe (''Giraffa camelopardalis reticulata'') |journal=[[Biochemical Systematics and Ecology]] |volume=30 |issue=10|pages=913–17 |doi=10.1016/S0305-1978(02)00037-6|bibcode=2002BioSE..30..913W }}</ref> ===Head=== [[File:Giraffa camelopardalis head (Profil).jpg|thumb|left|alt=Closeup photograph of a giraffe head|Closeup of the head of a [[northern giraffe]]]] Both sexes have prominent horn-like structures called [[ossicone]]s, which can reach {{cvt|13.5|cm}}. They are formed from [[Ossification|ossified]] [[cartilage]], covered in skin, and fused to the [[skull]] at the [[parietal bone]]s.<ref name=estes/><ref name=Dagg2014/>{{rp|95–97}} Being [[Blood vessel|vascular]]ised, the ossicones may have a role in [[thermoregulation]],<ref name=Mitchell2004/> and are used in combat between males.<ref name="sim2010"/> Appearance is a reliable guide to the sex or age of a giraffe: the ossicones of females and young are thin and display tufts of hair on top, whereas those of adult males tend to be bald and knobbed on top.<ref name=estes/> A lump, which is more prominent in males, emerges in the middle of the skull.<ref name=Dagg1971/> Males develop [[calcium]] deposits that form bumps on their skulls as they age.<ref name="Prothero 2003"/> Multiple [[Sinus (anatomy)|sinuses]] lighten a giraffe's skull.<ref name=Dagg2014/>{{rp|103}} However, as males age, their skulls become heavier and more club-like, helping them become more dominant in combat.<ref name=estes/> The [[occipital condyles]] at the bottom of the skull allow the animal to tip its head over 90 degrees and grab food on the branches directly above them with the tongue.<ref name=Dagg2014/>{{rp|103, 110}}<ref name=Dagg1971/> With eyes located on the sides of the head, the giraffe has a broad [[Field of view|visual field]] from its great height.<ref name=Dagg2014>{{cite book|author=Dagg, A. I.|year=2014|title=Giraffe: Biology, Behaviour, and Conservation|publisher=[[Cambridge University Press]] |isbn=978-1107610170}}</ref>{{rp|85, 102}} Compared to other [[ungulate]]s, giraffe vision is more [[binocular vision|binocular]] and the eyes are larger with a greater [[retina]]l surface area.<ref>{{cite journal|last1=Mitchell|first1=G.|last2=Roberts|first2=D. G.|last3=van Sittert |first3=S. J.|last4=Skinner|first4=J. D.|year=2013|title=Orbit orientation and eye morphometrics in giraffes (''Giraffa camelopardalis'') |journal=[[African Zoology]] |volume=48|issue=2|pages=333–339 |doi=10.1080/15627020.2013.11407600 |hdl=2263/37109 |s2cid=219292664 |hdl-access=free}}</ref> Giraffes may see in colour,<ref name=Dagg2014/>{{rp|85}} and their senses of hearing and [[olfaction|smell]] are sharp.<ref name="Prothero 2003"/> The ears are movable.<ref name=Dagg2014/>{{rp|95}} The nostrils are slit-shaped, possibly to withstand blowing sand.<ref>{{cite book|author=Peterson, D.|year=2013 |title=Giraffe Reflections |publisher=[[University of California Press]] |page=30 |isbn=978-0520266858}}</ref> The giraffe's [[tongue]] is about {{cvt|45|cm}} long. It is black, perhaps to protect against sunburn, and can [[prehensile|grasp]] foliage and delicately pick off leaves.<ref name=Dagg2014/>{{rp|109–110}} The upper lip is flexible and hairy to protect against sharp prickles.<ref name="Dagg1971"/> The upper jaw has a hard [[palate]] instead of front teeth. The [[Molar (tooth)|molar]]s and [[premolar]]s are wide with low [[Crown (tooth)|crowns]] on the surface.<ref name=Dagg2014/>{{rp|106}} ===Neck=== The giraffe has an extremely elongated neck, which can be up to {{cvt|2.4|m|ft}} in length.<ref name="Taylor&Wedel2013">{{Cite journal |last1=Taylor |first1=M. P. |last2=Wedel |first2=M. J. |doi=10.7717/peerj.36 |title=Why sauropods had long necks; and why giraffes have short necks |journal=[[PeerJ]] |volume=1 |page=e36 |year=2013 |pmid=23638372 |pmc=3628838 |doi-access=free}}</ref> Along the neck is a mane made of short, erect hairs.<ref name="Dagg1971"/> The neck typically rests at an angle of 50–60 degrees, though juveniles are closer to 70 degrees.<ref name=Dagg2014/>{{rp|72–73}} The long neck results from a disproportionate lengthening of the [[cervical vertebrae]], not from the addition of more vertebrae. Each cervical vertebra is over {{cvt|28|cm}} long.<ref name=anatomy>{{cite book |author=Swaby, S. |year=2010 |contribution=Giraffe |editor=Harris, T.|title=Mammal Anatomy: An Illustrated Guide |publisher=[[Marshall Cavendish]] |pages=64–84 |isbn=978-0-7614-7882-9}}</ref>{{rp|71}} They comprise 52–54 per cent of the length of the giraffe's [[vertebral column]], compared with the 27–33 percent typical of similar large ungulates, including the giraffe's closest living relative, the [[okapi]].<ref name=bada>{{Cite journal |doi=10.1111/j.1096-3642.2008.00458.x |last1=Badlangana |first1=L. N. |last2=Adams |first2=J. W. |last3=Manger |first3=P. R. |title=The giraffe ''(Giraffa camelopardalis)'' cervical vertebral column: A heuristic example in understanding evolutionary processes? |journal=[[Zoological Journal of the Linnean Society]] |volume=155 |issue=3 |pages=736–757 |year=2009 |doi-access=free}}</ref> This elongation largely takes place after birth, perhaps because giraffe mothers would have a difficult time [[giving birth]] to young with the same neck proportions as adults.<ref name=van>{{Cite journal |doi=10.1002/jez.b.21353 |pmid=20700891 |last1=Van Sittert |first1=S. J. |last2=Skinner |first2=J. D. |last3=Mitchell |first3=G. |title=From fetus to adult – An allometric analysis of the giraffe vertebral column |journal=[[Journal of Experimental Zoology Part B: Molecular and Developmental Evolution]] |volume=314B |issue=6 |pages=469–479 |year=2010|bibcode=2010JEZB..314..469V }}</ref> The giraffe's head and neck are held up by large muscles and a [[nuchal ligament]], which are anchored by long [[thoracic vertebrae]] spines, giving them a hump.<ref name=Dagg1971/><ref name=Solounias1999/><ref name=MacDonald/> [[File:Flickr - Rainbirder - High-rise living.jpg|thumb|left|upright|alt=Photograph of an adult male giraffe with its next fully extended feeding on an acacia|Adult male [[reticulated giraffe]] feeding high on an acacia, in Kenya]] The giraffe's neck vertebrae have [[ball and socket joint]]s.<ref name=anatomy/>{{rp|71}} The point of articulation between the cervical and thoracic vertebrae of giraffes is shifted to lie between the first and second thoracic vertebrae (T1 and T2), unlike in most other ruminants, where the articulation is between the seventh cervical vertebra (C7) and T1.<ref name=bada/><ref name=van/> This allows C7 to contribute directly to increased neck length and has given rise to the suggestion that T1 is actually C8, and that giraffes have added an extra cervical vertebra.<ref name=Solounias1999>{{Cite journal|author=Solounias, N.|year=1999|title=The remarkable anatomy of the giraffe's neck |journal=[[Journal of Zoology]] |volume=247 |issue=2 |pages=257–268 |doi=10.1111/j.1469-7998.1999.tb00989.x |url=http://www.ikhebeenvraag.be/mediastorage/FSDocument/73/download.pdf |archive-url=https://web.archive.org/web/20090325215447/http://www.ikhebeenvraag.be/mediastorage/FSDocument/73/download.pdf |archive-date=25 March 2009 |url-status=live}}</ref> However, this proposition is not generally accepted, as T1 has other [[morphology (biology)|morphological]] features, such as an articulating [[rib]], deemed diagnostic of thoracic vertebrae, and because exceptions to the mammalian limit of seven cervical vertebrae are generally characterised by increased [[neurological disorder|neurological anomalies]] and maladies.<ref name=bada/> There are several hypotheses regarding the evolutionary origin and maintenance of elongation in giraffe necks.<ref name=sim2010>{{Cite journal |doi=10.1111/j.1469-7998.2010.00711.x |last1=Simmons |first1= R. E. |last2=Altwegg |first2=R. | title=Necks-for-sex or competing browsers? A critique of ideas on the evolution of giraffe |journal=Journal of Zoology |volume=282 |issue=1 |pages=6–12 |year=2010}}</ref> [[Charles Darwin]] originally suggested the "competing [[browsing (predation)|browsers]] hypothesis", which has been challenged only recently. It suggests that competitive pressure from smaller browsers, like [[kudu]], [[steenbok]] and [[impala]], encouraged the elongation of the neck, as it enabled giraffes to reach food that competitors could not. This advantage is real, as giraffes can and do feed up to {{cvt|4.5|m}} high, while even quite large competitors, such as kudu, can feed up to only about {{cvt|2|m}} high.<ref name=dt1990>{{Cite journal |doi=10.1111/j.1365-2028.1990.tb01136.x |last=du Toit | first=J. T. |title=Feeding-height stratification among African browsing ruminants |journal=[[African Journal of Ecology]] |volume=28 |issue=1 |pages=55–62 |year=1990 |bibcode=1990AfJEc..28...55D |url=http://courses.biology.utah.edu/goller/7406/Goller7406/duToitPdfs/Feedingheightstrat_1990.pdf |access-date=21 November 2011 |url-status=dead|archive-date=10 November 2011 |archive-url=https://web.archive.org/web/20111110224355/http://courses.biology.utah.edu/goller/7406/Goller7406/duToitPdfs/Feedingheightstrat_1990.pdf}}</ref> There is also research suggesting that browsing competition is intense at lower levels, and giraffes feed more efficiently (gaining more leaf biomass with each mouthful) high in the canopy.<ref>{{Cite journal |last1=Cameron |first1=E. Z.|author-link=Elissa Cameron |last2=du Toit|first2=J. T.|year=2007|title=Winning by a Neck: Tall Giraffes Avoid Competing with Shorter Browsers |url=http://ecite.utas.edu.au/67394 |journal=[[American Naturalist]] |volume=169 |issue=1 |pages=130–135 |doi=10.1086/509940|pmid=17206591|bibcode=2007ANat..169..130C |s2cid=52838493|access-date=5 December 2019|archive-date=2 June 2020 |archive-url=https://web.archive.org/web/20200602031844/http://ecite.utas.edu.au/67394|url-status=live}}</ref><ref>{{Cite journal |author1=Woolnough, A. P. |author2=du Toit, J. T.| title=Vertical zonation of browse quality in tree canopies exposed to a size-structured guild of African browsing ungulates |journal=[[Oecologia]]| volume=129| issue=1| pages=585–590 |year=2001 |doi=10.1007/s004420100771| pmid=24577699 |bibcode=2001Oecol.129..585W|s2cid=18821024 |url=http://courses.biology.utah.edu/goller/7406/Goller7406/duToitPdfs/Verticalzonation_2001.pdf| access-date=7 March 2012| archive-url=https://web.archive.org/web/20111110224921/http://courses.biology.utah.edu/goller/7406/Goller7406/duToitPdfs/Verticalzonation_2001.pdf| archive-date=10 November 2011 |url-status=dead}}</ref> However, scientists disagree about just how much time giraffes spend feeding at levels beyond the reach of other browsers,<ref name=sim1996/><ref name=sim2010/><ref name=dt1990/><ref name=sexdiff>{{Cite journal| author1=Young, T. P.| author2=Isbell, L. A.| title=Sex differences in giraffe feeding ecology: energetic and social constraints |journal=[[Ethology (journal)|Ethology]]| volume=87| issue=1–2| pages=79–89| year=1991 |doi=10.1111/j.1439-0310.1991.tb01190.x | bibcode=1991Ethol..87...79Y|url=http://tpyoung.ucdavis.edu/publications/1991GiraffesEthology.pdf| access-date=2 February 2012| archive-url=https://web.archive.org/web/20130516034426/http://tpyoung.ucdavis.edu/publications/1991GiraffesEthology.pdf| archive-date=16 May 2013| url-status=dead}}</ref> and a 2010 study found that adult giraffes with longer necks actually suffered higher mortality rates under drought conditions than their shorter-necked counterparts. This study suggests that maintaining a longer neck requires more nutrients, which puts longer-necked giraffes at risk during a food shortage.<ref name=demography>{{cite journal|author1=Mitchell, G. |author2=van Sittert, S. |author3=Skinner, J. D. |year=2010 |title=The demography of giraffe deaths in a drought|journal=Transactions of the Royal Society of South Africa |volume=65|issue=3|pages=165–168|doi=10.1080/0035919X.2010.509153|bibcode=2010TRSSA..65..165M |hdl=2263/18957 |s2cid=83652889 |hdl-access=free}}</ref> Another theory, the [[sexual selection]] hypothesis, proposes that long necks evolved as a secondary [[sexual dimorphism|sexual characteristic]], giving males an advantage in "necking" contests (see below) to establish dominance and obtain access to sexually receptive females.<ref name=sim1996>{{Cite journal |author1=Simmons, R. E.|author2=Scheepers, L. |title=Winning by a Neck: Sexual Selection in the Evolution of Giraffe |journal=[[The American Naturalist]] |volume=148 |issue=5| pages=771–786 |year=1996 |url=http://bill.srnr.arizona.edu/classes/182/Giraffe/WinningByANeck.pdf |doi=10.1086/285955 |bibcode=1996ANat..148..771S |s2cid=84406669 |url-status=dead| archive-url=https://web.archive.org/web/20040823200801/http://bill.srnr.arizona.edu/classes/182/Giraffe/WinningByANeck.pdf| archive-date=23 August 2004}}</ref> In support of this theory, some studies have stated that necks are longer and heavier for males than females of the same age,<ref name=sim1996/><ref name=sim2010/> and that males do not employ other forms of combat.<ref name=sim1996/> However, a 2024 study found that, while males have thicker necks, females actually have proportionally longer ones, which is likely because of their greater need to find more food to sustain themselves and their dependent young.<ref name=Cavener2024>{{cite journal|last1=Cavener|first1=D. R.|last2=Bond|first2=M. L.|last3=Wu-Cavener|first3=L|last4=Lohay|first4=G. G.|last5=Cavener|first5=M. W.|last6=Hou|first6=X|last7=Pearce|first7=D. L.|last8=Lee|first8=D. E.|year=2024|title=Sexual dimorphisms in body proportions of Masai giraffes and the evolution of the giraffe's neck|journal=Mammalian Biology|volume=104 |issue=5 |pages=513–527 |doi=10.1007/s42991-024-00424-4|doi-access=free}}</ref> It has also been proposed that the neck serves to give the animal greater vigilance.<ref>{{cite journal|author=Brownlee, A. |year=1963|title=Evolution of the Giraffe|journal=[[Nature (journal)|Nature]] |volume=200 |issue=4910 |page=1022 |doi=10.1038/2001022a0 |bibcode=1963Natur.200.1022B|s2cid=4145785|doi-access=free}}</ref><ref>{{cite journal |last1=Williams|first1=E. M.|title=Giraffe Stature and Neck Elongation: Vigilance as an Evolutionary Mechanism |journal=Biology |volume=5 |issue=3 |page=35 |doi=10.3390/biology5030035 |year=2016 |pmc=5037354 |pmid=27626454 |doi-access=free}}</ref> ===Legs, locomotion and posture=== [[File:Masai Giraffe right-rear foot.jpg|thumb|upright|left|alt=Photograph of giraffe's hind leg|Right hind leg of a Masai giraffe at the [[San Diego Zoo]]]] The front legs tend to be longer than the hind legs,<ref name=Dagg2014/>{{rp|109}} and males have proportionally longer front legs than females, which gives them better support when swinging their necks during fights.<ref name=Cavener2024/> The leg bones lack first, second and fifth [[metapodial]]s.<ref name=Dagg2014/>{{rp|109}} It appears that a [[suspensory ligament]] allows the lanky legs to support the animal's great weight.<ref>{{cite web |author=Wood, C. |date=2014 |title=Groovy giraffes...distinct bone structures keep these animals upright|publisher=[[Society for Experimental Biology]] |url=http://www.eurekalert.org/pub_releases/2014-07/sfeb-ggd070314.php|access-date=7 May 2014|archive-date=25 November 2018|archive-url=https://web.archive.org/web/20181125074805/https://www.eurekalert.org/pub_releases/2014-07/sfeb-ggd070314.php|url-status=live}}</ref> The [[hoof|hooves]] of large male giraffes reach {{cvt|31|x|23|cm}} in diameter.<ref name=Dagg2014/>{{rp|98}} The [[fetlock]] of the leg is low to the ground, allowing the hoof to better support the animal's weight. Giraffes lack [[dewclaw]]s and interdigital glands. While the pelvis is relatively short, the [[Ilium (bone)|ilium]] has stretched-out crests.<ref name=Dagg1971/> A giraffe has only two [[gait]]s: walking and galloping. Walking is done by moving the legs on one side of the body, then doing the same on the other side.<ref name=estes/> When galloping, the hind legs move around the front legs before the latter move forward,<ref name="Prothero 2003"/> and the tail will curl up.<ref name=estes/> The movements of the head and neck provide balance and control momentum while galloping.<ref name=Kingdon1988/>{{rp|327–29}} The giraffe can reach a sprint speed of up to {{cvt|60|km/h}},<ref>{{Cite journal |last1=Garland |first1=T. |last2=J. |first2=C. M. |year=1993 | title=Does metatarsal/femur ratio predict maximal running speed in cursorial mammals? |url=http://www.biology.ucr.edu/people/faculty/Garland/GarlandJanis1993.pdf |journal=Journal of Zoology |volume=229 |issue=1 |pages=133–51 |doi=10.1111/j.1469-7998.1993.tb02626.x |access-date=25 April 2010 |archive-date=20 November 2018 | archive-url=https://web.archive.org/web/20181120033719/https://biology.ucr.edu/people/faculty/Garland/GarlandJanis1993.pdf | url-status=dead}}</ref> and can sustain {{cvt|50|km/h}} for several kilometres.<ref>{{Cite book|author=Rafferty, J. P. |year=2011|title=Grazers (Britannica Guide to Predators and Prey) |publisher=[[Britannica Educational Publishing]] |page=194 |isbn=978-1-61530-336-6 |url=https://archive.org/details/grazers0000raff/page/194}}</ref> Giraffes would probably not be competent swimmers as their long legs would be highly cumbersome in the water,<ref name=swim>{{cite journal |doi=10.1016/j.jtbi.2010.04.007 |author1=Henderson, D. M. |author2=Naish, D. |year=2010 |title=Predicting the buoyancy, equilibrium and potential swimming ability of giraffes by computational analysis |journal=[[Journal of Theoretical Biology]] |volume=265 |issue=2 |pages=151–59 |pmid=20385144 |bibcode=2010JThBi.265..151H}}</ref> although they might be able to float.<ref name=Naish>{{cite journal |last=Naish |first=D. |author-link=Darren Naish |date=2011 |title=Will it Float? |periodical=[[Scientific American]] |volume=304 |issue=1 |page=22 |url=http://www.scientificamerican.com/article.cfm?id=will-it-float |doi=10.1038/scientificamerican0111-22 |bibcode=2011SciAm.304a..22N |access-date=30 December 2010 | archive-date=4 December 2013 |archive-url=https://web.archive.org/web/20131204212753/http://www.scientificamerican.com/article.cfm?id=will-it-float |url-status=live}}</ref> When swimming, the thorax would be weighed down by the front legs, making it difficult for the animal to move its neck and legs in harmony<ref name=swim/><ref name=Naish/> or keep its head above the water's surface.<ref name="swim"/> [[File:Juvenile Giraffe - walking - Malawi - 2018 Aug.webm|thumb|right|A juvenile giraffe walking in Malawi]] A giraffe rests by lying with its body on top of its folded legs.<ref name=Kingdon1988/>{{rp|329}} To lie down, the animal kneels on its front legs and then lowers the rest of its body. To get back up, it first gets on its front knees and positions its backside on top of its hindlegs. It then pulls the backside upwards, and the front legs stand straight up again. At each stage, the animal swings its head for balance.<ref name=Dagg2014/>{{rp|67}} If the giraffe wants to reach down to drink, it either spreads its front legs or bends its knees.<ref name=estes/> Studies in captivity found the giraffe sleeps intermittently around 4.6 hours per day, mostly at night. It usually sleeps lying down; however, standing sleeps have been recorded, particularly in older individuals. Intermittent short "deep sleep" phases while lying are characterised by the giraffe bending its neck backwards and resting its head on the hip or thigh, a position believed to indicate [[paradoxical sleep]].<ref name=sleep>{{cite journal |last1=Tobler |first1=I. |last2=Schwierin |first2=B. |title=Behavioural sleep in the giraffe (''Giraffa camelopardalis'') in a zoological garden |year=1996 |journal=Journal of Sleep Research |volume=5 |issue=1 |pages=21–32 |doi=10.1046/j.1365-2869.1996.00010.x |pmid=8795798|s2cid=34605791 |doi-access=free}}</ref> ===Internal systems=== [[File:GiraffaRecurrEn.svg|thumb|right|alt=Sketch of the path of the recurrent laryngeal nerve in giraffe|Scheme of path of the recurrent laryngeal nerve in giraffe]] In mammals, the left [[recurrent laryngeal nerve]] is longer than the right; in the giraffe, it is over {{cvt|30|cm}} longer. These nerves are longer in the giraffe than in any other living animal;<ref name="Wedel">{{Cite journal|author=Wedel, M. J.|year=2012|title=A monument of inefficiency: the presumed course of the recurrent laryngeal nerve in sauropod dinosaurs|journal=[[Acta Palaeontologica Polonica]] |volume=57 |issue=2 |pages=251–256 |doi=10.4202/app.2011.0019|s2cid=43447891 |doi-access=free |archive-date=29 October 2013 |url-status=live |url=http://www.app.pan.pl/archive/published/app57/app20110019.pdf |archive-url=https://web.archive.org/web/20131029184627/http://www.app.pan.pl/archive/published/app57/app20110019.pdf}}</ref> the left nerve is over {{cvt|2|m}} long.<ref>{{cite book|author=Harrison, D. F. N. |year=1995 |title=The Anatomy and Physiology of the Mammalian Larynx|publisher=Cambridge University Press |page=165 |isbn=978-0-521-45321-9}}</ref> Each nerve cell in this path begins in the [[brainstem]] and passes down the neck along the [[vagus nerve]], then branches off into the recurrent laryngeal nerve which passes back up the neck to the larynx. Thus, these nerve cells have a length of nearly {{cvt|5|m}} in the largest giraffes.<ref name=Wedel/> Despite its long neck and large skull, the [[brain]] of the giraffe is typical for an ungulate.<ref>{{cite journal |author1=Graïc, J.-M.|author2=Peruffo, A. |author3=Ballarin, C.|author4=Cozzi, B. |year=2017|title=The brain of the giraffe (''Giraffa camelopardalis''): surface configuration, encephalization quotient, and analysis of the existing literature|journal=The Anatomical Record |volume=300 |issue=8 |pages=1502–1511 |doi=10.1002/ar.23593 |pmid=28346748 |s2cid=3634656|doi-access=free}}</ref> Evaporative heat loss in the nasal passages keep the giraffe's brain cool.<ref name=Mitchell2004/> The shape of the skeleton gives the giraffe a small [[lung]] volume relative to its mass. Its long neck gives it a large amount of [[Dead space (physiology)|dead space]], though this is limited by its narrow windpipe. The giraffe also has a high [[tidal volume]], so the balance of dead space and tidal volume is much the same as other mammals. The animal can still provide enough oxygen for its tissues, and it can increase its [[Breathing|respiratory]] rate and oxygen diffusion when running.<ref name=SkinnerJD2011>{{Cite journal |author1=Skinner, J. D.|author2= Mitchell, G.|year= 2011|title=Lung volumes in giraffes, ''Giraffa camelopardalis'' |journal=Comparative Biochemistry and Physiology A |volume=158 |issue=1 |pages=72–78 |doi=10.1016/j.cbpa.2010.09.003 |pmid=20837156 |url=http://137.215.9.22/bitstream/handle/2263/16472/Mitchell_Lung(2011).PDF?sequence=1 |hdl=2263/16472|hdl-access= free|access-date= 27 November 2011|archive-date=20 November 2018 |url-status=live |archive-url=https://web.archive.org/web/20181120033132/https://137.215.9.22/bitstream/handle/2263/16472/Mitchell_Lung(2011).PDF?sequence=1}}</ref> [[File:Flickr - Rainbirder - Reticulated Giraffe drinking.jpg|thumb|left|[[Reticulated giraffe]] bending down to drink in Kenya. The circulatory system is adapted to deal with blood flow rushing down its neck.|alt=Photograph of a giraffe bending down to drink]] The giraffe's [[circulatory system]] has several adaptations to compensate for its great height.<ref name=Holmes>{{cite magazine |last1=Holmes |first1=B. |title=Heads up! The cardiovascular secrets of giraffes |magazine=Knowable Magazine |date=2021 |doi=10.1146/knowable-051821-2 |s2cid=236354545 |doi-access=free |url=https://knowablemagazine.org/article/living-world/2021/heads-up-cardiovascular-secrets-giraffes |access-date=1 August 2022 |archive-date=6 July 2022 |archive-url=https://web.archive.org/web/20220706111325/https://knowablemagazine.org/article/living-world/2021/heads-up-cardiovascular-secrets-giraffes |url-status=live}}</ref> Its {{cvt|25|lb|kg|order=flip}} and {{cvt|2|ft|cm|order=flip|-1}} [[heart]] must generate approximately double the blood pressure required for a human to maintain blood flow to the brain. As such, the wall of the heart can be as thick as {{cvt|7.5|cm}}.<ref name="Prothero 2003"/> Giraffes have relatively high heart rates for their size, at 150 beats per minute.<ref name=anatomy/>{{rp|76}} When the animal lowers its head, the blood rushes down fairly unopposed and a [[rete mirabile]] in the upper neck, with its large [[Cross section (geometry)|cross-sectional]] area, prevents excess blood flow to the brain. When it raises again, the blood vessels constrict and push blood into the brain so the animal does not faint.<ref>{{cite journal|author1=Mitchell, G. |author2=Skinner, J. D. |year=1993|title=How giraffe adapt to their extraordinary shape |journal=[[Transactions of the Royal Society of South Africa]]|volume=48|issue=2 |pages=207–218 |doi=10.1080/00359199309520271|bibcode=1993TRSSA..48..207M }}</ref> The [[jugular vein]]s contain several (most commonly seven) valves to prevent blood flowing back into the head from the [[inferior vena cava]] and [[right atrium]] while the head is lowered.<ref>{{cite journal |doi=10.3957/056.039.0210 |author1=Mitchell, G.|author2=van Sittert, S. J.|author3=Skinner, J. D. |title=The structure and function of giraffe jugular vein valves |journal=South African Journal of Wildlife Research |volume=39 |issue=2|pages=175–180 |url=http://137.215.9.22/bitstream/handle/2263/13994/Mitchell_Structure(2009).pdf?sequence=1 |year=2009 |hdl=2263/13994 |s2cid=55201969 |hdl-access=free|access-date=21 November 2011|archive-date=20 November 2018|archive-url=https://web.archive.org/web/20181120033146/https://137.215.9.22/bitstream/handle/2263/13994/Mitchell_Structure(2009).pdf?sequence=1|url-status=live}}</ref> Conversely, the blood vessels in the lower legs are under great pressure because of the weight of fluid pressing down on them. To solve this problem, the skin of the lower legs is thick and tight, preventing too much blood from pouring into them.<ref name=MacDonald/> Giraffes have [[Esophagus|oesophageal muscles]] that are strong enough to allow regurgitation of food from the stomach up the neck and into the mouth for [[wikt:ruminate|rumination]].<ref name=anatomy/>{{rp|78}} They have [[Ruminant#Digestive system of ruminants|four-chambered stomachs]], which are adapted to their specialized diet.<ref name=Dagg1971/> The [[intestine]]s of an adult giraffe measure more than {{cvt|70|m}} in length and have a relatively small ratio of [[small intestine|small]] to [[large intestine]].<ref name="PerezEtAl2009">{{cite journal |author1=Pérez, W. |author2=Lima, M. |author3=Clauss, M. |year=2009 |title=Gross anatomy of the intestine in the giraffe (''Giraffa camelopardalis'') |url=https://www.zora.uzh.ch/id/eprint/24084/5/P%C3%A9rez_giraffe_intestine_AHE_revised-1.pdf |archive-url=https://web.archive.org/web/20180722150740/http://www.zora.uzh.ch/id/eprint/24084/5/P%C3%A9rez_giraffe_intestine_AHE_revised-1.pdf |archive-date=22 July 2018 |url-status=live |journal=Anatomia, Histologia, Embryologia |volume=38 |issue=6 |pages=432–435 |pmid=19681830 |doi=10.1111/j.1439-0264.2009.00965.x|s2cid=28390695}}</ref> The giraffe has a small, compact liver.<ref name=anatomy/>{{rp|76}} In fetuses there may be a small [[gallbladder]] that vanishes before birth.<ref name="Dagg1971"/><ref>{{cite journal |author=Cave, A. J. E. | title=On the liver and gall-bladder of the Giraffe |journal=[[Proceedings of the Zoological Society of London]] |volume=120 |pages=381–93 |year=1950 |doi=10.1111/j.1096-3642.1950.tb00956.x | issue=2}}</ref><ref>{{cite journal |author1=Oldham-Ott, C. K. |author2=Gilloteaux, J. |title=Comparative morphology of the gallbladder and biliary tract in vertebrates: variation in structure, homology in function and gallstones |journal=[[Microscopy Research and Technique]] |volume=38 |issue=6 |pages=571–579 |year=1997 |doi=10.1002/(SICI)1097-0029(19970915)38:6<571::AID-JEMT3>3.0.CO;2-I |pmid=9330347 |s2cid=20040338}}</ref> ==Behaviour and ecology== ===Habitat and feeding=== {{multiple image | direction =vertical | image1 =Giraffe feeding, Tanzania.jpg | caption1 =A [[Masai giraffe]] extending its tongue to feed, in Tanzania. | image2 =Giraffe in Malawi - 2018 Aug.webm | caption2 =A giraffe in Malawi eating leaves from a tree }} Giraffes usually inhabit savannahs and open [[woodland]]s. They prefer areas dominated by [[Acacieae]], ''[[Commiphora]]'', ''[[Combretum]]'' and ''[[Terminalia (plant)|Terminalia]]'' trees over ''[[Brachystegia]]'' which are more densely spaced.<ref name=Kingdon1988/>{{rp|322}} The Angolan giraffe can be found in desert environments.<ref>{{cite thesis|author=Fennessy, J. |year=2004 |title=Ecology of desert-dwelling giraffe ''Giraffa camelopardalis angolensis'' in northwestern Namibia|publisher=[[University of Sydney]]|degree=[[Doctor of Philosophy|PhD]]|url=http://ses.library.usyd.edu.au/handle/2123/910|access-date=30 January 2012|archive-date=20 November 2018|archive-url=https://web.archive.org/web/20181120033134/https://ses.library.usyd.edu.au/handle/2123/910|url-status=live}}</ref> Giraffes browse on the twigs of trees, preferring those of the subfamily Acacieae and the genera ''Commiphora'' and ''Terminalia'',<ref name=kingdon>{{Cite book|title=The Kingdon Field Guide to African Mammals|author=Kingdon, J.|publisher=[[Academic Press]] |chapter=Giraffes Giraffidae |pages=339–344 |year=1997 |isbn=978-0-12-408355-4 |chapter-url=https://archive.org/details/kingdonfieldguid00jona/page/339}}</ref> which are important sources of calcium and protein to sustain the giraffe's growth rate.<ref name="Mitchell20003"/> They also feed on shrubs, grass and fruit.<ref name=Kingdon1988/>{{rp|324}} A giraffe eats around {{cvt|34|kg}} of plant matter daily.<ref name=estes/> When stressed, giraffes may chew on large branches, stripping them of bark.<ref name=Kingdon1988/>{{rp|325}} Giraffes are also recorded to [[Osteophagy|chew old bones]].<ref name=Dagg2014/>{{rp|102}} During the wet season, food is abundant and giraffes are more spread out, while during the dry season, they gather around the remaining evergreen trees and bushes.<ref name=kingdon/> Mothers tend to feed in open areas, presumably to make it easier to detect predators, although this may reduce their feeding efficiency.<ref name=sexdiff /> As a [[ruminant]], the giraffe first chews its food, then swallows it for processing and then visibly passes the half-digested cud up the neck and back into the mouth to chew again.<ref name=anatomy/>{{rp|78–79}} The giraffe requires less food than many other herbivores because the foliage it eats has more concentrated nutrients and it has a more efficient digestive system.<ref name=kingdon/> The animal's faeces come in the form of small pellets.<ref name="Dagg1971"/> When it has access to water, a giraffe will go no more than three days without drinking.<ref name=estes/> Giraffes have a great effect on the trees that they feed on, delaying the growth of young trees for some years and giving "waistlines" to particularly tall trees. Feeding is at its highest during the first and last hours of daytime. Between these hours, giraffes mostly stand and ruminate. Rumination is the dominant activity during the night, when it is mostly done lying down.<ref name=estes/> ===Social life=== [[File:Giraffe (Giraffa camelopardalis) females.jpg|thumb|left|alt=Photograph of a gathering of four female giraffes|Gathering of female South African giraffes in [[Tswalu Kalahari Reserve]], South Africa. These animals commonly gather in herds.]] Giraffes usually form groups that vary in size and composition according to ecological, anthropogenic, temporal, and social factors.<ref name=":5">{{Cite journal|last1=Bond|first1=M. L.|last2=Lee|first2=Derek E. |last3=Ozgul |first3=A. |last4=König |first4=B. |date=2019 |title=Fission–fusion dynamics of a megaherbivore are driven by ecological, anthropogenic, temporal, and social factors |journal=[[Oecologia]] |volume=191 |issue=2|pages=335–347 |doi=10.1007/s00442-019-04485-y |pmid=31451928 |bibcode=2019Oecol.191..335B|s2cid=201732871 |url=https://scholarsphere.psu.edu/resources/80d99e21-8a05-4eaf-bb4e-ba8bcddd109d |access-date=22 August 2023|archive-date=22 September 2023 |url-status=live |archive-url=https://web.archive.org/web/20230922074125/https://scholarsphere.psu.edu/resources/80d99e21-8a05-4eaf-bb4e-ba8bcddd109d}}</ref> Traditionally, the composition of these groups had been described as open and ever-changing.<ref name="Manyara">{{Cite journal |author=van der Jeugd, H. P. |author2=Prins, H. H. T. |title=Movements and group structure of giraffe (''Giraffa camelopardalis'') in Lake Manyara National Park, Tanzania |journal=[[Journal of Zoology]] |volume=251 |issue=1 |pages=15–21 |year=2000 |doi=10.1111/j.1469-7998.2000.tb00588.x |url=http://www.resource-ecology.org/publ/2000_Jeugd,Prins_MovementsAndGroupStructureOfGiraffeInLakeManyara.pdf |archive-url=https://web.archive.org/web/20131106025238/http://www.resource-ecology.org/publ/2000_Jeugd,Prins_MovementsAndGroupStructureOfGiraffeInLakeManyara.pdf |archive-date=6 November 2013 |url-status=dead}}</ref> For research purposes, a "group" has been defined as "a collection of individuals that are less than a kilometre apart and moving in the same general direction".<ref name="Pratt 1985">{{cite journal |doi=10.1080/00222938500770471 |author1=Pratt, D. M. |author2=Anderson, V. H. |year=1985 |title=Giraffe social behavior |journal=[[Journal of Natural History]] |volume=19 |issue=4 |pages=771–781|bibcode=1985JNatH..19..771P }}</ref> More recent studies have found that giraffes have long-lasting social groups or cliques based on kinship, sex or other factors, and these groups regularly associate with other groups in larger communities or sub-communities within a [[fission–fusion society]].<ref name=":7">{{Cite journal|last1=Bond|first1=M. L.|last2=König|first2=B. |last3=Lee |first3=D. E. |last4=Ozgul |first4=A. |last5=Farine |first5=D. R. |title=Proximity to humans affects local social structure in a giraffe metapopulation |journal=[[Journal of Animal Ecology]] |year=2020 |volume=90 |issue=1 |pages=212–221|doi=10.1111/1365-2656.13247 |pmid=32515083 |doi-access=free}}</ref><ref name="Bercovitch 2013">{{cite journal|author1=Bercovitch, F. B. |author2=Berry, P. S. M. |year=2013|title=Herd composition, kinship and fission–fusion social dynamics among wild giraffe |journal=[[African Journal of Ecology]] |volume=51 |issue=2 |pages=206–216 |doi=10.1111/aje.12024 |doi-access=free|bibcode=2013AfJEc..51..206B }}</ref><ref name=Carter>{{cite journal|author1=Carter, K. D. |author2=Seddon, J. M. |author3=Frèreb, C. H. |author4=Carter, J. K. |year=2013|title=Fission–fusion dynamics in wild giraffes may be driven by kinship, spatial overlap and individual social preferences |journal=[[Animal Behaviour (journal)|Animal Behaviour]]|volume=85|issue=2|pages=385–394 |doi=10.1016/j.anbehav.2012.11.011 |s2cid=53176817}}</ref><ref name=VanderWaal>{{cite journal|author1=VanderWaal, K. L. |author2=Wang, H. |author3=McCowan, B. |author4=Fushing, H. |author5=Isbell, L. A. |year=2014|title=Multilevel social organization and space use in reticulated giraffe (''Giraffa camelopardalis'') |journal=[[Behavioral Ecology (journal)|Behavioral Ecology]] |volume=25 |issue=1 |pages=17–26 |doi=10.1093/beheco/art061|doi-access=}}</ref> Proximity to humans can disrupt social arrangements.<ref name=":7" /> Masai giraffes in Tanzania sort themselves into different subpopulations of 60–90 adult females with overlapping ranges, each of which differ in reproductive rates and calf mortality.<ref name=":9">{{Cite journal |last1=Bond |first1=M. L. |last2=König |first2=B. |last3=Ozgul |first3=A. |last4=Farine |first4=D. R. |last5=Lee |first5=D. E. |title=Socially defined subpopulations reveal demographic variation in a Giraffe metapopulation |journal=[[The Journal of Wildlife Management]] |year=2021 |volume=85 |issue=5 |pages=920–931 |doi=10.1002/jwmg.22044 |bibcode=2021JWMan..85..920B |s2cid=233600744 |url=http://nbn-resolving.de/urn:nbn:de:bsz:352-2-127dcuwx9z4bj0 |access-date=27 January 2024 |archive-date=8 March 2022 |archive-url=https://web.archive.org/web/20220308221559/https://wildlife.onlinelibrary.wiley.com/doi/abs/10.1002/jwmg.22044 |url-status=dead }}</ref> [[Biological dispersal|Dispersal]] is male biased, and can include spatial and/or social dispersal.<ref>{{Cite journal |last1=Bond |first1=M. L. |last2=Lee |first2=D. E. |last3=Ozgul |first3=A. |last4=Farine |first4=D. R. |last5=König |first5=B. |date=2021 |title=Leaving by staying: Social dispersal in giraffes |journal=Journal of Animal Ecology |volume=90 |issue=12 |pages=2755–2766 |doi=10.1111/1365-2656.13582 |pmc=9291750|bibcode=2021JAnEc..90.2755B }}</ref> Adult female subpopulations are connected by males into super communities of around 300 animals.<ref>{{Cite journal |last1=Lavista Ferres |first1=J. M. |last2=Lee |first2=D. E. |last3=Nasir |first3=Md. |last4=Chen |first4=Y.-C. |last5=Bijral |first5=A. S. |last6=Bercovitch |first6=F. B. |last7=Bond |first7=M. L. |date=2021 |title=Social connectedness and movements among communities of giraffes vary by sex and age class |journal=Animal Behaviour |volume=180 |pages=315–328 |s2cid=237949827 |doi=10.1016/j.anbehav.2021.08.008|url=https://scholarsphere.psu.edu/resources/4e92c3e0-e2f6-47b6-8a25-d5c78210af75 }}</ref> {{listen |filename=Giraffe Hum.oga |title=Giraffe hum |description=Giraffe hum, [[ogg]]/[[Vorbis]] format. |format=Vorbis |filename2=Giraffe snort.oga |title2=Giraffe snort |description2=Giraffe snort ogg/Vorbis format. |format2=Vorbis |filename3=Giraffe grunt.oga |title3=Giraffe grunt |description3=Giraffe grunt ogg/Vorbis format. |format3=Vorbis |filename4= Giraffe bursts.oga |title4=Giraffe bursts |description4=Giraffe bursts ogg/Vorbis format |format4=Vorbis}} The number of giraffes in a group can range from one up to 66 individuals.<ref name=":5" /><ref name=VanderWaal/> Giraffe groups tend to be sex-segregated<ref name=VanderWaal/> although mixed-sex groups made of adult females and young males also occur.<ref name="Pratt 1985" /> Female groups may be [[matrilineally]] related.<ref name=VanderWaal/> Generally, females are more selective than males when deciding which individuals of the same sex they associate with.<ref name=Carter/> Particularly stable giraffe groups are those made of mothers and their young,<ref name="Pratt 1985" /> which can last weeks or months.<ref name="Leuthold 1979" /> Young males also form groups and will engage in playfights. However, as they get older, males become more solitary but may also associate in pairs or with female groups.<ref name=VanderWaal/><ref name="Leuthold 1979" /> Giraffes are not [[Territory (animal)|territorial]],<ref name="Dagg1971" /> but they have [[home range]]s that vary according to rainfall and proximity to human settlements.<ref>{{Cite journal |last1=Knüsel |first1=M. |last2=Lee |first2=D. |last3=König |first3=B. |last4=Bond |first4=M. |date=2019 |title=Correlates of home range sizes of giraffes, ''Giraffa camelopardalis'' |doi=10.1016/j.anbehav.2019.01.017 |journal=[[Animal Behaviour (journal)|Animal Behaviour]] |volume=149 |pages=143–151 |s2cid=72332291 |url=https://www.zora.uzh.ch/id/eprint/171320/2/ZORA_Knuesel_et_al_2019_Correlates_of_home_range_size_of_giraffes.pdf |archive-url=https://web.archive.org/web/20200210212658/https://www.zora.uzh.ch/id/eprint/171320/2/ZORA_Knuesel_et_al_2019_Correlates_of_home_range_size_of_giraffes.pdf |archive-date=10 February 2020 |url-status=live}}</ref> Male giraffes occasionally roam far from areas that they normally frequent.<ref name=Kingdon1988/>{{rp|329}} Early biologists suggested giraffes were mute and unable to create enough air flow to vibrate their [[vocal fold]]s.<ref name=Kasozi/> This has been proved to the contrary; they have been recorded to [[Animal communication|communicate]] using snorts, sneezes, coughs, snores, hisses, bursts, moans, grunts, growls and flute-like sounds.<ref name=estes/><ref name=Kasozi>{{cite journal|last1=Kasozi|first1=H.|last2=Montgomery|first2=R. A. |year=2018|title=How do giraffes locate one another? A review of visual, auditory, and olfactory communication among giraffes |journal=Journal of Zoology |volume=306|issue=3|pages=139–146|doi=10.1111/jzo.12604|doi-access=free}}</ref> During courtship, males emit loud coughs. Females call their young by bellowing. Calves will emit bleats, mooing and mewing sounds.<ref name=estes/> Snorting and hissing is associated with vigilance.<ref>{{cite journal|last1=Volodina|first1=Elena V.|last2=Volodin|first2=Ilya A.|last3=Chelysheva|first3=Elena V.|last4=Frey|first4=Roland|year=2018|title=Hiss and snort call types of wild-living giraffes ''Giraffa camelopardalis'': acoustic structure and context|journal=[[BMC Research Notes]]|volume=11|issue=12|page=12|doi=10.1186/s13104-017-3103-x|pmid=29316966|pmc=5761111 |doi-access=free}}</ref> During nighttime, giraffes appear to [[humming|hum]] to each other.<ref name="hum">{{cite journal|author1=Baotic, A. |author2=Sicks, F. |author3=Stoeger, A. S. |year=2015|title=Nocturnal "humming" vocalizations: adding a piece to the puzzle of giraffe vocal communication |journal=BMC Research Notes |volume=8 |page=425 |doi=10.1186/s13104-015-1394-3 |pmc=4565008 |pmid=26353836 |doi-access=free}}</ref> There is some evidence that giraffes use [[Helmholtz resonance]] to create [[infrasound]].<ref>{{cite journal|last1=Von Muggenthaler|first1=E. |year=2013|title=Giraffe Helmholtz resonance |journal=Proceedings of Meetings on Acoustics |volume=19|issue=1|page=010012|doi=10.1121/1.4800658|doi-access=free}}</ref> They also communicate with body language. [[Dominance hierarchy|Dominant]] males display to other males with an erect posture; holding the chin and head up while walking stiffly and displaying their side. The less dominant show submissiveness by dropping the head and ears, lowering the chin and fleeing.<ref name=estes/> ===Reproduction and parental care=== [[File:Giraffa camelopardalis angolensis (mating).jpg|thumb|left|upright|alt=Photograph of giraffes mating|Angolan giraffes mating in Namibia]] Reproduction in giraffes is broadly [[Animal sexual behaviour#Polygamy|polygamous]]: a few older males mate with the fertile females.<ref name="Pratt 1985"/> Females can reproduce throughout the year and experience [[oestrus]] cycling approximately every 15 days.<ref name=":8">{{Cite journal|last1=del Castillo|first1=S. M.|last2=Bashaw|first2=M. J.|last3=Patton|first3=M. L. |last4=Rieches|first4=R. R.|last5=Bercovitch|first5=F. B.|date=2005 |title=Fecal steroid analysis of female giraffe (''Giraffa camelopardalis'') reproductive condition and the impact of endocrine status on daily time budgets |url=https://pubmed.ncbi.nlm.nih.gov/15804514/|journal=[[General and Comparative Endocrinology]] |volume=141|issue=3|pages=271–281 |doi=10.1016/j.ygcen.2005.01.011 |pmid=15804514|access-date=10 February 2021 |archive-date=25 May 2021 |archive-url=https://web.archive.org/web/20210525182409/https://pubmed.ncbi.nlm.nih.gov/15804514/|url-status=live}}</ref><ref>{{Cite journal |last1=Bercovitch|first1=F. B.|last2=Bashaw|first2=M. J.|last3=del Castillo|first3=S. M.|date=2006 |title=Sociosexual behavior, male mating tactics, and the reproductive cycle of giraffe ''Giraffa camelopardalis'' |journal=[[Hormones and Behavior]] |volume=50 |issue=2 |url=https://www.sciencedirect.com/science/article/pii/S0018506X06000900 |pages=314–321 |doi=10.1016/j.yhbeh.2006.04.004 |pmid=16765955 |s2cid=45843281}}</ref> Female giraffes in oestrus are dispersed over space and time, so reproductive adult males adopt a strategy of roaming among female groups to seek mating opportunities, with periodic hormone-induced rutting behaviour approximately every two weeks.<ref>{{Cite journal|last1=Seeber|first1=P. A. |last2=Duncan|first2=P. |last3=Fritz|first3=H. |last4=Ganswindt |first4=A. |date=2013 |title=Androgen changes and flexible rutting behaviour in male giraffes |url= |journal=[[Biology Letters]] |volume=9 |issue=5 |page=20130396 |doi=10.1098/rsbl.2013.0396 |pmc=3971675 |pmid=23925833}}</ref> Males prefer young adult females over juveniles and older adults.<ref name="Pratt 1985"/> Male giraffes assess female fertility by tasting the female's urine to detect oestrus, in a multi-step process known as the [[flehmen response]].<ref name="Pratt 1985"/><ref name="Leuthold 1979">{{Cite journal|author=Leuthold, B. M.|year=1979|title=Social organization and behaviour of giraffe in Tsavo East National Park|journal=African Journal of Ecology |volume=17|issue=1|pages=19–34 |doi=10.1111/j.1365-2028.1979.tb00453.x|bibcode=1979AfJEc..17...19L }}</ref> Once an oestrous female is detected, the male will attempt to court her. When courting, dominant males will keep subordinate ones at bay.<ref name="Leuthold 1979"/> A courting male may lick a female's tail, lay his head and neck on her body or nudge her with his ossicones. During copulation, the male stands on his hind legs with his head held up and his front legs resting on the female's sides.<ref name=estes/> Giraffe [[pregnancy (mammals)|gestation]] lasts 400–460 days, after which a single calf is normally born, although twins occur on rare occasions.<ref name=":8" /> The mother gives birth standing up. The calf emerges head and front legs first, having broken through the [[fetal membrane]]s, and falls to the ground, severing the [[umbilical cord]].<ref name="Dagg1971"/> A newborn giraffe is {{cvt|1.7|-|2|m |sigfig=2}} tall.<ref name=Nowak1999/> Within a few hours of birth, the calf can run around and is almost indistinguishable from a one-week-old. However, for the first one to three weeks, it spends most of its time hiding,<ref name="Langman 1977">{{Cite journal |author=Langman, V. A. |year=1977 |title=Cow-calf relationships in giraffe (''Giraffa camelopardalis giraffa'') |journal=Zeitschrift für Tierpsychologie |volume=43 |issue=3 |pages=264–286 |doi=10.1111/j.1439-0310.1977.tb00074.x |url=https://onlinelibrary.wiley.com/doi/abs/10.1111/j.1439-0310.1977.tb00074.x |access-date=12 January 2020 |archive-date=12 January 2020 |archive-url=https://web.archive.org/web/20200112100644/https://onlinelibrary.wiley.com/doi/abs/10.1111/j.1439-0310.1977.tb00074.x |url-status=dead }}</ref> its coat pattern providing camouflage. The ossicones, which have lain flat in the womb, raise up in a few days.<ref name=estes/> [[File:Angolan giraffe (Giraffa camelopardalis angolensis) female with young 2 months.jpg|thumb|alt=A female giraffe with her calf|Female Angolan giraffe with calf]] Mothers with calves will gather in nursery herds, moving or browsing together. Mothers in such a group may sometimes leave their calves with one female while they forage and drink elsewhere. This is known as a "[[Crèche (zoology)|calving pool]]".<ref name="Langman 1977"/> Calves are at risk of predation, and a mother giraffe will stand over them and kick at an approaching predator.<ref name=estes/> Females watching calving pools will only alert their own young if they detect a disturbance, although the others will take notice and follow.<ref name="Langman 1977"/> Allo-sucking, where a calf will suckle a female other than its mother, has been recorded in both wild and captive giraffes.<ref>{{cite journal|last1=Saito|first1=M. |last2=Idani|first2=G. |year=2018|title=Suckling and allosuckling behavior in wild giraffe (''Giraffa camelopardalis tippelskirchi'') |journal=[[Mammalian Biology]]|volume=93|pages=1–4 |doi=10.1016/j.mambio.2018.07.005 |bibcode=2018MamBi..93....1S |s2cid=91472891}}</ref><ref>{{cite journal |last1=Gloneková |first1=M. |last2=Brandlová |first2=K. |last3=Pluháček |first3=J. |year=2021 |title=Further behavioural parameters support reciprocity and milk theft as explanations for giraffe allonursing |journal=Scientific Reports |volume=11 |issue=1 |page=7024 |doi=10.1038/s41598-021-86499-2 |pmid=33782483 |pmc=8007720|bibcode=2021NatSR..11.7024G}}</ref> Calves first ruminate at four to six months and stop [[lactation|nursing]] at six to eight months. Young may not reach independence until they are 14 months old.<ref name=Dagg2014/>{{rp|49}} Females are able to reproduce at four years of age,<ref name=estes/> while [[spermatogenesis]] in males begins at three to four years of age.<ref>{{cite journal |author=Hall-Martin, A. J. |author2=Skinner, J. D. |author3=Hopkins, B. J. |year=1978 |title=The development of the reproductive organs of the male giraffe, ''Giraffa camelopardalis'' |url=http://www.reproduction-online.org/content/52/1/1.full.pdf |journal=[[Journal of Reproduction and Fertility]] |volume=52 |issue=1 |pages=1–7 |doi=10.1530/jrf.0.0520001 |pmid=621681 |doi-access=free |access-date=12 June 2017 |archive-date=19 July 2018 |archive-url=https://web.archive.org/web/20180719113129/http://www.reproduction-online.org/content/52/1/1.full.pdf |url-status=live}}</ref> Males must wait until they are at least seven years old to gain the opportunity to mate.<ref name=estes/> {{Clear}} ===Necking=== [[File:Giraffe Ithala KZN South Africa Luca Galuzzi 2004.JPG|thumb|right|alt=Photograph of two male giraffes necking to establish dominance|Here, male South African giraffes engage in low intensity necking to establish dominance, in [[Ithala Game Reserve]], [[KwaZulu-Natal|Kwa-Zulu-Natal]], South Africa.]] Male giraffes use their necks as [[weapon (biology)|weapons]] in combat, a behaviour known as "necking". Necking is used to establish dominance, and males that win necking bouts have greater [[reproductive success]].<ref name=sim1996/> This behaviour occurs at low or high intensity. In low-intensity necking, the combatants rub and lean on each other. The male that can keep itself more upright wins the bout. In high-intensity necking, the combatants will spread their front legs and swing their necks at each other, attempting to land blows with their ossicones. The contestants will try to dodge each other's blows and then prepare to counter. The power of a blow depends on the weight of the skull and the arc of the swing.<ref name=estes/> A necking duel can last more than half an hour, depending on how well matched the combatants are.<ref name=Kingdon1988/>{{rp|331}} Although most fights do not lead to serious injury, there have been records of broken jaws, broken necks, and even deaths.<ref name=sim1996/> After a duel, it is common for two male giraffes to caress and court each other. Such interactions between males have been found to be more frequent than heterosexual coupling.<ref>{{Cite journal|author=Coe, M. J.|year=1967|title='Necking' behavior in the giraffe |journal=Journal of Zoology |volume=151|issue=2|pages=313–21|doi= 10.1111/j.1469-7998.1967.tb02117.x}}</ref> In one study, up to 94 percent of observed mounting incidents took place between males. The proportion of [[Homosexual behavior in animals|same-sex activities]] varied from 30 to 75 percent. Only one percent of same-sex mounting incidents occurred between females.<ref>{{Cite book |author=Bagemihl, B. |year=1999|title=Biological Exuberance: Animal Homosexuality and Natural Diversity|publisher=St. Martin's Press |pages=391–393 |isbn=978-0-312-19239-6|url=https://archive.org/details/biologicalexuber00bage/page/391}}</ref> ===Mortality and health=== [[File:Lioness with giraffe kill, jackal lurking, kenya, august 9th 2012.jpg|thumb|alt=Photograph of a lioness with at an adult giraffe kill|[[Lion]]ess seen with an adult [[Masai giraffe]] kill]] Giraffes have high adult survival probability,<ref name=":0">{{Cite book|doi=10.1016/B978-0-12-409548-9.09721-9|title=Reference Module in Earth Systems and Environmental Sciences|last1=Lee|first1=D. E.|last2=Strauss|first2=M. K. L. |date=2016 |publisher=Elsevier |isbn=9780124095489}}</ref> and an unusually long lifespan compared to other ruminants, up to 38 years.<ref>{{Cite journal |author1=Müller, D.W. |author2=Zerbe, P. |author3=Codron, D. |author4=Clauss, M. |author5=Hatt, J.M. |s2cid=10687135 |year=2011 |title=A long life among ruminants: giraffids and other special cases|journal=Schweizer Archiv für Tierheilkunde |volume=153 |issue=11 |pages=515–519 |pmid=22045457 |doi=10.1024/0036-7281/a000263}}</ref> Adult female survival is significantly correlated with the number of social associations.<ref>{{Cite journal|last1=Bond|first1=M. L.|last2=Lee|first2=D. E.|last3=Farine|first3=D. R. |last4=Ozgul |first4=A. |last5=König|first5=B. |date=2021 |title=Sociability increases survival of adult female giraffes |journal=Proceedings of the Royal Society B: Biological Sciences |volume=288 |issue=1944 |pages=20202770 |doi=10.1098/rspb.2020.2770 |pmid=33563118 |pmc=7893237 |doi-access=free}}</ref> Because of their size, eyesight and powerful kicks, adult giraffes are mostly safe from predation,<ref name=estes/> with [[lion]]s being their only major threats.<ref name=Dagg2014/>{{rp|55}} Calves are much more vulnerable than adults and are also preyed on by [[leopard]]s, [[spotted hyena]]s and [[African wild dog|wild dogs]].<ref name="Prothero 2003"/> A quarter to a half of giraffe calves reach adulthood.<ref name=":0" /><ref>{{Cite journal |last1=Lee|first1=D. E.|last2=Bond|first2=M. L. |last3=Kissui |first3=B. M. |last4=Kiwango|first4=Y. A. |last5=Bolger|first5=T. |date=2016 |title=Spatial variation in giraffe demography: a test of 2 paradigms |journal=[[Journal of Mammalogy]] |pages=1015–1025 |doi=10.1093/jmammal/gyw086 |volume=97 |issue=4 |s2cid=87117946 |doi-access=free}}</ref> Calf survival varies according to the season of birth, with calves born during the [[dry season]] having higher survival rates.<ref>{{Cite journal|last1=Lee|first1=D. E. |last2=Bond|first2=M. L. |last3=Bolger|first3=D. T. |date=2017 |title=Season of birth affects juvenile survival of giraffe |journal=[[Population Ecology]] |volume=59 |issue=1|pages=45–54 |doi=10.1007/s10144-017-0571-8 |bibcode=2017PopEc..59...45L |s2cid=7611046 |url-status=live |access-date=22 August 2023 |url=https://scholarsphere.psu.edu/resources/d7aaa4fd-9365-431c-8a9c-824e1f54d185|archive-date=22 September 2023 |archive-url=https://web.archive.org/web/20230922074127/https://scholarsphere.psu.edu/resources/d7aaa4fd-9365-431c-8a9c-824e1f54d185}}</ref> The local, seasonal presence of large herds of migratory [[wildebeest]]s and [[zebra]]s reduces predation pressure on giraffe calves and increases their survival probability.<ref>{{Cite journal|last1=Lee|first1=D. E.|last2=Kissui|first2=B. M. |last3=Kiwango|first3=Y. A. |last4=Bond |first4=M. L.|year=2016|title=Migratory herds of wildebeests and zebras indirectly affect calf survival of giraffes |journal=[[Ecology and Evolution]] |volume=6 |issue=23 |pages=8402–8411 |doi=10.1002/ece3.2561 |pmid=28031792 |pmc=5167056|bibcode=2016EcoEv...6.8402L }}</ref> In turn, it has been suggested that other ungulates may benefit from associating with giraffes, as their height allows them to spot predators from further away. Zebras were found to assess predation risk by watching giraffes and spend less time looking around when giraffes are present.<ref>{{cite journal|author=Schmitt, M. H. |author2=Stears, K.|author3=Shrader, A. M. |year=2016 |title=Zebra reduce predation risk in mixed-species herds by eavesdropping on cues from giraffe |journal=Behavioral Ecology |volume=27 |issue=4|pages=1073–1077 |doi=10.1093/beheco/arw015 |doi-access=free}}</ref> [[File:Giraffe Oxpeckers Lupande Zambia Jul23 A7R 06194.jpg|thumb|upright|left|Red-billed oxpeckers on a giraffe, Zambia]] Some parasites feed on giraffes. They are often [[host (biology)|hosts]] for [[tick]]s, especially in the area around the genitals, which have thinner skin than other areas. Tick species that commonly feed on giraffes are those of genera ''[[Hyalomma]]'', ''[[Amblyomma]]'' and ''[[Rhipicephalus]]''. [[Red-billed Oxpecker|Red-billed]] and [[yellow-billed oxpecker]]s clean giraffes of ticks and alert them to danger. Giraffes host numerous species of internal parasites and are susceptible to various diseases. They were victims of the (now eradicated) viral illness [[rinderpest]].<ref name=Dagg1971/> Giraffes can also suffer from a skin disorder, which comes in the form of wrinkles, [[Cutaneous condition#Lesions|lesions]] or raw [[skin fissure|fissures]]. As much as 79% of giraffes have symptoms of the disease in [[Ruaha National Park]], but it did not cause mortality in [[Tarangire National Park|Tarangire]] and is less prevalent in areas with fertile soils.<ref>{{Cite journal |last1=Lee|first1=D. E.|last2=Bond|first2=M. L.|date=2016 |title=The occurrence and prevalence of Giraffe skin disease in protected areas of northern Tanzania |journal=[[Journal of Wildlife Diseases]] |volume=52|issue=3|pages=753–755 |doi=10.7589/2015-09-247 |pmid=27310168|s2cid=10736316}}</ref><ref>{{Cite journal|last1=Bond|first1=M. L. |last2=Strauss |first2=M. K. L. |last3=Lee|first3=D. E.|date=2016 |title=Soil Correlates and Mortality from Giraffe Skin Disease in Tanzania |journal=Journal of Wildlife Diseases |doi=10.7589/2016-02-047|pmid=27529292 |volume=52|issue=4|pages=953–958 |s2cid=46776142}}</ref><ref>{{Cite journal |last1=Muneza |first1=A. B. |last2=Montgomery|first2=R. A.|last3=Fennessy|first3=J. T.|last4=Dickman|first4=A. J.|last5=Roloff|first5=G. J. |last6=Macdonald |first6=D. W.|date=2016 |title=Regional variation of the manifestation, prevalence, and severity of giraffe skin disease: A review of an emerging disease in wild and captive giraffe populations |journal=[[Biological Conservation]] |volume=198|pages=145–156 |doi=10.1016/j.biocon.2016.04.014|bibcode=2016BCons.198..145M }}</ref> {{-}} ==Human relations== ===Cultural significance=== With its lanky build and spotted coat, the giraffe has been a source of fascination throughout human history, and its image is widespread in culture. It has represented flexibility, far-sightedness, femininity, fragility, passivity, grace, beauty and the continent of Africa itself.<ref name=Williams/>{{rp|7, 116}} [[File:Giraffe cave art.jpg|thumb|alt=Photograph of a giraffe painted on a rock face|[[San rock art]] in Namibia depicting a giraffe]] Giraffes were depicted in art throughout the African continent,.<ref name=Williams>{{Cite book|author=Williams, E.|year=2011|title=Giraffe|publisher=[[Reaktion Books]]|isbn=978-1-86189-764-0}}</ref>{{rp|45–47}} The [[Kiffian culture|Kiffians]] were responsible for a life-size rock engraving of two giraffes, dated 8,000 years ago, that has been called the "world's largest rock art petroglyph".<ref name=Williams/>{{rp|45}}<ref>{{cite web|url=http://www.bradshawfoundation.com/giraffe/|title=The Dabous Giraffe rock art petrograph|publisher=The [[Bradshaw Foundation]]|access-date=6 November 2011|archive-date=7 November 2011|archive-url=https://web.archive.org/web/20111107203318/http://www.bradshawfoundation.com/giraffe/|url-status=live}}</ref> Judging from evidence including incised clay pots, archaeologists now believe that, in the ancient [[Kingdom of Kush|Kushite]] societies located in [[Nubia]], or what is now northern Sudan, giraffes may have featured in popular religion and women's religion, though not in elite or royal religion, and may have also had a connection to beliefs about the sun.<ref>{{Cite journal |last=Kilroe |first=Loretta |date=2023-12-18 |title=Giants of the Sands: The Giraffe and its Place in Symbolic Vocabulary in the Kingdom of Kush, Sudan |url=https://brill.com/view/journals/jaa/aop/article-10.1163-21915784-bja10032/article-10.1163-21915784-bja10032.xml |journal=Journal of African Archaeology |pages=1–20 |doi=10.1163/21915784-bja10032 |issn=1612-1651}}</ref> The [[Tugen people]] of modern Kenya used the giraffe to depict their god Mda.<ref>{{cite book|author=Shorrocks, B.|year=2016|url=https://books.google.com/books?id=747RDAAAQBAJ|title=The Giraffe: Biology, Ecology, Evolution and Behaviour|publisher=[[Wiley (publisher)|Wiley]]|page=3|isbn=9781118587478}}</ref> The [[Ancient Egypt|Egyptians]] gave the giraffe its own [[Egyptian hieroglyphs|hieroglyph]]; 'sr' in [[Old Egyptian language|Old Egyptian]] and 'mmy' in later periods.<ref name=Williams/>{{rp|49}} How the giraffe got its height has been the subject of various [[Culture of Africa#Folklore and religion|African folktales]].<ref name=sim1996/> Giraffes have a presence in modern [[Western culture]]. [[Salvador Dalí]] depicted them with burning manes in some surrealist paintings. Dali considered the giraffe to be a masculine symbol. A flaming giraffe was meant to be a "masculine cosmic apocalyptic monster".<ref name=Williams/>{{rp|123}} Several children's books feature the giraffe, including David A. Ufer's ''The Giraffe Who Was Afraid of Heights'', [[Giles Andreae]]'s ''Giraffes Can't Dance'' and [[Roald Dahl]]'s ''[[The Giraffe and the Pelly and Me]]''. Giraffes have appeared in animated films as minor characters in [[Disney]]'s ''[[Dumbo]]'' and ''[[The Lion King]]'', and in more prominent roles in ''[[The Wild]]'' and the ''[[Madagascar (franchise)|Madagascar]]'' films. [[Sophie the Giraffe]] has been a popular [[teether]] since 1961. Another famous fictional giraffe is the Toys "R" Us mascot [[Toys "R" Us#Mascot|Geoffrey the Giraffe]].<ref name=Williams/>{{rp|127}} The giraffe has also been used for some scientific experiments and discoveries. Scientists have used the properties of giraffe skin as a model for [[space suit|astronaut]] and [[flight suit|fighter pilot suits]] because the people in these professions are in danger of passing out if blood rushes to their legs.<ref name=anatomy/>{{rp|76}} Computer scientists have modeled the coat patterns of several subspecies using [[reaction–diffusion]] mechanisms.<ref>{{Cite book |author1=Walter, M. |author2=Fournier, A. |author3=Menevaux, D. |title=Proceedings of the 28th annual conference on Computer graphics and interactive techniques |chapter=Integrating shape and pattern in mammalian models |year=2001 |pages=[https://archive.org/details/siggraph2001conf00fium/page/317 317–26] |doi=10.1145/383259.383294 |chapter-url=https://archive.org/details/siggraph2001conf00fium/page/317 |isbn=978-1-58113-374-5 |citeseerx=10.1.1.10.7622 |s2cid=13488215 |access-date=16 November 2011 |url-status=dead |archive-url=https://web.archive.org/web/20150923224559/http://www.csun.edu/~renzo/GraphicsResources/Articles/walter.pdf |archive-date=23 September 2015 }}</ref> The [[constellation]] of [[Camelopardalis]], introduced in the 17th century, depicts a giraffe.<ref name=Williams/>{{rp|119–20}} The [[Tswana people]] of Botswana traditionally see the constellation [[Crux]] as two giraffes—[[Acrux]] and [[Beta Crucis|Mimosa]] forming a male, and [[Gacrux]] and [[Delta Crucis]] forming the female.<ref>{{cite journal|author=Clegg, A.|year=1986|title=Some Aspects of Tswana Cosmology|journal=Botswana Notes and Records|volume=18|pages=33–37|jstor=40979758 }}</ref> [[File:Tribute Giraffe with Attendant.jpg|thumb|right|upright|alt=Photograph of a painting of a giraffe and a man holding its leash|Painting of a giraffe imported to China during the [[Ming dynasty]]]] ===Captivity=== The Egyptians were among the earliest people to keep giraffes in captivity and shipped them around the Mediterranean.<ref name=Williams/>{{rp|48–49}} The giraffe was among the many animals collected and [[Mosaic Fragment with Man Leading a Giraffe (Art Institute of Chicago)|displayed]] by the Romans. The first one in Rome was brought in by [[Julius Caesar]] in 46 BC.<ref name=Williams/>{{rp|52}} With the [[fall of the Western Roman Empire]], the housing of giraffes in Europe declined.<ref name=Williams/>{{rp|54}} During the [[Middle Ages]], giraffes were known to Europeans through contact with the Arabs, who revered the giraffe for its peculiar appearance.<ref name="Prothero 2003">{{Cite book|author1=Prothero, D. R. |author2=Schoch, R. M. |year=2003|title=Horns, Tusks, and Flippers: The Evolution of Hoofed Mammals|publisher=[[Johns Hopkins University Press]]|pages=67–72|isbn=978-0-8018-7135-1}}</ref> Individual captive giraffes were given celebrity status throughout history. In 1414, a giraffe from [[Malindi]] was [[Treasure voyages|taken to China]] by explorer [[Zheng He]] and placed in a [[Ming dynasty]] zoo. The animal was a source of fascination for the Chinese people, who associated it with the mythical [[Qilin]].<ref name=Williams/>{{rp|56}} The [[Medici giraffe]] was a giraffe presented to [[Lorenzo de' Medici]] in 1486. It caused a great stir on its arrival in [[Florence]].<ref>{{Cite journal|url=https://archive.org/download/ErikRingmaraudienceForAGiraffeEuropeanExceptionalismAndTheQuest_744/ErikRingmarAudienceForAGiraffe.pdf |archive-url=https://web.archive.org/web/20080910082557/http://www.archive.org/download/ErikRingmaraudienceForAGiraffeEuropeanExceptionalismAndTheQuest_744/ErikRingmarAudienceForAGiraffe.pdf |archive-date=10 September 2008 |url-status=live |title=Audience for a Giraffe: European Expansionism and the Quest for the Exotic|author= Ringmar, E.|journal= [[Journal of World History]]| year = 2006| volume = 17| issue = 4| pages = 353–97 |jstor = 20079397|doi=10.1353/jwh.2006.0060|s2cid=143808549}}</ref> [[Zarafa (giraffe)|Zarafa]], another famous giraffe, was brought from [[History of Egypt under the Muhammad Ali dynasty|Egypt]] to Paris in the early 19th century as a gift for [[Charles X of France]]. A sensation, the giraffe was the subject of numerous memorabilia or "giraffanalia".<ref name=Williams/>{{rp|81}} Giraffes have become popular attractions in modern [[zoo]]s, though keeping them is difficult as they prefer large areas and need to eat large amounts of browse. Captive giraffes in North America and Europe appear to have a higher mortality rate than in the wild, the most common causes being poor husbandry, nutrition, and management.<ref name=Dagg2014/>{{rp|153}} Giraffes in zoos display [[Stereotypy (non-human)|stereotypical behaviours]], particularly the licking of inanimate objects and pacing.<ref name=Dagg2014/>{{rp|164}} Zookeepers may offer various activities to stimulate giraffes, including training them to take food from visitors.<ref name=Dagg2014/>{{rp|167, 176}} [[Stable]]s for giraffes are built particularly high to accommodate their height.<ref name=Dagg2014/>{{rp|183}} ===Exploitation=== Giraffes were probably common targets for hunters throughout Africa.<ref name="Kingdon1988" />{{rp|337}} Different parts of their bodies were used for different purposes.<ref name="Dagg1971" /> Their meat was used for food. The tail hairs were [[flyswatter]]s, bracelets, necklaces, and threads. Shields, sandals, and drums were made using the skin, and the strings of musical instruments were from the tendons.<ref name="Dagg1971" /><ref name="Kingdon1988" />{{rp|337}} In [[Buganda]], the smoke of burning giraffe skin was traditionally used to treat nosebleeds.<ref name="Kingdon1988" />{{rp|337}} The [[Humur|Humr]] people of [[Kordofan]] consume the drink Umm Nyolokh, which is prepared from the [[liver]] and [[bone marrow]] of giraffes. [[Richard Rudgley]] hypothesised that Umm Nyolokh might contain [[N,N-Dimethyltryptamine|DMT]].<ref>Rudgley, Richard ''The Encyclopedia of Psychoactive Substances'', pub. Abacus 1998 {{ISBN|0 349 11127 8}} pps. 20–21.</ref> The drink is said to cause hallucinations of giraffes, believed to be the giraffes' ghosts, by the [[Humr]].<ref>{{cite journal|author=Ian Cunnison|title= Giraffe hunting among the Humr tribe|journal=[[Sudan Notes and Records]]|volume=39|year=1958}}</ref> ==Conservation status== In 2016, giraffes were assessed as [[Vulnerable species|Vulnerable]] from a conservation perspective by the IUCN.<ref name=iucn/> In 1985, it was estimated there were 155,000 giraffes in the wild. This declined to over 140,000 in 1999.<ref name="wildstatus">{{cite web|url=http://www.giraffeconservation.org/giraffe_facts.php?pgid=40|title=Giraffe – The Facts: Current giraffe status?|publisher=Giraffe Conservation Foundation|access-date=21 December 2010|archive-url=https://web.archive.org/web/20160319183600/http://www.giraffeconservation.org/giraffe_facts.php?pgid=40|archive-date=19 March 2016|url-status=dead}}</ref> Estimates as of 2016 indicate there are approximately 97,500 members of ''Giraffa'' in the wild.<ref>{{cite news|author=Matt McGrath|date=8 December 2016|title=Giraffes facing 'silent extinction' as population plunges|publisher=[[BBC News]]|url=https://www.bbc.co.uk/news/science-environment-38240760|access-date=8 December 2016|archive-date=21 May 2019|archive-url=https://web.archive.org/web/20190521190734/https://www.bbc.co.uk/news/science-environment-38240760|url-status=live}}</ref><ref>{{cite web|title=New bird species and giraffe under threat – IUCN Red List|date=8 December 2016|url=https://www.iucn.org/news/new-bird-species-and-giraffe-under-threat-%E2%80%93-iucn-red-list|access-date=8 December 2016|archive-date=5 March 2017|archive-url=https://web.archive.org/web/20170305022533/https://www.iucn.org/news/new-bird-species-and-giraffe-under-threat-%E2%80%93-iucn-red-list|url-status=live}}</ref> The Masai and reticulated subspecies are [[Endangered species|endangered]],<ref>{{cite iucn|author=Bolger, D.|author2=Ogutu, J.|author3=Strauss, M.|author4=Lee, D.|author5=Muneza, A.|author6=Fennessy, J.|author7=Brown, D.|year=2019|title=Masai Giraffe|doi=10.2305/IUCN.UK.2019-1.RLTS.T88421036A88421121.en}}</ref><ref>{{cite iucn|title=Reticulated Giraffe|author=Muneza, A.|author2=Doherty, J. B.|author3=Hussein Ali, A.|author4=Fennessy, J.|author5=Marais, A.|author6=O'Connor, D.|author7=Wube, T.|year=2018|doi=10.2305/IUCN.UK.2018-2.RLTS.T88420717A88420720.en}}</ref> and the Rothschild subspecies is [[Near-threatened species|near threatened]].<ref name="IUCNrothschildi" /> The Nubian subspecies is [[critically endangered]].<ref name="IUCNnubian">{{cite iucn|author=Wube, T.|author2=Doherty, J. B.|author3=Fennessy, J.|author4=Marais, A.|year=2018|title=''Giraffa camelopardalis'' ssp. ''camelopardalis''|doi=10.2305/IUCN.UK.2018-2.RLTS.T88420707A88420710.en}}</ref> [[File:Giraffe koure niger 2006.jpg|thumb|right|alt=Photograph of an endangered West African giraffe|Endangered [[West African giraffe]] near [[Kouré|Koure]], Niger]] The primary causes for giraffe population declines are [[habitat destruction|habitat loss]] and direct killing for [[bushmeat]] markets. Giraffes have been [[Local extinction|extirpated]] from much of their historic range, including [[Eritrea]], [[Guinea]], [[Mauritania]] and [[Senegal]].<ref name="iucn" /> They may also have disappeared from [[Angola]], [[Mali]], and [[Nigeria]], but have been introduced to [[Rwanda]] and [[Eswatini]].<ref name=iucn/><ref name=IUCNnubian/> {{As of|2010}}, there were more than 1,600 in captivity at [[Species360]]-registered zoos.<ref name="ISIS" /> Habitat destruction has hurt the giraffe. In the [[Sahel]], the need for firewood and grazing room for livestock has led to [[deforestation]]. Normally, giraffes can coexist with livestock, since they avoid direct competition by feeding above them.<ref name="MacDonald" /> In 2017, severe droughts in northern Kenya led to increased tensions over land and the killing of wildlife by herders, with giraffe populations being particularly hit.<ref>{{cite journal|author=Qiu, Jane|author-link=Jane Qiu|title=Surge in wildlife killings is wiping out giraffes|journal=[[Science (journal)|Science]]|url=https://www.science.org/content/article/surge-wildlife-killings-wiping-out-giraffes?et_rid=305708592&et_cid=1399786|date=2017|doi=10.1126/science.aan7000|access-date=30 June 2022|archive-date=9 February 2023|archive-url=https://web.archive.org/web/20230209060153/https://www.science.org/content/article/surge-wildlife-killings-wiping-out-giraffes?et_rid=305708592&et_cid=1399786|url-status=live}}</ref> Protected areas like national parks provide important habitat and anti-[[poaching]] protection to giraffe populations.<ref name=iucn/> Community-based conservation efforts outside national parks are also effective at protecting giraffes and their habitats.<ref>{{Cite journal|last=Lee|first=Derek E.|date=2018|title=Evaluating conservation effectiveness in a Tanzanian community wildlife management area|url=https://wildlife.onlinelibrary.wiley.com/doi/abs/10.1002/jwmg.21549|journal=The Journal of Wildlife Management|language=en|volume=82|issue=8|pages=1767–1774|doi=10.1002/jwmg.21549|bibcode=2018JWMan..82.1767L |s2cid=91251633|issn=1937-2817|access-date=16 November 2020|archive-date=26 October 2020|archive-url=https://web.archive.org/web/20201026202649/https://wildlife.onlinelibrary.wiley.com/doi/abs/10.1002/jwmg.21549|url-status=live}}</ref><ref>{{Cite journal|last1=Lee|first1=Derek E|last2=Bond|first2=Monica L. |date=2018 |title=Quantifying the ecological success of a community-based wildlife conservation area in Tanzania|url= |journal=[[Journal of Mammalogy]]|volume=99|issue=2|pages=459–464|doi=10.1093/jmammal/gyy014|issn=0022-2372|pmc=5965405|pmid=29867255}}</ref> Private game reserves have contributed to the preservation of giraffe populations in eastern and southern Africa.<ref name="MacDonald" /> The giraffe is a protected species in most of its range. It is the [[national animal]] of Tanzania,<ref>{{Cite book|author=Knappert, J|title=East Africa: Kenya, Tanzania & Uganda|publisher=Vikas Publishing House|year=1987|isbn=978-0-7069-2822-8|page=57|author-link=Jan Knappert}}</ref> and is protected by law,<ref name="FoleyFoley2014">{{cite book|author1=Charles Foley|url=https://books.google.com/books?id=dt6QAwAAQBAJ&pg=PA179|title=A Field Guide to the Larger Mammals of Tanzania|author2=Lara Foley|author3=Alex Lobora|author4=Daniela De Luca|author5=Maurus Msuha|author6=Tim R. B. Davenport|author7=Sarah M. Durant|date=2014|publisher=[[Princeton University Press]]|isbn=978-1-4008-5280-2|page=179|access-date=13 July 2016|archive-date=22 September 2023|archive-url=https://web.archive.org/web/20230922074122/https://books.google.com/books?id=dt6QAwAAQBAJ&pg=PA179|url-status=live}}</ref> and unauthorised killing can result in imprisonment.<ref>{{cite web|title=National Symbols: National Animal|url=http://tanzania.go.tz/home/pages/260|url-status=dead|archive-url=https://web.archive.org/web/20150118195350/http://tanzania.go.tz/home/pages/260|archive-date=18 January 2015|access-date=14 January 2015|website=tanzania.go.tz|publisher=Tanzania Government Portal}}</ref> The UN-backed Convention of Migratory Species selected giraffes for protection in 2017.<ref>{{cite news|date=28 October 2017|title=Chimpanzees among 33 breeds selected for special protection|publisher=[[BBC News]]|url=https://www.bbc.com/news/world-asia-41791842|access-date=30 October 2017|archive-date=29 October 2017|archive-url=https://web.archive.org/web/20171029055127/http://www.bbc.com/news/world-asia-41791842|url-status=live}}</ref> In 2019, giraffes were listed under Appendix II of the Convention on International Trade in Endangered Species ([[CITES]]), which means international trade including in parts/derivatives is regulated.<ref>{{Cite web|title=Good News for Giraffes at CITES CoP18 > Newsroom|url=https://newsroom.wcs.org/News-Releases/articleType/ArticleView/articleId/12930/Good-News-for-Giraffes-at-CITES-CoP18.aspx|access-date=16 November 2020|website=newsroom.wcs.org|archive-date=27 September 2020|archive-url=https://web.archive.org/web/20200927225103/https://newsroom.wcs.org/News-Releases/articleType/ArticleView/articleId/12930/Good-News-for-Giraffes-at-CITES-CoP18.aspx|url-status=live}}</ref> [[Translocation (Wildlife conservation)|Translocations]] are sometimes used to augment or re-establish diminished or extirpated populations, but these activities are risky and difficult to undertake using the best practices of extensive pre- and post-translocation studies and ensuring a viable founding population.<ref>{{Cite journal|last1=Muller|first1=Zoe|last2=Lee|first2=Derek E.|last3=Scheijen|first3=Ciska P. J.|last4=Strauss|first4=Megan K. L.|last5=Carter|first5=Kerryn D.|last6=Deacon|first6=Francois|date=2020|title=Giraffe translocations: A review and discussion of considerations|journal=[[African Journal of Ecology]] |volume=58 |issue=2 |pages=159–171 |doi=10.1111/aje.12727 |doi-access=free|bibcode=2020AfJEc..58..159M }}</ref><ref>{{Cite journal |last1=Lee |first1=De |last2=Fienieg |first2=E. |last3=Van Oosterhout |first3=C. |last4=Muller |first4=Z. |last5=Strauss |first5=M. |last6=Carter |first6=Kd. |last7=Scheijen |first7=Cpj |last8=Deacon |first8=F. |date=27 February 2020 |title=Giraffe translocation population viability analysis |url=https://www.int-res.com/abstracts/esr/v41/p245-252/ |journal=Endangered Species Research |volume=41 |pages=245–252 |doi=10.3354/esr01022 |doi-access=free |access-date=16 November 2020 |archive-date=24 September 2020 |archive-url=https://web.archive.org/web/20200924020906/https://www.int-res.com/abstracts/esr/v41/p245-252/ |url-status=live }}</ref> Aerial survey is the most common method of monitoring giraffe population trends in the vast roadless tracts of African landscapes, but aerial methods are known to undercount giraffes. Ground-based survey methods are more accurate and can be used in conjunction with aerial surveys to make accurate estimates of population sizes and trends.<ref name=":4">{{Cite journal|last1=Lee|first1=Derek E.|last2=Bond|first2=Monica L.|date=2016|title=Precision, accuracy, and costs of survey methods for giraffe ''Giraffa camelopardalis''|journal=[[Journal of Mammalogy]] |volume=97 |issue=3|pages=940–948|doi=10.1093/jmammal/gyw025 |s2cid=87384776 |doi-access=free}}</ref> ==See also== * [[Fauna of Africa]] * [[Giraffe Centre]] * [[Giraffe Manor]] - hotel in Nairobi with giraffes == References == {{Reflist}} ==External links== {{Commons|Giraffa}} * {{official website|https://giraffeconservation.org/}} of the Giraffe Conservation Foundation {{Artiodactyla|R.}} {{Taxonbar|from=Q862089}} {{Authority control}} [[Category:Giraffes| ]] [[Category:Articles containing video clips]] [[Category:Fauna of Sub-Saharan Africa]] [[Category:Herbivorous mammals]] [[Category:Mammal genera]] [[Category:Mammals of Africa]] [[Category:National symbols of Tanzania]] [[Category:Vulnerable animals]] [[Category:Vulnerable biota of Africa]]
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