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長頸鹿
化石时期:11.61–0 Ma
中新世 to 全新世
Giraffe Mikumi National Park.jpg
位于坦桑尼亚米庫米國家公園的一只马赛长颈鹿(G. c. tippelskirchi)
保护状况
科学分类 编辑
界: 动物界 Animalia
门: 脊索动物门 Chordata
纲: 哺乳纲 Mammalia
目: 偶蹄目 Artiodactyla
科: 长颈鹿科 Giraffidae
属: 长颈鹿属 Giraffa
马蒂兰·雅克·布里松, 1772
模式种
努比亚长颈鹿

G. c. camelopardalis
(卡尔·林奈, 自然系统第十版)

物种

长颈鹿

Giraffa camelopardalis distribution2.png
现存长颈鹿亚种分布

长颈鹿 (Giraffa) 是非洲偶蹄目哺乳动物的一个。 该属目前仅由一个物种组成,但已经有七种已经灭绝的长颈鹿的化石被发现,目前仅存的努比亚长颈鹿(Giraffa camelopardalis)即为该物种的模式種,而基于线粒体DNA和核DNA分析和形态学比较,现存的长颈鹿可以被生物分类为一至八个种, 但是IUCN目前只承认长颈鹿为单一个物种(但拥有九个亚种)。长颈鹿也是最高的陆生动物和最大的反芻动物。

长颈鹿的显著特征在于长脖子、长腿、类角的听小骨、以及它们特有的毛发形态。在科 (生物)划分上,它与最近的远亲㺢㹢狓共同列为 长颈鹿科. 长颈鹿的分布北起乍得,南达南非, 西到尼日尔,东至索马里,常居于疏林草原或是疏林。它们以食用含羞草亚科为主的木本植物的树叶、水果以及花朵为生,此类植物生长于其他草食动物够不到的高度。它们也可能成为斑鬣狗非洲野犬的猎物。 长颈鹿为群居动物,与配偶子女为群,或与成年雄性同类为群。雄鹿划分社会等级通过以脖子为武器的角斗,其中,优胜者得以与雌鹿交配。雌鹿的唯一责任为养育幼崽。

长颈鹿以其独特的外观和行为成为了从古至今的各种文化的组成元素,经常出现在绘画,书籍和漫画之中。國際自然保護聯盟认定长颈鹿为易危物種,在部分地区已经局部地區滅絕。 长颈鹿是众多國家公園和主题公园的常见动物,2016年的估计表明,野生长颈鹿大约有97,500只,被饲养的约有1,144只。

命名编辑

長頸鹿的名字最早起源於阿拉伯语詞語zarāfah (زرافة),[2] 可能取自與索馬里語中的動物geri.[3] 阿拉伯語中這個詞的意思是“走得快的東西”。[4]中古英语中長頸鹿有多種拼法,例如jarraf, ziraphgerfauntz[2] 意大利语中的giraffa出現與20世紀90年代。[2] 現代英語中的長頸鹿在1600年左右出現,源自法语中的girafe[2] 驼豹(Camelopard) 是古英语中对长颈鹿的称呼,这个词起源于古希臘語,用于代指拥有骆驼一样的身体和一样的花纹的动物。[5][6]

中国典籍关于长颈鹿的记载,最早出自晋代李石所著《续博物志》,记录非洲索马里沿岸拨拔力古国出产的异兽,身高一丈余颈长九尺。宋代赵汝适著《诸蕃志》中称非洲长颈鹿为徂蜡:“状如驼驰,而大如,色黄,前脚高五尺,后低三尺,头高向上。”[7]明朝永乐十二年(1414),时值郑和第四次出海,榜葛剌国(今孟加拉)使臣将一只长颈鹿进贡给明成祖,以“麒麟”为名,据说乃东非土语中的长颈鹿名称giri之音译[8][9];在中国传说中,麒麟为祥瑞之兽,明成祖便将其视为祥瑞之兆,命人撰文并绘图记录,以此标榜自己施政的伟大。存世的图像记录仅有传沈度所绘的《麒麟图》。[9]该典故流传甚广。长颈鹿在日語韓語中的名称皆源于麒麟(日语:キリン)[10],在閩南話中也称为「麒麟鹿」。

分类编辑

将现存的长颈鹿曾被卡尔·林奈于1758年根据二名法归入鹿屬 。但摩顿·思伦·布兰尼奇(Morten Thrane Brünnich)于1772年重㺢㹢狓将其分类为长颈鹿属[11] 种名camelopardalis 来源于拉丁语.[12]

演化编辑

反刍亚目
鼷鹿下目英语Tragulina

鼷鹿科  

有角下目

叉角羚科  

长颈鹿科  

鹿科  

牛科  

麝科  

Cladogram英语Cladogram based on a 2003 study by Hassanin and Douzery.[13][14]

长颈鹿是偶蹄目长颈鹿科仅有的两个现存的属中的一个,剩下的一个是㺢㹢狓. 该科曾经一度分布非常广泛,有超过10个属。他们关系最近的已知的近亲是已经灭绝的类鹿的梯角鹿科 。他们和叉角羚科一样(现仅存的物种只有叉角羚)都从属于长颈鹿总科 。这些动物很可能是从已经灭绝的古鹿科演化过来,而古鹿科很可能是鹿科的祖先[15]

长颈鹿脖子的延长似乎开始于长颈鹿谱系的早期。与远古时期的近亲相比,现在的长颈鹿接近颅骨的椎骨较早延长,随后椎骨进一步向下延长。[16] Canthumeryx 是早期长颈鹿的一个祖先,该物种的历史可以追溯到2500万至2000万年前,并且在利比亚找到了他们的化石。该物种是中等大小,体型苗条,形态类似羚羊。始长颈鹿属出现在1500万年前的印度次大陆,类似于霍加或小长颈鹿,有较长的脖子和类似的听小骨。[15] 始长颈鹿属可能与与大量的长颈鹿比如:西瓦鹿布拉马兽共享同一个进化分支。[16]

 
欧亚大陆和非洲的中新世和上新世的灭绝属Samotherium(中)生物与㺢㹢狓(下)和现代长颈鹿相比的图片。 Samotherium属的生物的解剖学特征明显表现出向长颈鹿状颈部的过渡。[17]

长颈鹿科於 1400 萬年前出現,並且居住於歐亞非大陸全境,代表性的有古長頸鹿山西獸屬以及薩摩獸英语Samotherium 。These animals had bare ossicones and small cranial sinuses and were longer with broader skulls.[15][16] Paleotragus resembled the okapi and may have been its ancestor.[15] Others find that the okapi lineage diverged earlier, before Giraffokeryx.[16] Samotherium was a particularly important 過渡化石 in the giraffe lineage as its cervical vertebrae was intermediate in length and structure between a modern giraffe and an okapi, and was more vertical than the okapi's.[17] Bohlinia英语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.[15]

Bohlinia entered China and northern India in response to climate change. From there, the genus 长颈鹿 evolved and, around 7 mya, entered Africa.[18] Further climate changes caused the extinction of the Asian giraffes, while the African giraffes survived and radiated into several new species. Living giraffes appear to have arisen around 1 mya in eastern Africa during the 更新世.[15] Some biologists suggest the modern giraffes descended from G. jumae英语Giraffa jumae;[19] others find G. gracilis a more likely candidate.[15] G. jumae was larger and more heavily built while G. gracilis was smaller and more lightly built. The main driver for the evolution of the giraffes is believed to have been the changes from extensive forests to more open habitats, which began 8 mya.[15] During this time, tropical plants disappeared and were replaced by arid C4类二氧化碳固定s, and a dry savannah emerged across eastern and northern Africa and western India.[20][21] 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.[22] 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.[15]

In the early 19th century, 让-巴蒂斯特·拉马克 believed the giraffe's long neck was an "acquired characteristic", developed as generations of ancestral giraffes strove to reach the leaves of tall trees.[23] This theory was eventually rejected, and scientists now believe the giraffe's neck arose through Darwinian 自然选择—that ancestral giraffes with long necks thereby had a competitive feeding advantage (competing browsers hypothesis)[24] that better enabled them to survive and reproduce to pass on their genes.[23]

The giraffe 基因組 is around 29億 碱基对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 two species are equally distantly related to cattle, suggesting the giraffe's unique characteristics are not because of faster evolution. The 趨異演化 of giraffe and okapi lineages dates to around 11.5 mya. A small group of regulatory gene英语regulatory genes in the giraffe appear to be responsible for the animal's stature and associated circulatory adaptations.[25]

种及亚种编辑

 
"Approximate geographic ranges, fur patterns, and 系统发生学 relationships between some giraffe subspecies based on 线粒体DNA sequences. Colored dots on the map represent sampling localities. The phylogenetic tree is a 最大似然估计 系統發生樹 based on samples from 266 giraffes. Asterisks along branches correspond to 演化支 values of more than 90% bootstrap英语Resampling (statistics) support. Stars at branch tips identify 並系群 haplotype英语haplotypes found in Maasai and reticulated giraffes".[26]

世界自然保护联盟目前仅承认具有9个亚种的一个长颈鹿物种。[27][28] 在2001年,有人提出了将长颈鹿分为两种物种的分类建议。[29] A 2007 study on the genetics of Giraffa, suggested they were six species: the West African, Rothschild's, reticulated, Masai, Angolan, and South African giraffe.[26] The study deduced from genetic differences in nuclear英语nuclear DNA and 线粒体DNA (mtDNA) that giraffes from these populations are 生殖隔离 and rarely interbreed, though no natural obstacles block their mutual access. This includes adjacent populations of Rothschild's, reticulated, and Masai giraffes. The Masai giraffe was also suggested to consist of possibly two species separated by the 東非裂谷.[26]

Reticulated and Masai giraffes have the highest mtDNA diversity, which is consistent with giraffes originating in eastern Africa. Populations further north are more closely related to the former, while those to the south are more related to the latter. Giraffes appear to select mates of the same coat type, which are imprinted on them as calves.[26] The implications of these findings for the conservation of giraffes were summarised by David Brown, lead author of the study, who told 英國廣播公司新聞: "Lumping all giraffes into one species obscures the reality that some kinds of giraffe are on the brink. Some of these populations number only a few hundred individuals and need immediate protection."[30]

A 2011 study using detailed analyses of the morphology of giraffes, and application of the 物种, described eight species of living giraffes.[31] The eight species are: G. angolensis英语Angolan giraffe, G.antiquorum英语Kordofan giraffe, G. camelopardalis英语Nubian giraffe, G. giraffa英语South African giraffe, G. peralta英语West African giraffe, 网纹长颈鹿, G. thornicrofti英语Rhodesian giraffe, and G. tippelskirchi英语Masai giraffe.

A 2016 study also concluded that living giraffes consist of multiple species.[32] The researchers suggested the existence of four species, which have not exchanged genetic information between each other for 1 million to 2 million years. Those four species are the 努比亚长颈鹿 (G. camelopardalis), southern giraffe英语southern giraffe (G. giraffa), 网纹长颈鹿 (G. reticulata), and Masai giraffe英语Masai giraffe (G. tippelskirchi).[32] Since then, a response to this publication has been published, highlighting seven problems in data interpretation, and concludes "the conclusions should not be accepted unconditionally".[33]

据估计,野生长颈鹿个体约有90,000,目前有1,144只长颈鹿处于被饲养状态(动物园、主题公园)。[32][34]

还有七种已灭绝的长颈鹿物种,如下所示:

G. attica,也灭绝了, 该物种一度也被认为是长颈鹿属(Giraffa)下的一个,但1929年被重新分类为Bohlinia attica英语Bohlinia

长颈鹿的可能分类
一种[28][27] 四种[32]   八种[31] 说明 图片
长颈鹿 (G. camelopardalis)[28][27] 努比亚长颈鹿 (G. camelopardalis) 科尔多凡长颈鹿英语Kordofan giraffe (G. antiquorum)[35] 科尔多凡长颈鹿(G. c. antiquorum)分布于乍得南部,中非共和国喀麦隆北部以及刚果(金)东北部。之前喀麦隆的种群错误地包括了西非长颈鹿(G. c. peralta)。与努比亚长颈鹿相比,科尔多凡长颈鹿有更小也更不规则的斑点图案,它的斑覆盖肘关节以下以及腿的内侧区域。雄性个体的头部中央会长有一个肿块。人们相信大约有两千只科尔多凡长颈鹿生活在野外,但是科尔多凡长颈鹿和西非长颈鹿在动物园中的数量有相当大的混淆,在2007年所有之前宣称的生存于欧洲动物园中的西非长颈鹿被证明实际上都是科尔多凡长颈鹿(G. c. peralta),随着这一修正,生活在动物园中的科尔多凡长颈鹿数量达到了大约65只。而之前的亚种刚果长颈鹿则被认为是科尔多凡长颈鹿的一部分。  
努比亚长颈鹿(G. camelopardalis)包括罗氏长颈鹿,也被称作巴林戈长颈鹿乌干达长颈鹿 努比亚长颈鹿 (G. c. camelopardalis)分布于南苏丹埃塞俄比亚西南部,以及肯尼亚乌干达[28] 努比亚长颈鹿的栗色斑点有着被白线包围的清晰边界,同时身体下方缺少斑点,[36] 雄性个体头部中央的肿块尤为发达。[37]:51 人们认为大约有2150只个体生活在野外,其中大约1500只属于罗氏生态型。[28] 随着罗氏长颈鹿并入努比亚长颈鹿亚种,使得努比亚长颈鹿在动物园中相当常见,但是其最初的表型仍然十分稀有,其中一小群生活在阿联酋的艾恩野生动物园,[38] 2003年仅有14只。[39]

罗氏长颈鹿 (G. c. rothschildi)可能是努比亚长颈鹿的一个生态型,它们的分布范围包括一部分肯尼亚和乌干达,[28] 并不确定的它们是否也生活在苏丹南部。[40] 罗氏长颈鹿的大块深色斑点通常有着完整边界,也可能有锋利的边缘,深色斑点也会有淡色的辐射线或者条痕,这些斑点长在肘关节以下时并不常见,而且完全不会到达蹄部。这一生态型可能会长出五个“角”。[37]:53 人们相信大约有1500只罗氏长颈鹿生活在野外,[28] 并且有450只生活在动物园。[34] 依据大约于2016年九月进行的基因分析,努比亚长颈鹿和罗氏长颈鹿 (G. c. camelopardalis)是同种的。[32]

 
西非长颈鹿英语West African giraffe (G. peralta),[41][42] 也被称作尼日尔长颈鹿尼日利亚长颈鹿 西非长颈鹿(G. c. peralta)是尼日尔西南部的特有种[28] 它们的毛色比其他亚种更浅,[43]:322 红色裂片状的斑点可以长到肘关节以下。西非长颈鹿的角比起其它亚种更为竖直,雄性头部中央有一个较大的肿块。[37]:52–53 这一亚种是长颈鹿属内最为濒危的亚种,仅有大约400只生活在野外。[28] 之前分布于喀麦隆的长颈鹿被认为属于西非长颈鹿但实际上是科尔多凡长颈鹿(G. c. antiquorum)。[44] 这一错误也导致对于动物园中长颈鹿的一些混乱认知,但是在2007年已经确定所有在欧洲动物园的西非长颈鹿(G. c. peralta)均为科尔多凡长颈鹿(G. c. antiquorum)。2007年同样的研究也发现比起科尔多凡长颈鹿,西非长颈鹿与罗氏长颈鹿的关系更近,它们的祖先可能随着撒哈拉沙漠的扩大逐渐从东非迁徙到北非它们现在的栖息地。而乍得湖则成了在全新世(公元前5000年以前)分隔科尔多凡长颈鹿和西非长颈鹿的屏障。[44]
 
网纹长颈鹿 (G. reticulata),[45] 又称作索马里长颈鹿 网纹长颈鹿 (G. c. reticulata) 分布于肯尼亚东北部,埃塞俄比亚南部以及索马里[28] 它有独特的红棕色多边形斑点,并由网状的细白线分割开来。它们的斑点有时会长在肘关节以下,雄性头部中央长有肿块。[37]:53 据估计有8660只网纹长颈鹿生活在野外,[28] 根据国际物种信息系统的记录,超过450只网纹长颈鹿生活在动物园中。[34]  
南部长颈鹿(G. giraffa) 安哥拉长颈鹿(G. angolensis),也被称作纳米比亚长颈鹿 安哥拉长颈鹿(G. c. angolensis)分布于 纳米比亚东北部,赞比亚西南部,波札那辛巴威西部。[28] 一项于2009年开展的基因研究表明 納米比沙漠北部和埃托沙國家公園 的种群生成了独立的亚种。[46] 这种长颈鹿有着大块的棕色斑点,其边缘有缺口或是棱角。它们的斑点一直长到腿上,但是面部上半部分却没有。脖子和臀部的斑点往往相当小。 此种长颈鹿在耳朵上也有白色斑点。[37]:51 现估计有13000只安哥拉长颈鹿生活在野外[28] 大约20只生活在动物园里。[34]  
南非长颈鹿 (G. giraffa)[47] 也被称作好望角长颈鹿 南非长颈鹿(G. c. giraffa)分布于南非北部,博茨瓦纳南部,津巴布韦南部以及莫桑比克西南部。[28] 在茶色的底色之上,南非长颈鹿的深色斑点有时会呈现圆形,延伸到腿上后斑点会变小,雄性头部的肿块也相对较小。[37]:52 估计最多有31500只生活在野外[28] 并有45只生活在动物园。[34]  
马赛长颈鹿(G. tippelskirchi) 马赛长颈鹿(G. tippelskirchi),[48] 也被称作乞力马扎罗长颈鹿 马赛长颈鹿(G. c. tippelskirchi)分布于肯尼亚中部和南部以及坦桑尼亚.[28] 它们延伸到蹄部的斑点非常独特,呈不规则的锯齿状或星状。雄性头部中央通常长有肿块。[37]:54[49] 总共有32550只马赛长颈鹿生活在野外,[28] 并且有大约100只生活在动物园中。[34]  
索氏长颈鹿("G. thornicrofti", 以哈里·斯科特·索尼克罗夫特命名),[50] 也被称作卢安瓜长颈鹿罗得西亚长颈鹿 索氏长颈鹿(G. c. thornicrofti) 仅仅生活在赞比亚东部的卢安瓜河谷区域.[28] 它们的斑点呈锯齿或星状,有时会长满整个腿部,雄性个体的头上没有肿块。[37]:54 不超过550只索氏长颈鹿生活在野外,[28]且没有一只生活在动物园中。[34]  

外观和解剖学编辑

 
墨爾本動物園的一只长颈鹿头部近照。
 
长颈鹿的骨骼,展出于 Museum of Osteology英语Museum of Osteology, 奧克拉荷馬市
 
一只马赛长颈鹿的右后腿,摄于 圣迭戈动物园

成年长颈鹿高度约 4.3~5.7米(14.1~18.7英尺),其中,雄性长颈鹿的平均身高要比雌性长颈鹿更高。[51][52][53] 目前记录到的雄性长颈鹿最高达 5.88米(19.3英尺) 而最高的雌性长颈鹿达 5.17米(17.0英尺) 高。[51][54] 雄性长颈鹿的平均重量约为 1,192公斤(2,628磅),而雌性长颈鹿的平均重量约为 828公斤(1,825磅) 。[55] 目前记录到的雄性和雌性长颈鹿最大体重分别是1,930公斤(4,250磅) 和 1,180公斤(2,600磅)。[52][53]尽管长颈鹿的脖子和腿很长,但它身体的主体部分相对较短。[56]:66 一双突出的大眼睛位于头部两侧,凭借身高优势,长颈鹿拥有良好的全方位视觉。[57]:25 长颈鹿拥有色觉[57]:26 并且它们的听力和嗅觉也十分敏锐。[23] 长颈鹿可以利用肌肉控制鼻孔的开闭以防止沙尘和蚂蚁的进入。[57]:27

长颈鹿的舌头长约45厘米(18英寸)。[52][53] It is purplish-black in colour, perhaps to protect against sunburn, and is useful for grasping foliage, as well as for grooming and cleaning the animal's nose.[57]:27 The upper lip of the giraffe is also prehensile and useful when foraging and is covered in hair to protect against thorns. The tongue, and inside of the mouth are covered in 味蕾.[11]

日本埼玉縣东武动物公园英语Tobu Zoo的一对长颈鹿

The coat has dark blotches or patches (which can be orange, chestnut英语Chestnut (color), brown, or nearly black in colour[23]) separated by light hair (usually white or cream英语Cream (colour) in colour[23]). Male giraffes become darker as they age.[49] The coat pattern has been claimed to serve as 偽裝 (動物學) in the light and shade patterns of savannah woodlands.[50] While adult giraffes standing among trees and bushes are hard to see at even a few metres' distance, they actively move into the open to gain the best view of an approaching predator, obviating any benefit that camouflage might bring. Instead, the adults rely on their size and ability to defend themselves. However, camouflage appears to be important for calves, which spend a large part of the day in hiding, away from their mothers; further, over half of all calves die within a year, so predation is certainly important. It appears, therefore, that the spotted coat of the giraffe functions as camouflage for the young, while adults simply inherit this coloration as a by-product.[15] The skin underneath the dark areas may serve as windows for 體溫調節, being sites for complex blood vessel systems and large sweat glands.[58] Each individual giraffe has a unique coat pattern.[49]

长颈鹿的皮肤大部分是灰色的。[55] 皮肤的厚度允许长颈鹿穿过刺灌木丛而不受伤。[57]:34 The fur may serve as a chemical defence, as its parasite repellents give the animal a characteristic scent. At least 11 main 芳香性 chemicals are in the fur, although 吲哚 and 糞臭素 are responsible for most of the smell. Because the males have a stronger odour than the females, the odour may also have sexual function.[59] Along the animal's neck is a mane made of short, erect hairs.[11] The one-metre (3.3-ft) tail ends in a long, dark tuft of hair and is used as a defense against insects.[57]:36

颅骨和鹿角状突起(ossicones)编辑

长颈鹿两性都有突出的角状结构,称为鹿角状突起(ossicones) which are formed from ossified cartilage, covered in skin and fused to the skull at the 顶骨s.[49] Being 血管ized, the ossicones may have a role in thermoregulation,[58] and are also used in combat between males.[60] 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 end in knobs and tend to be bald on top.[49] Also, a median lump, which is more prominent in males, emerges at the front of the 颅骨.[11] Males develop deposits that form bumps on their skulls as they age.[23] A giraffe's skull is lightened by multiple 竇 (動物構造).[56]:70 However, as males age, their skulls become heavier and more club-like, helping them become more dominant in combat.[49] The upper jaw has a grooved and lacks front teeth.[57]:26 The giraffe's 大臼齿s have a rough surface.[57]:27

腿部, 运动系统和形态姿势编辑

The front and back legs of a giraffe are about the same length. The 桡骨 and 尺骨 of the front legs are articulated by the 腕骨, which, while structurally equivalent to the human wrist, functions as a knee.[61] It appears that a suspensory ligament英语suspensory ligament allows the lanky legs to support the animal's great weight.[62] The foot of the giraffe reaches a diameter of 30 cm(12英寸), and the is 15 cm(5.9英寸) high in males and 10 cm(3.9英寸) in females.[57]:36 The rear of each hoof is low and the fetlock英语fetlock is close to the ground, allowing the foot to provide additional support to the animal's weight.[11] Giraffes lack dewclaw英语dewclaws and interdigital glands. The giraffe's pelvis, though relatively short, has an 髂骨 that is outspread at the upper ends.[11]

A giraffe has only two gait英语gaits: walking and galloping. Walking is done by moving the legs on one side of the body at the same time, then doing the same on the other side.[49] When galloping, the hind legs move around the front legs before the latter move forward,[23] and the tail will curl up.[49] The animal relies on the forward and backward motions of its head and neck to maintain balance and the counter momentum while galloping.[43]:327–29 The giraffe can reach a sprint speed of up to 60 km/h(37 mph),[63] and can sustain 50 km/h(31 mph) for several kilometres.[64]

A giraffe rests by lying with its body on top of its folded legs.[43]: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 knees and spreads its hind legs to raise its hindquarters. It then straightens its front legs. With each step, the animal swings its head.[57]:31 In captivity, the giraffe sleeps intermittently around 4.6 hours per day, mostly at night.[65] 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 快速動眼期.[65] If the giraffe wants to bend down to drink, it either spreads its front legs or bends its knees.[49] Giraffes would probably not be competent swimmers as their long legs would be highly cumbersome in the water,[66] although they could possibly float.[67] 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[66][67] or keep its head above the surface.[66]

颈部编辑

 
长颈鹿(右)与其近亲物种㺢㹢狓(左)的颈椎都是七块骨头

长颈鹿的脖子非常长,可以达到 2~2.4米(6.6~7.9英尺),占其身长的很大一部分[52][53][57]:29[68] 它的长颈是由于颈椎不成比例的延长而不是颈椎骨的增多导致的。 每一節頸椎長度都超過 28 cm(11英寸) 。[56]:71 They comprise 52–54 per cent of the length of the giraffe's 脊椎, compared with the 27–33 percent typical of similar large ungulates, including the giraffe’s closest living relative, the 㺢㹢狓.[22] 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.[69] The giraffe's head and neck are held up by large muscles and a strengthened nuchal ligament英语nuchal ligament, which are anchored by long dorsal spines on the anterior 胸椎, giving the animal a hump.[11][70]

 
一头成年雄性网纹长颈鹿正在金合欢树下仰头进食,拍摄于肯尼亚

The giraffe's neck vertebrae have 杵臼關節s.[56]:71 In particular, the atlas英语Atlas (anatomy)axis英语Axis (anatomy) joint (C1 and C2) allows the animal to tilt its head vertically and reach more branches with the tongue.[57]:29 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 most other ruminants where the articulation is between the seventh cervical vertebra (C7) and T1.[22][69] 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.[70] However, this proposition is not generally accepted, as T1 has other morphological features, such as an articulating 肋骨, deemed diagnostic of thoracic vertebrae, and because exceptions to the mammalian limit of seven cervical vertebrae are generally characterised by increased neurological anomalies英语neurological disorder and maladies.[22]

There are several hypotheses regarding the evolutionary origin and maintenance of elongation in giraffe necks.[60] The "competing browsers英语browsing (predation) hypothesis" was originally suggested by 查尔斯·达尔文 and challenged only recently. It suggests that competitive pressure from smaller browsers, such as kudu英语kudu, 小岩羚 and 高角羚, 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 4.5米(15英尺) high, while even quite large competitors, such as kudu, can feed up to only about 2米(6英尺7英寸) high.[71] 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.[72][73] However, scientists disagree about just how much time giraffes spend feeding at levels beyond the reach of other browsers,[19][60][71][74] 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.[75]

Another theory, the 性選擇 hypothesis, proposes that the long necks evolved as a secondary 两性异形, giving males an advantage in "necking" contests (see below) to establish dominance and obtain access to sexually receptive females.[19] In support of this theory, necks are longer and heavier for males than females of the same age,[19][60] and the former do not employ other forms of combat.[19] However, one objection is that it fails to explain why female giraffes also have long necks.[76] It has also been proposed that the neck serves to give the animal greater vigilance.[77][78]

内部系统编辑

 
一只肯尼亚网纹长颈鹿 弯下来喝水。循环系统能够很好适应从颈部冲下来的血流。

In mammals, the left 喉返神經 is longer than the right; in the giraffe it is over 30 cm(12英寸) longer. These nerves are longer in the giraffe than in any other living animal;[79] the left nerve is over 2米(6英尺7英寸) long.[80] Each nerve cell in this path begins in the 腦幹 and passes down the neck along the 迷走神经, 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 5米(16英尺) in the largest giraffes.[79] The structure of a giraffe's brain resembles that of domestic cattle.[57]:31 It is kept cool by evaporative heat loss in the nasal passages.[58] 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英语Dead space (physiology), in spite of its narrow windpipe. These factors increase the resistance to airflow. Nevertheless, the animal can still supply enough oxygen to its tissues and it can increase its respiratory rate and oxygen diffusion when running.[81]

 
长颈鹿喝水时的吻端

The 循环系统 of the giraffe has several adaptations for its great height. Its heart, which can weigh more than 11公斤(25磅) and measures about 60 cm(2英尺) long, 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 7.5 cm(3.0英寸).[23] Giraffes have unusually high heart rates for their size, at 150 beats per minute.[56]:76 When the animal lowers its head the blood rushes down fairly unopposed and a 迷網 in the upper neck, with its large 截面 (幾何) area, prevents excess blood flow to the brain. When it raises again, the blood vessels constrict and direct blood into the brain so the animal does not faint.[82] The jugular vein英语jugular veins contain several (most commonly seven) valves to prevent blood flowing back into the head from the 下腔靜脈 and 心房 while the head is lowered.[83] 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.[50]

Giraffes have 食道 that are unusually strong to allow regurgitation of food from the stomach up the neck and into the mouth for rumination.[56]:78 They have four chambered stomachs, as in all ruminants, and the first chamber has adapted to their specialised diet.[11] The intestines of an adult giraffe measure more than 70米(230英尺) in length and have a relatively small ratio of small to large intestine.[84] The liver of the giraffe is small and compact.[56]:76 A gallbladder is generally present during fetal life, but it may disappear before birth.[11][85][86]

行为与生态编辑

栖息地和饮食编辑

 
在坦桑尼亚,一只马赛长颈鹿伸出舌头进食。 它的舌头,嘴唇和上颚都很厚实,足以应对树上的尖刺。

Giraffes usually inhabit 疏林草原s and open 疏林s. They prefer 含羞草亚科, 没药属, 风车藤属 and open 榄仁树属 woodlands over denser environments like Brachystegia英语Brachystegia woodlands.[43]:322 The Angolan giraffe can be found in desert environments.[87] Giraffes browse on the twigs of trees, preferring trees of the subfamily Acacieae and the genera Commiphora and Terminalia,[4] which are important sources of calcium and protein to sustain the giraffe's growth rate.[15] They also feed on shrubs, grass and fruit.[43]:324 A giraffe eats around 34公斤(75磅) of foliage daily.[49] When stressed, giraffes may chew the bark off branches. Although 食草动物, the giraffe has been known to visit carcasses and lick dried meat off bones.[43]:325

在雨季,食物丰富,长颈鹿分布更广,而在旱季,它们聚集在剩下的常绿乔木和灌木丛周围。[4] Mothers tend to feed in open areas, presumably to make it easier to detect predators, although this may reduce their feeding efficiency.[74] As a 反芻, 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.[56]:78–79 It is common for a giraffe to salivate while feeding.[57]:27 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.[4] The animal's faeces come in the form of small pellets.[11] When it has access to water, a giraffe drinks at intervals no longer than three days.[49]

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 trees that are too tall.[49] 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.[49]

社会性编辑

 
南非Tswalu Kalahari保护区内的一群南部长颈鹿。长颈鹿通常聚集成群。

长颈鹿通常成群出现。一般这些群体的组成被认为是开放且不断变化的。[88] Giraffes were thought to have few social bonds and 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."[89] More recent studies have found that giraffes do have long-term social associations and may form groups or pairs based on kinship, sex or other factors. These groups may regularly associate with one another in larger communities or sub-communities within a fission–fusion society英语fission–fusion society.[90][91][92] The number of giraffes in a group can range up to 44 individuals.[92]

长颈鹿哼叫声Ogg / Vorbis格式

长颈鹿喷气声Ogg / Vorbis格式

长颈鹿咕噜声Ogg / Vorbis格式

长颈鹿爆发声Ogg / Vorbis格式

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Giraffe groups tend to be sex-segregated[92] although mixed-sex groups made of adult females and young males are known to occur. Particularly stable giraffe groups are those made of mothers and their young,[89] which can last weeks or months.[93] Social cohesion in these groups is maintained by the bonds formed between calves.[43]:330[89] Female association appears to be based on space-use英语Philopatry and individuals may be 母系制度 related.[92] In general, females are more selective than males in who they associate with in regards to individuals of the same sex.[91] 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.[92][93] Giraffes are not 领域 (动物),[11] but they have home range英语home ranges.[49] Male giraffes occasionally wander far from areas that they normally frequent.[43]:329

尽管长颈鹿通常安静无声,但已听到长颈鹿使用各种声音进行交流。During courtship, males emit loud coughs.[49] Females call their young by bellowing. Calves will emit snorts, bleats, mooing and mewing sounds. Giraffes also snore, hiss, moan, grunt and make flute-like sounds,[49][94] and possibly communicate over long distances using 次声波[95]—though this is disputed.[94] During nighttime, giraffes appear to hum英语humming to each other above the infrasound range for purposes which are unclear.[94]

繁殖和父母监护行为编辑

纳米比亚的南部长颈鹿在正在求爱(交配)。通常,只有占优势的雄性才能够与雌性交配。

Reproduction in giraffes is broadly 多配偶制: a few older males mate with the fertile females. Male giraffes assess female fertility by tasting the female's urine to detect 動情週期, in a multi-step process known as the 裂唇嗅反應.[89][93] Males prefer young adult females over juveniles and older adults.[89] Once an oestrous female is detected, the male will attempt to court her. When courting, dominant males will keep subordinate ones at bay.[93] A courting male may lick a female's tail, rest his head and neck on her body or nudge her with his horns. During copulation, the male stands on his hind legs with his head held up and his front legs resting on the female's sides.[49]

Giraffe gestation英语gestation lasts 400–460 days, after which a single calf is normally born, although twins occur on rare occasions.[96] The mother gives birth standing up. The calf emerges head and front legs first, having broken through the fetal membrane英语fetal membranes, and falls to the ground, severing the 脐带.[11] The mother then grooms the newborn and helps it stand up.[57]:40 A newborn giraffe is 1.7~2米(5.6~6.6英尺) tall.[51][52][53] Within a few hours of birth, the calf can run around and is almost indistinguishable from a one-week-old. However, for the first 1–3 weeks, it spends most of its time hiding;[97] its coat pattern providing camouflage. The ossicones, which have lain flat while it was in the womb, become erect within a few days.[49]

 
位于肯尼亚的马赛长颈鹿幼崽和成年雌性在进食。抚养后代的主要是雌性长颈鹿。 长颈鹿家庭经常出现在在人类的苗圃中。

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 "calving pool英语Crèche (zoology)".[97] Adult males play almost no role in raising the young,[43]:337 although they appear to have friendly interactions.[89] Calves are at risk of predation, and a mother giraffe will stand over her calf and kick at an approaching predator.[49] Females watching calving pools will only alert their own young if they detect a disturbance, although the others will take notice and follow.[97]

长颈鹿后代与母亲在一起的时间长短不一,尽管可以一直呆到雌性下一次产犊。[97] 小长颈鹿的哺乳期一般从最短一个月[43]:335 到最长一年。[49][93] 雌性长颈鹿四岁时性成熟,雄性在四到五岁成熟。雄性长颈鹿的精子发生在三到四岁已经开始了。[98] 但是雄性必须等到至少七岁才有机会交配。[49][57]:40

脖击编辑

 
南非夸祖魯-納塔爾省野生动物保护区,雄性南部长颈鹿通过低强度脖击来展示自己优势地位。
 
纳米比亚的一对冲突中的南部长颈鹿,他们采用了更积极的脖击方式。

雄性长颈鹿在战斗中以脖子为武器,这种行为被称为“脖击”。脖击用于建立支配地位,赢得脖击的雄性具有更大的生殖成功率。[19] This behaviour occurs at low or high intensity. In low intensity necking, the combatants rub and lean against each other. The male that can hold itself more erect 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 get ready to counter. The power of a blow depends on the weight of the skull and the arc of the swing.[49] A necking duel can last more than half an hour, depending on how well matched the combatants are.[43]:331 Although most fights do not lead to serious injury, there have been records of broken jaws, broken necks, and even deaths.[19]

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.[99] In one study, up to 94 percent of observed mounting incidents took place between males. The proportion of same-sex activities varied from 30–75 percent. Only one percent of same-sex mounting incidents occurred between females.[100]

死亡率和健康编辑

 
一只和一头被猎杀的成年马赛长颈鹿( Masai giraffe英语Masai giraffe

长颈鹿有很高的成活率,[9] 与其他反刍动物相比,寿命异常长,[101] 在野外可存活长达25年。[50] Because of their size, eyesight and powerful kicks, adult giraffes are usually not subject to predation,[49] aside from s.[102] Giraffes are the most common prey for the big cats in 克留格爾國家公園.[103][104] 尼羅鱷s can also be a threat to giraffes when they bend down to drink.[57] Calves are much more vulnerable than adults, and are additionally preyed on by s, 斑鬣狗s and 非洲野犬.[23] A quarter to a half of giraffe calves reach adulthood.[9][105] Calf survival varies according to the season of birth, with calves born during the dry season having higher survival rates.[106] The local, seasonal presence of large herds of migratory 角马s and 斑馬s reduces predation pressure on giraffe calves and increases their survival probability.[107]

Some parasites feed on giraffes. They are often hosts for 真蜱科s, especially in the area around the genitals, which has thinner skin than other areas.[11] Tick species that commonly feed on giraffes are those of genera Hyalomma英语Hyalomma, Amblyomma英语Amblyomma and Rhipicephalus英语Rhipicephalus. Giraffes may rely on red-billed英语Red-billed Oxpecker and yellow-billed oxpecker英语yellow-billed oxpeckers to clean them of ticks and alert them to danger. Giraffes host numerous species of internal parasite and are susceptible to various diseases. They were victims of the (now eradicated) viral illness 牛瘟病毒.[11] Giraffes can also suffer from a skin disorder, which comes in the form of wrinkles, 皮膚病 or raw fissures英语skin fissure. In Tanzania, it appears to be caused by a 线虫动物门, and may be further affected by secondary infections. As much as 79% of giraffes show signs of the disease in 鲁阿哈国家公园, but is less prevalent in areas with fertile soils.[108][109][110]

与人类的联系编辑

 
纳米比亚描绘长颈鹿的古老岩画

人类与长颈鹿相处的历史已经有数千年。南非的萨恩人 有将医疗舞蹈(巫舞)命名为动物的传统,其中就有治疗头部疾病的长颈鹿舞。[111]长颈鹿为何长得这么高一直是非洲民俗中热议的话题,[19] 比如东非某民俗认为这是因为长颈鹿吃太多神奇的草药。[112]在整个非洲范围内,长颈鹿是重要的艺术元素,尤其是早期文明比如Kiffians英语Kiffian culture, 古埃及麦罗埃努比亚[57]:45–47 Kiffians英语Kiffian culture人曾制作了一个包含两个长颈鹿的等身大小的岩石雕刻,这也是世界上最大的岩石岩画.[57]:45[113] 古埃及人为长颈鹿创造了圣书体写法𓃱, 在埃及语中代表“sr”之后又代表 “mmy”。[57]:49 古埃及人还将长颈鹿作为宠物饲养并将它们运送到地中海沿岸各地。[57]:48–49

 
清代陈璋根据明代沈度的《瑞应麒麟图》临摹而成的《榜葛剌进麒麟图》,现藏于中国国家博物馆。该图描绘永乐十二年(1414年)榜葛剌(今孟加拉)使臣向明成祖进贡长颈鹿的场景。

古希腊古罗马的记述中也存在长颈鹿,但长颈鹿被认为是骆驼和猎豹的不自然杂交体。[57]:50 长颈鹿是被收集并被送回罗马展览的众多动物之一。已知第一只被带到罗马对公共展览的长颈鹿是公元前46年尤利烏斯·凱撒带回的。[57]:52 随着罗马帝国的衰落, 在欧洲被饲养的长颈鹿逐渐绝种[57]:54 中世纪期间,欧洲人对于长颈鹿的认识主要来自于和阿拉伯人的交流,阿拉伯人非常独特地尊重长颈鹿,[23]

在各地古代史中,被圈养的长颈鹿(数量并不多)一般享有崇高的地位。1414年,一只长颈鹿被从馬林迪运至孟加拉 (地區)。之后被郑和下西洋的船队带回并安置在明朝的皇室动物园。这种动物令中国人着迷,他们将其与神话中的麒麟联系在一起。[57]:56 1486年,一只被命名为美第奇长颈鹿(Medici giraffe英语Medici giraffe)的长颈鹿被敬献给洛伦佐·德·美第奇,它一抵达佛罗伦萨便引起了轰动。[114] 另外一只著名的长颈鹿,Zarafa英语Zarafa (giraffe), 在十九世纪早期,被穆罕默德·阿里帕夏送给查理十世 (法兰西)。这只长颈鹿在巴黎了引起了轰动,市面上一时流行大量的纪念品,甚至有词汇"giraffanalia"来专门描述它。[57]:81

长颈鹿继续在现代文化中占有一席之地。艺术家萨尔瓦多·达利在他的超现实主义绘画中将长颈鹿描绘得一种拥有燃烧的鬃毛的生物。达利认为长颈鹿是男子气概的象征,而燃烧的长颈鹿则是“男性宇宙的末日怪物(masculine cosmic apocalyptic monster)”。[57]:123 长颈鹿也是儿童读物中常见的元素,包括大卫·A·乌弗的《怕高的长颈鹿(The Giraffe Who Was Afraid of Heights)》;吉尔斯·安德烈的《长颈鹿不会跳舞(Giraffes Can't Dance)》; 羅爾德·達爾的《长颈鹿,佩利和我(The Giraffe and the Pelly and Me)》。长颈鹿多次作为动画中的次要角色出现在华特迪士尼公司的动画电影中,比如《狮子王》和《小飞象》,并在《野蛮任务》和《馬達加斯加系列電影》中扮演相当重要的角色。苏菲小鹿宝宝(Sophie the Giraffe英语Sophie the Giraffe)自1961年推出以来,都是奶嘴的热门产品。另一个著名的长颈鹿玩具是玩具反斗城的吉祥物 ”R“[57]:127

长颈鹿也已经用于一些科学实验和研究。科学家在为宇航员和战斗机飞行员研制服装的时候广泛研究了长颈鹿的特征,[56]:76 因为这些人士因为处于高重力加速度环境有血液急速流向下半身的危险。 计算机科学家利用反应-扩散机制对几种长颈鹿的外皮模式进行了建模。[115]

鹿豹座的星座,于十七世纪问世,描绘的就是一只长颈鹿。[57]:119–20 茨瓦纳人的传统将南十字座看作两只长颈鹿。其中十字架二十字架三组成了一只雄性,而十字架一十字架四构成了雌性长颈鹿。[116]

开发及保护状况编辑

 
在二十世纪早期于德屬東非被当地部落猎杀的马赛长颈鹿(Masai giraffe英语Masai giraffe

长颈鹿是整个非洲的猎人的共同目标。[43]被猎杀后,它们身体的各部分都有不同用途:[11]肉主要用于食用;尾毛主要用于制作苍蝇拍,手镯,项链和线;[11][43] 皮肤可以被制成盾牌,凉鞋和鼓;部分的筋被用于制作乐器的弦;[11] 传统布干达的医师使用燃烧长颈鹿皮肤产生的烟雾来治疗流鼻血。[43]位于苏丹的Humr人常常饮用认为蕴藏长颈鹿灵魂的Umm Nyolokhh饮料,这种液体主要由长颈鹿的肝脏和骨髓制作,通常包含二甲基色胺 和长颈鹿吃的植物中所包含的精神活性物质,例如长颈鹿常见食物金合欢中一些生物碱已经被证实会引起长颈鹿的幻觉。[117][118] 在19世纪,欧洲探险家开始将猎杀长颈鹿作为一种打猎运动。[57]

栖息地破坏也影响了长颈鹿的生存:在萨赫勒,对牲畜的放牧空间和木柴的需求导致了森林砍伐。但通常,长颈鹿可以很好与牲畜共存,因为它们不直接与牲畜竞争。[50]2017年,肯尼亚北部的严重干旱导致牧场稀缺的局势加剧,牧民因此杀害了大量野生动物,长颈鹿种群尤其受到打击。[119]

航空测量法是监测非洲广袤地区长颈鹿种群情况的最方便也是最常用的方法,但空中方法会低估长颈鹿的数量。[120] 地面调查法相对更加准确,与航空调查结合使用的话,可以准确估算种群规模和趋势.[120]2010年,长颈鹿被國際自然保護聯盟 (IUCN)评估为无危物种,但在2016年又被认定为易危物種[1] 在一些特定地区包括厄立特里亚几内亚毛里塔尼亚塞内加尔长颈鹿已经局部地區滅絕。长颈鹿也有可能从安哥拉马里奈及利亞消失, 但是长颈鹿已经被引进到卢旺达斯威士兰[1] 长颈鹿的两个亚种,西非长颈鹿和羅氏長頸鹿已经被认定为瀕危物種[41][40] 因为它们的野外种群数量都已经只剩数百。

1997年,Jonathan Kingdon英语Jonathan Kingdon认为努比亚长颈鹿是最受威胁的长颈鹿;[4] 2010年不精确的估计显示,该模式种数量已经少于250。私人野生动物保护区对于保护非洲南部的长颈鹿种群起来很大帮助[50] 比如长颈鹿庄园(Giraffe Manor)就是奈洛比的热门酒店,也是罗氏长颈鹿的避难所。

参考编辑

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