犬齿窝与人类中面部骨骼的演化

doi:10.16359/j.1000-3193/AAS.2020.0016

摘要/Abstract

摘要：

犬齿窝是包括现代人类在内的许多人族成员面部骨骼的重要性状,但在分类学上的意义仍存在争议。有学者认为该性状是一个发生于基础面形的近祖性状,除了一些例外,在灭绝和现生的大猿及人属中都存在。另有学者认为,犬齿窝是仅存在于智人及其直系祖先的衍生性状,在发育上与颧齿槽突嵴有关。这种关系并非总是成立,在智人中存在着明显的差异：弧型颧齿槽突嵴和直斜型颧齿槽突嵴与犬齿窝有时共存,有时不共存。我们由此推测,犬齿窝的发生和形态与上颌窦的前部发育有关,颧齿槽突嵴的形态与鼻窦的侧面发育有关。在人类演化的过程中,犬齿窝经历了不同的变形,比如上颌沟(如南方古猿非洲种、傍人粗壮种)、上颌小窝(如傍人粗壮种)、上颌沟(如匠人)或犬齿窝缺如(如傍人埃塞俄比亚种、傍人鲍氏种、肯尼亚扁脸人、人属鲁道夫种)。犬齿窝消失的原因各类群并不相同,如中新世和早更新世人属以及中更新世人属(如人属海德堡种/人属罗得西亚种、人属尼安德特种)。人属罗得西亚种具有弱化的犬齿窝,不具备演化为智人的可能,因此被排除在智人的演化支之外。

关键词: 面部骨骼, 中面部, 眶下板, 人类演化, 智人种

Abstract:

The canine fossa is an important feature in the facial skeleton of many hominins, including modern humans. However, its phylogenetic significance is debated. Some researchers consider it as a plesiomorphic characteristic in a generalized face that, with some exceptions, is found in both extinct and extant great apes and in the hominins. Others consider that the canine fossa is a derived characteristic only found in Homo sapiens and its direct ancestors, and that it is related to an arched zygomatic-alveolar crest (ZAC). However, this relationship is not always fulfilled, and in Homo sapiens, there is a notable variability: An arched ZAC with the presence or absence of a canine fossa, and straight oblique ZAC with presence or absence of a canine fossa. In this sense, we hypothesize that the canine fossa is related to the degree of anterior extension of the maxillary sinus and that the morphology of the ZAC is related to the degree of lateral extension of said sinus. During the hominine evolution, the canine fossa has undergone different transformations, such as the maxillary furrow (Australopithecus africanus, Paranthropus robustus), maxillary fossula (P. robustus), and the sulcus maxillaris (Homo ergaster), or has been obliterated (Paranthropus aethiopicus, Paranthropus boisei, Kenyanthropus platyops, Homo rudolfensis). In the taxa where it has been obliterated, the mechanism of obliteration is different in the hominins of the Pliocene and Early Pleistocene compared to those of the Middle Pleistocene (Homo heidelbergensis/rhodesiensis, Homo nenaderthalensis). The facial morphology of H. rhodesiensis would exclude it from the line evolving to H. sapiens.

Key words: Facial skeleton, Midface, Infraorbital plate, Hominin evolution, Homo sapiens

中图分类号:

Q987

Francesc RIBOT Trafí, Mario GARCÍA Bartual, Alfredo José ALTAMIRANO Enciso, 王谦. 犬齿窝与人类中面部骨骼的演化[J]. 人类学学报, 2022, 41(02): 193-217.

Francesc RIBOT Trafí, Mario GARCÍA Bartual, Alfredo José ALTAMIRANO Enciso, Qian WANG. Canine fossa and evolution of the human mid-facial bones[J]. Acta Anthropologica Sinica, 2022, 41(02): 193-217.

图/表 10

图1 现代人犬齿窝的变异从左到右依次为：轻微扩张、浅犬齿窝From left to right: little expanded and shallow canine fossa(Y1 CAH86 EXP-ESTE—Q.W.拍摄);扩张且浅的犬齿窝expanded and shallow(LPPMANC collection—F.R.T.拍摄);扩张且相对深的犬齿窝expanded and relatively deep(SON 02.03.21—A.J.A.E.拍摄);扩张且深的犬齿窝expanded and deep(LPPMANC collection—F.R.T.拍摄)

Fig.1 Variability of the canine fossa in current humans

图2 现代类人猿基础面形的犬齿窝左：雄性猩猩 L: Pongo pygmaeus male (NMNH153828 —Q.W.拍摄);右：雄性大猩猩 R: Gorilla gorilla male (Museum Reverte collection —M.G.B.拍摄)

Fig.2 Generalized morphology of the canine fossa in current hominids

表1 东非和南非不同年代和地区的南方古猿种属

Tab.1 Localities and chronologies for the different Australopithecus species of East Africa and South Africa

区域Area	种群Taxon	地点Locality	年代Age (MaBP)	参考文献Reference
东非East Africa	南方古猿湖畔种A. anamensis	Asa Issie(埃塞俄比亚)	4.2-4.1	[189]
	南方古猿湖畔种A. anamensis	Kanapoi(肯尼亚)	4.1	[190]
		Allia Bay(肯尼亚)	3.9	[190]
		Woranso Mille(埃塞俄比亚)	3.8-3.6	[191]
	莱托里的Garusi I 标本	Laetoli(坦桑尼亚)	3.7-3.5	[192]
	南方古猿阿法种A. afarensis	Woranso Mille(埃塞俄比亚)	3.6	[193]
	南方古猿阿法种A. afarensis	Tulu Bor(肯尼亚)	3.7-3.4	[194]
		Maka(埃塞俄比亚)	3.4	[195]
		Hadar(埃塞俄比亚)	3.4-3	[188]
		Dikika(埃塞俄比亚)	3.3	[196]
	南方古猿近亲种 A. deiyremeda	Woranso Mille(埃塞俄比亚)	3.5-3.3	[73]
	南方古猿惊奇种A. garhi	Bouri(埃塞俄比亚)	2.5	[74]
南非South Africa	南方古猿普罗米修斯种 A. prometheus	Sterkfontein member 2	3.67	[197]
	南方古猿普罗米修斯种 A. prometheus	Makapansgat	2.9-2.6	[198]
		Sterkfontein member 4	2.6-2	[198]
	南方古猿非洲种 A. africanus	Taung	3.0-2.6	[198]
	南方古猿非洲种 A. africanus	Makapansgat	2.9-2.6	[198]
		Sterkfontein member 4	2.6-2	[198]
		Gladysvale	2.4-2	[198]
	南方古猿源泉种A.sediba	Malapa	1.95-1.78	[80]

图3 存在颧齿槽突嵴但不存在犬齿窝的人属上排,从左到右：OH5(傍人鲍氏种;铸模);KGA10-525(傍人鲍氏种;修改自[91]);KNM-WT 40000(肯尼亚扁脸人;铸模)。下排,从左到右：KNM-ER 1470(人属鲁道夫种;铸模);Sangiran 17(人属匠人种;铸模);Sima de los Huesos 5号(可能是人属海德堡种;铸模);Kabwe 1(可能是人属罗得西亚种;铸模)。比例尺不同

Fig.3 Hominins with the ZAC arched and without a canine fossa Top, from left to right: OH 5 (P. boisei, cast); KGA 10-525 (P. boisei, modified from [91]); KNM-WT 40000 (K. platyops, cast). Bottom, from left to right: KNM-ER 1470 (H. rudolfensis, cast); Sangiran 17 (H. ergaster, cast); Sima de los Huesos cranium 5 (probable H. heidelbergensis, cast); Kabwe 1 (probable H. rhodesiensis, cast). Not at the same scale.

表2 早更新世人属颅脑性状的特殊分类及匠人与直立人的区别

Tab.2 Craniofacial features that allow a specific classification in Homo of the lower Pleistocene and the differentiation between Homo ergaster and Homo erectus

性状Feature	鲁道夫种H. rudolfensis	能人种H.habilis	匠人种H.ergaster	直立人种H. erectus
眶下面	朝向冠状面,平坦, 向前向下倾斜	朝向冠状面, 相对屈曲, 向后向下倾斜	朝向冠状面, 相对屈曲, 向后向下倾斜	朝向冠状面,屈曲,向后向下倾斜
犬齿窝	缺失	存在	上颌沟	存在
上颌拐角	非常微弱	微弱	微弱	显著
颧齿槽突嵴	起源低且朝前,多变	起源高且朝后,弧形	多变,多变	起源高且朝后,弧形
颧切迹	缺失	缺失	缺失^a	存在
颧上颌结节*	缺失	缺失	缺失^b	存在
鼻侧外翻	非常微弱	非常微弱	存在	存在
鼻下面	直角	突颌	中间	中间
鼻下缘形态*	2	2 和 3	2 和3	3^c
颞鳞上缘*	直线	直线	直线	屈曲

图4 存在颧齿槽突嵴但不存在犬齿窝的现代人样本从左到右：LPPMANC,collection(F.R.T.拍摄);新店34,500BCE(Q.W.拍摄);图片来自[200];井沟子03LJM46B,400BCE(Q.W.拍摄)。比例尺不同

Fig.4 Examples of current humans with the ZAC arched and without a canine fossa From left to right: LPPMANC collection (photo F.R.T.); Xindianzi 34 500BCE (photo Q.W.); photo taken from [200]; Jinggouzi 03LJM46B 400BCE (photo Q.W.). LPPMAN: Paleoanthropology and Paleopathology Laboratory of the National Museum of Archeology of Catalonia. Not at the same scale

图5 颧齿槽突嵴直斜且存在犬齿窝的人属(上,灭绝;下,现生) 上排,从左到右：Sts52(南方古猿非洲种;改自 [201]);MH1(南方古猿源泉种;铸模);Sima de los Huesos AT-404(可能是人属海德堡种;引自[116]);Steinheim(可能是人属海德堡种;铸模)。下排,从左到右：SON 02.03.29(A.J.A.E.拍摄);SON 02.02.13(A.J.A.E.拍摄);图片引自[202];图片由Dr. Domènech Campillo提供。

Fig.5 Hominins (top, extinct; bottom, extant) with the ZAC straight oblique and with a canine fossa Top, from left to right: Sts 52 (A. africanus, modified from [201]); MH1 (A. sediba, cast); Sima de los Huesos AT-404 (a probable H. heidelbergensis, taken from [116]); Steinheim (a probable H. heidelbergensis, cast). Bottom, from left to right: SON 02.03.29 (photo A.J.A.E.); SON 02.02.13 (photo A.J.A.E.); photo taken from [202]; photo courtesy of Dr. Domènech Campillo.

图6 欧洲直立人的犬齿窝欧洲直立人ATD6-69 (A)和ATD6-58 (B)的窝位于眶下孔外侧但不低于眶下孔,两种性状无关联。在有颧颌窝的个体中可以观察到类似的形态(Oschinsky,黄色箭头)：(C)南方古猿源泉种MH1,蓝色箭头指向犬齿窝;(D)人属能人种KNM-ER 1813(翻转,铸模);(E)SH5(铸模);(F)大荔人(翻转,铸模)。ATD6-69来自于FAQ at the talk.origins Archive;ATD6-58来自于Schwartz & Tattersall, 2002[199]。在阿塔普尔卡直立人中,犬齿窝更宽阔、更深,但在个体发生过程中由于上颌窦的前伸其深度逐渐减少。注意ATD6-69眶下孔下部平坦或略微突出,与智人的形态不同。

Fig.6 Hominins with zygomaxillary fossa (Oschinsky fossa) in European H. erectus An unusual form of presentation of the canine fossa (?) in European H. erectus [ATD6-69 (A) and ATD6-58 (B)]: it is not inferior to the infraorbital foramen, but is lateral to it and without both features having contact. A similar morphology can be observed in specimens with zygomaxillary fossa, Oschinsky (yellow arrows): A. sediba MH1 (C) ; the blue arrow indicates the canine fossa; H. habilis KNM-ER 1813 (flipped, cast) (D); SH5 (cast) (E); Dali (flipped, cast) (F). ATD6-69 taken from FAQ at the talk.origins Archive; ATD6-58 taken from Schwartz & Tattersall, 2002. In the Atapuerca H. erectus, the fossa is more widespread and deeper, but during ontogeny it is reduced in depth by the anterior expansion of the maxillary sinus. Note that the region inferior to the infraorbital foramen in ATD6-69 is flat or somewhat convex, morphology different from H. sapiens.

表3 不同人类种群犬齿窝的变异

Tab.3 Variation of the canine fossa in the different human species

分类群Taxon		犬齿窝形态Morphology of the canine fossa
乍得撒海尔人Sahelanthropus tchadensis		宽而浅
始祖地猿Ardipithecus ramidus		未观察到
更新纪灵长类动物Australopiths	南方古猿湖畔种Australopithecus anamensis	位于犬齿突和P⁴突之间
更新纪灵长类动物Australopiths	莱托里个体, Garusi I	位于犬齿突和P⁴突之间
	阿法种Australopithecus afarensis	两种形态：深而凹陷 (可能是男性);表面的凹陷 (可能是女性)
	近亲种Australopithecus deyiremeda	初步发育
	惊奇种Australopithecus ghari	位于犬齿突和P⁴突之间
	普罗米修斯种Australopithecus prometheus	三种形态：类上颌沟Like-sulcus maxillaris^a;非常微弱的上颌沟Very weak maxillary furrow^b;上颌小窝Maxillary fossula^c
	非洲种Australopithecus africanus	上颌沟^d
	源泉种Australopithecus sediba	位于犬齿突和P⁴突之间
	肯尼亚扁脸人Kenyanthropus platyops	完全消失
	傍人埃塞俄比亚种Paranthropus aethiopicus	完全消失
	鲍氏种Paranthropus boisei	完全消失
	粗壮种Paranthropus robustus	上颌小窝
人属Homo	能人种Homo habilis	小,位于犬齿突和P⁴突之间
	鲁道夫种Homo rudolfensis	完全消失
	匠人种Homo ergaster	上颌沟^e
	直立人种Homo erectus	犬齿窝宽阔^f
	海德堡罗得西亚种Homo heidelbergensisrhodesiensis	完全消失^g
	尼安德特种Homo neanderthalensis	完全消失
	纳莱迪种Homo naledi	浅,P³突上难以定义的犬齿窝
	智人种Homo sapiens	犬齿窝宽阔

图7 犬齿窝两侧的不同表型

Fig.7 Phenotypes of the canine fossa at two sides LPPMANC collection—photo F.R.T.

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