A skull of Early Pleistocene Paracamelus gigas (Artiodactyla, Mammalia) from Luotuo Hill in Dalian, Northeast China

doi:10.19615/j.cnki.2096-9899.230616

Abstract

Abstract:

Originated in North America in the Middle Eocene, camelids were a successful group with very large diversity. But the camels emigrated to the Old World from North America, probably during the middle stage of the Middle Miocene, and did not radiate much as those in North America, represented by only two genera Paracamelus and Camelus. The former was considered as giving rise to the latter, but the detailed relationship of the Old World camelines was controversial. The new camel material unearthed from Layer 4 in the Jinyuan Cave at Luotuo Hill in Dalian, Liaodong peninsula in Northeast China, was described and referred to as Paracamelus gigas. Its dentition length is slightly longer than that of Camelus knoblochi but evidently larger than that of C. ferus and C. dromedarius. Based on the fossil records and morphometric evidences, P. gigas originated from a form similar to P. alexejevi in the Late Pliocene in the Old World, instead of from Megatylopus gigas of North America and then migrated into Asia as previously thought. The morphometric similarities between the Early Pleistocene Dalian specimens and those of the Middle and Late Pleistocene C. knoblochi indicate that P. gigas probably gave rise to C. knoblochi as formerly postulated and likely in the late Early Pleistocene by reduction or simplifying of P3 and P4, disappearance of p3 and shortening of dentition length. P. gigas inhabited in the forest steppe environment of Liaodong peninsula from 1.1 to 1.52 Ma based on paleomagnetic dating and pollen evidence.

Key words: Dalian, Early Pleistocene, cave deposits, camel, Paracamelus, evolution

摘要：

起源于中始新世的骆驼科(Camelidae)是一支在北美的新生代期间演化非常成功的类群，具有很大的多样性。其中的一类约在中中新世期间经白令陆桥迁徙到亚洲，随后扩散到欧洲和非洲。虽然骆驼科在北美新生代期间演化辐射出很多种类，但从北美扩散到旧大陆的骆驼只有一个族(Camelini)的两个属：副驼(Paracamelus)和骆驼(Camelus)。巨副驼(Paracamelus gigas)是我国境内最早发现的化石骆驼，一直被认为是起源于北美的Megatylopus之类的大型骆驼，然后扩散到旧大陆其他地区，最后演化成诺氏驼(Camelus knoblochi)及双峰驼(C. bactrianus)。但也有学者认为骆驼属起源于非洲。最近在大连复州湾骆驼山金远洞第四纪堆积剖面的第4层中出土了一些骆驼化石，其中1件破损的头骨和带有两枚下臼齿的残破下颌骨经研究被归入巨副驼。根据对巨副驼及其他副驼种的地理及年代分布的研究，巨副驼的直接祖先应该类似于晚上新世分布在欧亚大陆的体型稍小的阿氏副驼(P. alexejevi)或相似类型。大连出产的巨副驼在形态及大小上与诺氏驼接近，但其齿列长度略大于诺氏驼且明显大于野生双峰驼(C. ferus)及单峰驼(C. dromedarius)。而巨副驼与诺氏驼在地理分布上的重叠范围较大，时代分布上呈先后关系，因此可以认为巨副驼是诺氏驼的直接祖先，巨副驼在中更新世晚期通过P3收缩或简化及p3消失而演化成诺氏驼。综合化石层位的古地磁测年及花粉分析结果判断，巨副驼在早更新世的1.1~1.52 Ma期间栖息在大连半岛的森林草原环境中。

关键词: 大连, 早更新世, 洞穴堆积, 骆驼, 巨副驼, 演化

CLC Number:

Q915.876

DONG Wei, LIU Wen-Hui, BAI Wei-Peng, LIU Si-Zhao, WANG Yuan, LIU Jin-Yuan, JIN Chang-Zhu. A skull of Early Pleistocene Paracamelus gigas (Artiodactyla, Mammalia) from Luotuo Hill in Dalian, Northeast China. Vertebrata Palasiatica, 2024, 62(1): 47-68.

董为, 刘文晖, 白炜鹏, 刘思昭, 王元, 刘金远, 金昌柱. 2024, 62(1): 47-68, 辽宁大连骆驼山早更新世巨副驼头骨化石. 古脊椎动物学报.

Figures/Tables 8

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	Paracamelus gigas					Camelus knoblochi			C. ferus	C. dromedarius
	DLJ168001		Honan¹⁾	Mianchi²⁾	Nihewan³⁾	Luchka⁴⁾	Razdorskaya⁴⁾	Sengiley⁴⁾	c/o.20.705.676	ov25
	left	right	Honan¹⁾	Mianchi²⁾	Nihewan³⁾	Luchka⁴⁾	Razdorskaya⁴⁾	Sengiley⁴⁾	c/o.20.705.676	ov25
I3 L	14.42	16.06		12.8		20	21*	20	6.1-6.9	(9.5)
I3 W	10.27	11.97		9.1		15	17*	16	4.3-6.9	(7.2)
I3 H	19.37	20.6							9.9-18.7
C L	39.63	40.18		16.4	20†	40*	22.0; 23.1	34; 37	11.9-12.1	12.4-12.7
C W	28.32	24.48		12.2	14†	29*			7.4-9.2	8.7-8.9
C H	80.05	56.22							17.6-29.3
P1 L	17.57	19.16				22*		24; 23.2	9.1-9.5	8.7-8.9
P1 W	14.11	14.23				18*		19; 17	7.2-7.4	6.3-6.5
P1 H	31.18	33.16							22.1-22.8	17.1-17.9
P3 L	28.72	29.51		(28)	31	26	18; 18.7	28; 27	17.3-18.2	18.7-20.1
P3 W	21.52	23.14		(20)	21	25.5	17; 17	20; 22	13.8-15.1	12.7-13.5
P3 H	24.91	25.71							16.2-19.2	27.1-29.2
P4 L	32.67	33.35		30	29	27; 28	22.4; 24	29.5; 31	19.1-22.9	23.6-24.3
P4 W	30.19	39.63		29.6	28	29.4; 29	27.1; 28	29; 29.3	18.4-25.9	22.3-23.3
P4 H	27.82	29.15							18.7-23.3	23.9-25.7
M1 L	37.46	37.32	47§	(36)	38	33.3; 31	34.3; 34.4	32.8; 33.8	25.8-32.1	26.9-30.6
M1 W	41.84	36.15	38§	38.5	37	34; 33	33; 34.1	34.2; 34.2	30.1-33.2	29.1-30.7
M1 H	24.82	24.97	25§						19.4-24.9	11.7-21.1
M2 L	48.48	47.36	50	(46)	45.5#	51	42; 41.6	54.3; 53	37.2-45.2	39.8-40.2
M2 W	39.34	38.35	41	42.4	35.0#	38	36.6; 37.9	37.3; 35.3	31.1-35.8	28.9-31.1
M2 H	30.75	31.87	36						19.3-28.9	24.3-24.4
M3 L	51.98	52.23		55	43.0#	59	52; 52.7	61.4; 61.3	39.9-46.2	42.2-44.7
M3 W	35.99	35.74		43.3	34.5#	35	36.1; 36.8	35; 25	24.3-32.3	27.1-31.6
M3 H	33.79	35.08							20.1-22.5	27.8-30.1
Di I3-C	20.51	23.28		18		21.0	14*		12.1-16.8	10.9
Di C-P1	33.4	37.61			72†		16*		30.1-41.7	35.4-35.9
Di P1-3	61.62	59.79			>60	51.0; 49.0	49*		24.1-43.8	43.9-47.8
P3-4 L	61.59	64.63		(58)			38.5; 39.2		37.2-39.8	35.1-39.1
M1-3 L	136.35	136.85		(131)		140	123; 124	144; 142.3	104.1-112.4	111.2-115.9
P3-M3 L	199.6	203.5		(186)		184			135.7-145.3	141.1-143.7

	Paracamelus gigas					Camelus knoblochi			C. ferus	C. dromedarius
	DLJ168001		Honan¹⁾	Mianchi²⁾	Nihewan³⁾	Luchka⁴⁾	Razdorskaya⁴⁾	Sengiley⁴⁾	c/o.20.705.676	ov25
	left	right	Honan¹⁾	Mianchi²⁾	Nihewan³⁾	Luchka⁴⁾	Razdorskaya⁴⁾	Sengiley⁴⁾	c/o.20.705.676	ov25
I3 L	14.42	16.06		12.8		20	21*	20	6.1-6.9	(9.5)
I3 W	10.27	11.97		9.1		15	17*	16	4.3-6.9	(7.2)
I3 H	19.37	20.6							9.9-18.7
C L	39.63	40.18		16.4	20†	40*	22.0; 23.1	34; 37	11.9-12.1	12.4-12.7
C W	28.32	24.48		12.2	14†	29*			7.4-9.2	8.7-8.9
C H	80.05	56.22							17.6-29.3
P1 L	17.57	19.16				22*		24; 23.2	9.1-9.5	8.7-8.9
P1 W	14.11	14.23				18*		19; 17	7.2-7.4	6.3-6.5
P1 H	31.18	33.16							22.1-22.8	17.1-17.9
P3 L	28.72	29.51		(28)	31	26	18; 18.7	28; 27	17.3-18.2	18.7-20.1
P3 W	21.52	23.14		(20)	21	25.5	17; 17	20; 22	13.8-15.1	12.7-13.5
P3 H	24.91	25.71							16.2-19.2	27.1-29.2
P4 L	32.67	33.35		30	29	27; 28	22.4; 24	29.5; 31	19.1-22.9	23.6-24.3
P4 W	30.19	39.63		29.6	28	29.4; 29	27.1; 28	29; 29.3	18.4-25.9	22.3-23.3
P4 H	27.82	29.15							18.7-23.3	23.9-25.7
M1 L	37.46	37.32	47§	(36)	38	33.3; 31	34.3; 34.4	32.8; 33.8	25.8-32.1	26.9-30.6
M1 W	41.84	36.15	38§	38.5	37	34; 33	33; 34.1	34.2; 34.2	30.1-33.2	29.1-30.7
M1 H	24.82	24.97	25§						19.4-24.9	11.7-21.1
M2 L	48.48	47.36	50	(46)	45.5#	51	42; 41.6	54.3; 53	37.2-45.2	39.8-40.2
M2 W	39.34	38.35	41	42.4	35.0#	38	36.6; 37.9	37.3; 35.3	31.1-35.8	28.9-31.1
M2 H	30.75	31.87	36						19.3-28.9	24.3-24.4
M3 L	51.98	52.23		55	43.0#	59	52; 52.7	61.4; 61.3	39.9-46.2	42.2-44.7
M3 W	35.99	35.74		43.3	34.5#	35	36.1; 36.8	35; 25	24.3-32.3	27.1-31.6
M3 H	33.79	35.08							20.1-22.5	27.8-30.1
Di I3-C	20.51	23.28		18		21.0	14*		12.1-16.8	10.9
Di C-P1	33.4	37.61			72†		16*		30.1-41.7	35.4-35.9
Di P1-3	61.62	59.79			>60	51.0; 49.0	49*		24.1-43.8	43.9-47.8
P3-4 L	61.59	64.63		(58)			38.5; 39.2		37.2-39.8	35.1-39.1
M1-3 L	136.35	136.85		(131)		140	123; 124	144; 142.3	104.1-112.4	111.2-115.9
P3-M3 L	199.6	203.5		(186)		184			135.7-145.3	141.1-143.7

Specimens	Paracamelus gigas			Camelus knoblochi			C. ferus	C. dromedarius
Specimens	DLJ1610-04	Yushe¹⁾	Kossom Bougoudi²⁾	Luchka³⁾	Sengiley³⁾	Salawusu⁴⁾	c/o.20,705,676	ov25
m2 L		37-40	39.7	43	49.2		34.1-42.3	39.6-40.4
m2 W	(28)	23-26	23.7	31.4	28.4		21.8-26.1	19.1-20.2
m2 H	26.28						15.2-23.3	27.7-28.3
m3 L	62.97	54-58	54.4	73	72	65	54.1-57.3	52.1-56.3
m3 W	28.52	22-25	21.5	29.4	25.5		19.5-24.5	23.8
m3 H	26.27						18.9-20.4	26.1

Specimens	Paracamelus gigas			Camelus knoblochi			C. ferus	C. dromedarius
Specimens	DLJ1610-04	Yushe¹⁾	Kossom Bougoudi²⁾	Luchka³⁾	Sengiley³⁾	Salawusu⁴⁾	c/o.20,705,676	ov25
m2 L		37-40	39.7	43	49.2		34.1-42.3	39.6-40.4
m2 W	(28)	23-26	23.7	31.4	28.4		21.8-26.1	19.1-20.2
m2 H	26.28						15.2-23.3	27.7-28.3
m3 L	62.97	54-58	54.4	73	72	65	54.1-57.3	52.1-56.3
m3 W	28.52	22-25	21.5	29.4	25.5		19.5-24.5	23.8
m3 H	26.27						18.9-20.4	26.1