Eco-functional divergence of Crocuta and Pachycrocuta: a Huainan specimen reappraisal

doi:10.19615/j.cnki.2096-9899.250912

Abstract

Abstract:

Crocuta and Pachycrocuta are widely regarded as the most prevalent and emblematic hyenas across Eurasia during Quaternary. They are easily distinguished by their distinctive carnassial teeth. However, the disparities in non-carnassial elements are less pronounced and have received minimal attention in previous studies. This has resulted in erroneous identifications when dealing with fragmented specimens, particularly in cases where carnassial teeth are poorly preserved or absent. Such misidentifications have the potential to give rise to erroneous inferences regarding the paleozoogeography and biochronology of the animals in question. The bone-cracking hyena specimens from Huainan, Anhui (Tseng et al., 2008) are re-examined and re-evaluated here through a series of morphological comparisons and data analyses (univariate, bivariate, and multivariate analyses etc.). The results provide unequivocal confirmation that the specimens from Xiliexi are not spotted hyenas, but belong to Pachycrocuta perrieri instead. Conversely, the specimen from Dadingshan is the only genuine representative of the spotted hyena, which is supposed to be a possible earliest fossil record for Crocuta ultima thus far in China. Furthermore, the disparities in dentognathic morphology between Crocuta and Pachycrocuta are systematically summarized and analyzed, with an explanation of their eco-functional significance. The present study hypothesizes that Pachycrocuta retains a certain degree of active predatory capability, attributable to the robustness of the canine and the symphysis of the jaw, among other factors. This finding indicates that Pachycrocuta exhibits a form of flexible foraging behavior, combining opportunistic scavenging and active hunting in a manner analogous to Crocuta. Finally, the dynamic evolutionary history of hyenas in East China since Pleistocene has also been broadly reconstructed.

Key words: Huainan, Anhui, Pleistocene, re-examination, Pachycrocuta perrieri, Crocuta honanensis, Crocuta ultima, morphological comparison, statistical analysis

摘要：

斑鬣狗(Crocuta)和硕鬣狗(Pachycrocuta)是第四纪期间欧亚大陆最常见、最具代表性的鬣狗，仅由裂齿的形态便可加以区分。然而，二者非裂齿的形态差异却不甚明显，过往的研究也没有予以足够的重视。在面对破碎的标本时，尤其是在裂齿破损或缺失的情况下，时常会导致分类鉴定错误，并进一步导致对有关类群在古动物地理分布和生物年代学上的错误认识和推断。通过一系列的形态学比较和数据统计分析(单元、双元和多元分析等), 对曾志杰等(2008)研究发表的安徽淮南碎骨型鬣狗进行了重新研究和评估，结果表明西裂隙的标本并非斑鬣狗，而是属于佩里硕鬣狗(Pachycrocuta perrieri); 大顶山的标本才是斑鬣狗的真正代表，它可能是迄今中国境内最后斑鬣狗(Crocuta ultima)的最早化石记录。此外，系统总结并分析了斑鬣狗与硕鬣狗在齿颌形态上的差异，并阐述了相应的生态功能意义。研究认为硕鬣狗仍保留了一定的主动捕杀能力，其强壮的犬齿和下颌联合部等充分展示了这一点，在觅食行为上表现出与斑鬣狗类似的灵活弹性：既能机会腐食也能主动捕猎。最后，大致恢复和重现了自更新世以来，华东地区鬣狗的动态演化历史。

关键词: 安徽淮南, 更新世, 重新研究, 佩里硕鬣狗, 河南斑鬣狗, 最后斑鬣狗, 形态学比较, 统计分析

CLC Number:

Q915.874

LIU Jin-Yi, Zhijie Jack TSENG, JIN Chang-Zhu, ZHENG Long-Ting, SUN Cheng-Kai, TONG Hao-Wen. Eco-functional divergence of Crocuta and Pachycrocuta: a Huainan specimen reappraisal. Vertebrata Palasiatica, 2025, 63(4): 277-322.

刘金毅, 曾志杰, 金昌柱, 郑龙亭, 孙承凯, 同号文. 2025, 63(4): 277-322, 斑鬣狗(Crocuta)和硕鬣狗(Pachycrocuta)的生态功能之分化——对淮南标本的重新评估. 古脊椎动物学报.

Figures/Tables 13

References 114

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	Huainan, Anhui					Crocuta																Pachycrocuta
	XLX				DDS	crocuta				honanensis				spelaea				ultima				pyrenaica				perrieri
	*3	*4.1-5	*4.6	X	*0	n	range	X	s	n	range	X	s	n	range	X	s	n	range	X	s	n	range	X	s	n	range	X	s
c L	18.7	16.9	17.4	17.67		32	13.4-19	15.47	1.28	19	12.4-15.1	13.94	0.78	52	14.9-19.8	16.80	1.02	33	15.2-22	17.47	1.56	14	15.1-18.24	17.05	0.90	21	15.8-19	17.58	0.71
c W	15.1	14.1	14.3	14.5		35	11.2-13.6	12.23	0.58	18	10.1-13.5	11.68	0.92	52	12.5-15.8	13.68	0.70	32	12.6-15.7	14.31	0.80	14	11-14.9	12.90	0.99	21	13-16.3	14.26	0.73
p2 L	15.7	15.3		15.5	16.1	187	12.3-18.1	14.78	0.98	19	12.4-15.3	13.83	0.76	434	13.2-19.3	16.70	0.94	40	15.1-18.8	16.92	0.90	16	14.3-17.4	15.64	0.75	48	13.3-17.7	15.43	1.05
p2 W	11.1	9.9		10.5	11.1	76	8.2-12.7	10.33	0.92	19	8.3-10.5	9.49	0.54	383	10.4-14.4	12.40	0.80	39	10.1-14	12.26	0.95	16	9.4-10.4	9.80	0.30	47	9.1-12.9	10.55	0.80
p3 L	21.8			21.8	22.7	190	17.5-23	20.15	0.96	22	16.2-19.2	18.24	0.76	528	19.6-26.5	22.57	1.01	49	21.2-26	23.24	1.11	21	15.7-21.5	20.11	1.15	52	19.2-22.9	20.78	0.76
p3 W	14.5			14.5	16.6	74	12.6-16.2	14.54	0.84	20	12.4-14.3	13.45	0.58	511	13.7-19	16.51	0.82	48	14.2-18.8	16.85	0.89	20	10.7-13.9	12.94	0.74	49	12.2-16.6	14.51	0.88
p4 L	23.9			23.9	24.3	177	19-24.6	21.97	0.97	22	19-21.9	20.68	0.77	528	21-26.8	24.19	1.01	48	20.4-26.4	24.28	1.14	23	19.7-24.4	22.51	1.09	56	21.4-25.2	23.14	0.99
p4 W	14.9			14.9	14	74	10.7-14	12.75	0.76	22	11.7-13.6	12.73	0.50	522	12.6-17.3	15.07	0.74	49	12.7-17.1	15.11	0.86	21	9.3-13.8	12.88	0.92	50	13.1-16.11	14.54	0.76
m1 L		24.5		24.5		176	23.3-31.2	27.26	1.71	20	23.1-26.5	24.91	0.91	442	27.5-38.4	32.45	1.46	47	29.7-37.4	32.77	1.71	23	22.7-26.3	24.33	0.99	60	23-27.6	25.26	1.26
m1 W		13.0		13		84	9.8-14	11.62	0.75	19	10.8-12.3	11.52	0.48	442	11.5-15.9	13.87	0.69	46	11.7-15.1	13.94	0.77	24	10.8-12.82	11.83	0.56	53	11-14.81	12.82	0.81
m1tri L		20.2		20.2		87	21.7-28.2	25.20	1.53	18	19.3-22	21.03	0.79	428	25.5-33.5	29.30	1.27	35	27.2-34.7	29.65	1.69	19	18.1-21.2	19.64	0.90	49	19-22.8	20.91	1.02
p2 L /m1 L		0.62		0.62		171	0.47-0.62	0.54	0.03	16	0.5-0.63	0.56	0.03	162	0.43-0.62	0.52	0.03	29	0.43-0.58	0.52	0.03	10	0.59-0.76	0.65	0.05	37	0.52-0.68	0.61	0.04
p3 W /p3 L	0.67			0.67	0.73	74	0.63-0.79	0.71	0.03	20	0.68-0.8	0.74	0.03	505	0.6-0.83	0.73	0.03	48	0.67-0.79	0.73	0.03	20	0.61-0.71	0.64	0.02	49	0.58-0.78	0.70	0.04
p3 W /p4 W	0.97			0.97	1.19	71	1.05-1.22	1.14	0.03	19	0.99-1.11	1.06	0.03	299	0.85-1.2	1.10	0.04	40	1.0-1.2	1.11	0.04	20	0.95-1.15	1.01	0.04	44	0.91-1.11	1.01	0.03
m1tri L /m1 L		0.82		0.82		84	0.88-0.95	0.91	0.02	17	0.81-0.88	0.85	0.02	422	0.86-0.95	0.90	0.02	35	0.85-0.96	0.90	0.02	18	0.76-0.83	0.80	0.02	49	0.74-0.87	0.83	0.02
m1tri L /p4 L				0.85		83	1.03-1.3	1.14	0.06	18	0.9-1.06	1.01	0.04	179	1.08-1.37	1.21	0.05	24	1.11-1.28	1.20	0.05	13	0.79-1.0	0.87	0.06	35	0.85-1.02	0.90	0.04
C L		16.95		16.95		40	13.4-19	16.36	1.13	6	13-15.9	14.47	0.86	31	15.1-22.5	17.37	1.30	22	16-20.5	18.25	1.41	7	17-18.2	17.33	0.43	14	16.1-20.8	18.61	1.25
C W		13.1		13.1		41	10.2-14	11.87	0.81	6	10.9-11.9	11.35	0.33	30	11.7-15.7	13.46	0.98	22	12.5-15.2	13.92	0.69	7	12.4-13.1	12.66	0.24	11	12.4-15.5	13.65	0.84
P3 L		22.2		22.2		190	19-25.4	21.53	1.17	7	20-21.5	20.61	0.54	374	20.7-27.7	24.51	1.16	55	19.7-28.3	25.38	1.62	11	20.6-23.6	22.23	0.79	55	21-25.8	23.51	1.08
P3 W		16.8		16.8		80	14.2-18.7	16.56	0.99	7	14.1-16	15.17	0.57	360	15.4-20.5	18.21	0.91	55	15.8-21.7	19.05	1.34	8	13.73-15.9	15.09	0.67	46	14-18.8	16.58	1.05
P4 W		21.1		21.1		51	16.6-22.5	20.28	1.36	8	15-20.4	19.08	1.65	337	18-26	22.16	1.23	52	19.5-27.9	22.57	1.78	12	18.06-22.7	19.91	1.35	47	17.9-24.1	20.86	1.32
P4ps L		11		11		15	6.1-8.9	7.48	0.74	8	7.9-9.7	8.98	0.54	29	7.4-11.4	9.52	1.19	37	7-12.7	8.98	1.08	6	9.3-11.3	10.43	0.71	33	9.72-13.5	11.57	1.06
P3 W /P3 L		0.76		0.76		80	0.68-0.86	0.75	0.03	7	0.66-0.76	0.74	0.03	345	0.67-0.86	0.74	0.03	53	0.69-0.83	0.75	0.04	8	0.62-0.77	0.68	0.04	46	0.6-0.78	0.71	0.04

	Huainan, Anhui					Crocuta																Pachycrocuta
	XLX				DDS	crocuta				honanensis				spelaea				ultima				pyrenaica				perrieri
	*3	*4.1-5	*4.6	X	*0	n	range	X	s	n	range	X	s	n	range	X	s	n	range	X	s	n	range	X	s	n	range	X	s
c L	18.7	16.9	17.4	17.67		32	13.4-19	15.47	1.28	19	12.4-15.1	13.94	0.78	52	14.9-19.8	16.80	1.02	33	15.2-22	17.47	1.56	14	15.1-18.24	17.05	0.90	21	15.8-19	17.58	0.71
c W	15.1	14.1	14.3	14.5		35	11.2-13.6	12.23	0.58	18	10.1-13.5	11.68	0.92	52	12.5-15.8	13.68	0.70	32	12.6-15.7	14.31	0.80	14	11-14.9	12.90	0.99	21	13-16.3	14.26	0.73
p2 L	15.7	15.3		15.5	16.1	187	12.3-18.1	14.78	0.98	19	12.4-15.3	13.83	0.76	434	13.2-19.3	16.70	0.94	40	15.1-18.8	16.92	0.90	16	14.3-17.4	15.64	0.75	48	13.3-17.7	15.43	1.05
p2 W	11.1	9.9		10.5	11.1	76	8.2-12.7	10.33	0.92	19	8.3-10.5	9.49	0.54	383	10.4-14.4	12.40	0.80	39	10.1-14	12.26	0.95	16	9.4-10.4	9.80	0.30	47	9.1-12.9	10.55	0.80
p3 L	21.8			21.8	22.7	190	17.5-23	20.15	0.96	22	16.2-19.2	18.24	0.76	528	19.6-26.5	22.57	1.01	49	21.2-26	23.24	1.11	21	15.7-21.5	20.11	1.15	52	19.2-22.9	20.78	0.76
p3 W	14.5			14.5	16.6	74	12.6-16.2	14.54	0.84	20	12.4-14.3	13.45	0.58	511	13.7-19	16.51	0.82	48	14.2-18.8	16.85	0.89	20	10.7-13.9	12.94	0.74	49	12.2-16.6	14.51	0.88
p4 L	23.9			23.9	24.3	177	19-24.6	21.97	0.97	22	19-21.9	20.68	0.77	528	21-26.8	24.19	1.01	48	20.4-26.4	24.28	1.14	23	19.7-24.4	22.51	1.09	56	21.4-25.2	23.14	0.99
p4 W	14.9			14.9	14	74	10.7-14	12.75	0.76	22	11.7-13.6	12.73	0.50	522	12.6-17.3	15.07	0.74	49	12.7-17.1	15.11	0.86	21	9.3-13.8	12.88	0.92	50	13.1-16.11	14.54	0.76
m1 L		24.5		24.5		176	23.3-31.2	27.26	1.71	20	23.1-26.5	24.91	0.91	442	27.5-38.4	32.45	1.46	47	29.7-37.4	32.77	1.71	23	22.7-26.3	24.33	0.99	60	23-27.6	25.26	1.26
m1 W		13.0		13		84	9.8-14	11.62	0.75	19	10.8-12.3	11.52	0.48	442	11.5-15.9	13.87	0.69	46	11.7-15.1	13.94	0.77	24	10.8-12.82	11.83	0.56	53	11-14.81	12.82	0.81
m1tri L		20.2		20.2		87	21.7-28.2	25.20	1.53	18	19.3-22	21.03	0.79	428	25.5-33.5	29.30	1.27	35	27.2-34.7	29.65	1.69	19	18.1-21.2	19.64	0.90	49	19-22.8	20.91	1.02
p2 L /m1 L		0.62		0.62		171	0.47-0.62	0.54	0.03	16	0.5-0.63	0.56	0.03	162	0.43-0.62	0.52	0.03	29	0.43-0.58	0.52	0.03	10	0.59-0.76	0.65	0.05	37	0.52-0.68	0.61	0.04
p3 W /p3 L	0.67			0.67	0.73	74	0.63-0.79	0.71	0.03	20	0.68-0.8	0.74	0.03	505	0.6-0.83	0.73	0.03	48	0.67-0.79	0.73	0.03	20	0.61-0.71	0.64	0.02	49	0.58-0.78	0.70	0.04
p3 W /p4 W	0.97			0.97	1.19	71	1.05-1.22	1.14	0.03	19	0.99-1.11	1.06	0.03	299	0.85-1.2	1.10	0.04	40	1.0-1.2	1.11	0.04	20	0.95-1.15	1.01	0.04	44	0.91-1.11	1.01	0.03
m1tri L /m1 L		0.82		0.82		84	0.88-0.95	0.91	0.02	17	0.81-0.88	0.85	0.02	422	0.86-0.95	0.90	0.02	35	0.85-0.96	0.90	0.02	18	0.76-0.83	0.80	0.02	49	0.74-0.87	0.83	0.02
m1tri L /p4 L				0.85		83	1.03-1.3	1.14	0.06	18	0.9-1.06	1.01	0.04	179	1.08-1.37	1.21	0.05	24	1.11-1.28	1.20	0.05	13	0.79-1.0	0.87	0.06	35	0.85-1.02	0.90	0.04
C L		16.95		16.95		40	13.4-19	16.36	1.13	6	13-15.9	14.47	0.86	31	15.1-22.5	17.37	1.30	22	16-20.5	18.25	1.41	7	17-18.2	17.33	0.43	14	16.1-20.8	18.61	1.25
C W		13.1		13.1		41	10.2-14	11.87	0.81	6	10.9-11.9	11.35	0.33	30	11.7-15.7	13.46	0.98	22	12.5-15.2	13.92	0.69	7	12.4-13.1	12.66	0.24	11	12.4-15.5	13.65	0.84
P3 L		22.2		22.2		190	19-25.4	21.53	1.17	7	20-21.5	20.61	0.54	374	20.7-27.7	24.51	1.16	55	19.7-28.3	25.38	1.62	11	20.6-23.6	22.23	0.79	55	21-25.8	23.51	1.08
P3 W		16.8		16.8		80	14.2-18.7	16.56	0.99	7	14.1-16	15.17	0.57	360	15.4-20.5	18.21	0.91	55	15.8-21.7	19.05	1.34	8	13.73-15.9	15.09	0.67	46	14-18.8	16.58	1.05
P4 W		21.1		21.1		51	16.6-22.5	20.28	1.36	8	15-20.4	19.08	1.65	337	18-26	22.16	1.23	52	19.5-27.9	22.57	1.78	12	18.06-22.7	19.91	1.35	47	17.9-24.1	20.86	1.32
P4ps L		11		11		15	6.1-8.9	7.48	0.74	8	7.9-9.7	8.98	0.54	29	7.4-11.4	9.52	1.19	37	7-12.7	8.98	1.08	6	9.3-11.3	10.43	0.71	33	9.72-13.5	11.57	1.06
P3 W /P3 L		0.76		0.76		80	0.68-0.86	0.75	0.03	7	0.66-0.76	0.74	0.03	345	0.67-0.86	0.74	0.03	53	0.69-0.83	0.75	0.04	8	0.62-0.77	0.68	0.04	46	0.6-0.78	0.71	0.04

	Huainan, Anhui		Crocuta																Pachycrocuta
	XLX	DDS	crocuta				honanensis				spelaea				ultima				pyrenaica				perrieri
	V15163	V15160	n	range	X	S	n	range	X	S	n	range	X	S	n	range	X	S	n	range	X	S	n	range	X	S
1	43.8	41.2	16	28-38.5	32.36	2.74	17	30.8-39.9	36.19	3.03	32	29.1-45	38.41	4.28	21	36.6-48.8	43.90	3.00	6	36.1-41.4	38.93	2.11	12	37.7-44.2	41.04	1.87
2	48.8		16	33.8-42.8	38.03	2.65	18	31.8-44.8	40.30	3.56	29	32.7-52.7	43.06	5.91	21	37.9-54.7	48.00	3.97	6	41.8-45	43.38	1.09	12	38.5-50.8	43.88	3.69
3	23.7	20.7	16	15.8-19.3	17.37	1.00	18	16.1-20.8	18.30	1.38	36	17.8-25.2	21.65	1.79	20	19.6-28.2	22.77	1.90	5	18.4-21.9	19.50	1.35	13	18.1-23.8	21.06	1.39

	Huainan, Anhui		Crocuta																Pachycrocuta
	XLX	DDS	crocuta				honanensis				spelaea				ultima				pyrenaica				perrieri
	V15163	V15160	n	range	X	S	n	range	X	S	n	range	X	S	n	range	X	S	n	range	X	S	n	range	X	S
1	43.8	41.2	16	28-38.5	32.36	2.74	17	30.8-39.9	36.19	3.03	32	29.1-45	38.41	4.28	21	36.6-48.8	43.90	3.00	6	36.1-41.4	38.93	2.11	12	37.7-44.2	41.04	1.87
2	48.8		16	33.8-42.8	38.03	2.65	18	31.8-44.8	40.30	3.56	29	32.7-52.7	43.06	5.91	21	37.9-54.7	48.00	3.97	6	41.8-45	43.38	1.09	12	38.5-50.8	43.88	3.69
3	23.7	20.7	16	15.8-19.3	17.37	1.00	18	16.1-20.8	18.30	1.38	36	17.8-25.2	21.65	1.79	20	19.6-28.2	22.77	1.90	5	18.4-21.9	19.50	1.35	13	18.1-23.8	21.06	1.39

	PC 1	PC 2	PC 3	PC 4	PC 5	PC 6	PC 7	PC 8	PC 9	PC 10	PC 11
c L	0.12857	0.52793	-0.51798	-0.4058	-0.1319	0.42808	-0.10251	0.18228	-0.14816	0.0031197	0.073732
c W	0.11519	0.41137	-0.2895	0.061824	0.19053	-0.40619	0.5479	-0.3734	0.27855	0.028902	0.10265
p2 L	0.19562	0.21439	0.60374	-0.35355	0.019271	0.090964	0.19582	-0.23824	-0.34804	0.44345	0.068881
p2 W	0.20677	0.14561	0.35952	-0.015387	0.3512	0.43567	0.23062	0.25402	0.47061	-0.33123	-0.21097
p3 L	0.26228	0.23653	0.21418	-0.34127	-0.051144	-0.48927	-0.49421	-0.095557	0.074532	-0.45689	-0.057054
p3 W	0.22325	0.10562	-0.01265	0.26965	0.33499	-0.22746	0.19058	0.45676	-0.63522	-0.22854	0.038555
p4 L	0.18885	0.25885	0.20094	0.31979	-0.72407	-0.12879	0.13943	0.37982	0.18512	0.10872	0.063554
p4 W	0.1571	0.28103	-0.066492	0.34547	0.27647	-0.061683	-0.42603	0.025896	0.09434	0.51327	-0.48766
m1 L	0.56868	-0.22759	-0.14619	0.18498	-0.26791	0.2328	0.1042	-0.44614	-0.21226	-0.22508	-0.37099
m1 W	0.16425	0.16438	0.10107	0.44869	0.12938	0.26572	-0.32376	-0.2717	0.024521	-0.081191	0.67917
m1tri L	0.59986	-0.43932	-0.17806	-0.24483	0.13613	-0.13176	-0.023896	0.26553	0.24489	0.31513	0.29878
Eigenvalue	29.9412	5.66701	0.970645	0.672578	0.502124	0.342199	0.295265	0.200275	0.181543	0.150677	0.108996
% variance	76.708	14.519	2.4868	1.7231	1.2864	0.8767	0.75646	0.5131	0.46511	0.38603	0.27924