青藏高原与横断山被子植物区系演化的 细胞地理学特征

doi:10.17520/biods.2016281

摘要/Abstract

摘要：

青藏高原与横断山集特殊的地理位置和复杂的地形地势以及高度丰富的植物区系成分于一体, 成为世界上研究生物多样性起源与演化的热点地区之一。我们对这一地区被子植物的染色体数据进行了全面的统计分析, 结果表明新多倍体在该地区只约占23%的比例, 远低于其他高山地区, 这可能是由于这一地区特殊的地质历史或者复杂的地形和生境等因素所导致的。低基数的二倍体占近一半的比例(43.3%), 说明了二倍体水平上的染色体结构和核型进化也是本地区物种分化的另一重要机制。古多倍体, 即高基数二倍体, 也占有较大的成分(33.7%)。本地区物种在细胞学遗传演化途径上的多样性与这一地区极高的物种多样性和特有性可能存在一定的联系。通过对青藏高原及横断山地区被子植物区系进化中染色体演化机制的研究, 为探讨东亚以及北半球其他植物区系物种进化、高山植物区系起源和演化等许多重大问题提供了有益的帮助。

关键词: 青藏高原, 细胞地理, 染色体进化, 多倍体, 被子植物

Abstract

The Qinghai-Tibet Plateau and Hengduan Mountains is an important hotspot to study the origin and evolution of plant biodiversity, which is characterized by high species richness and endemism, as well as a unique location and complicated topography. Based on a comprehensive survey of the chromosome numbers of angiosperms from the Qinghai-Tibet Plateau and Hengduan Mountains, our results indicated that the frequency of neopolyploids (23%) was much lower than expected. This was likely caused by the unique history, geology, and complicated topography and habitat. Another mechanism of chromosomal evolution is on the diploidy level (ca. 43.3%), with variations of chromosomal restructure and karyotype. The number of paleopolyploids (diploids with high base numbers) was large in plants from this mountain area (33.7%). The diversity of chromosomal evolution patterns of angiosperms from this region may be correlated to the high level of species richness and endemism. This study provides useful insight into our understanding of speciation, the origins of alpine flora and other important issues in eastern Asia and other regions in the Northern Hemisphere.

Key words: Qinghai-Tibet Plateau, cytogeography, chromosomal evolution, polyploidy, angiosperm

王家坚, 彭智邦, 孙航, 聂泽龙, 孟盈 (2017) 青藏高原与横断山被子植物区系演化的细胞地理学特征. 生物多样性, 25, 218-225. DOI: 10.17520/biods.2016281.

Jia-Jian Wang, Zhi-Bang Peng, Hang Sun, Ze-Long Nie, Ying Meng (2017) Cytogeographic patterns of angiosperms flora of the Qinghai-Tibet Plateau and Hengduan Mountains. Biodiversity Science, 25, 218-225. DOI: 10.17520/biods.2016281.

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链接本文: https://www.biodiversity-science.net/CN/10.17520/biods.2016281

https://www.biodiversity-science.net/CN/Y2017/V25/I2/218

图/表 3

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	报道条数 No. of counts	种数 No. of species (%)
	报道条数 No. of counts	总数 Total	新多倍体 Neopolyploids	古多倍体 Paleopolyploids	二倍体 Diploids
分布型 Areal-type
世界分布 Cosmopolitan	177	139	38 (27.3)	21 (15.1)	80 (57.6)
泛热带分布 Pantropic	37	33	3 (9.1)	15 (45.5)	15 (45.5)
东亚(热带、亚热带)及热带南美间断 East Asia (Tropical & Subtropical) & Tropical South America disjuncted	1	1	0 (0.0)	1 (100)	0 (0.0)
旧世界热带分布 Old World Tropics	111	83	26 (31.3)	28 (33.7)	29 (34.9)
旧世界热带分布 Old World Tropics	111	83	26 (31.3)	28 (33.7)	29 (34.9)	热带亚洲至热带大洋洲分布 Tropical Asia to Tropical Australasia Oceania	5	5	2 (40.0)	2 (40.0)	1 (20.0)
热带亚洲至热带非洲分布 Tropical Asia to Tropical Africa	18	11	2 (18.2)	4 (36.4)	5 (45.5)	热带亚洲至热带大洋洲分布 Tropical Asia to Tropical Australasia Oceania	5	5	2 (40.0)	2 (40.0)	1 (20.0)
热带亚洲(印度-马来西亚)分布 Tropical Southeast Asia to Indo-Malaya & Tropical Southwest Pacific Islands	43	26	7 (26.9)	16 (61.5)	3 (11.5)
北温带分布 North Temperate	838	516	118 (22.9)	185 (35.8)	213 (41.3)
东亚和北美洲间断分布 East Asia & North America disjuncted	18	12	2 (16.7)	6 (50)	4 (33.3)
旧世界温带 Old World Temperate	11	11	2 (18.2)	3 (27.3)	6 (54.5)
温带亚洲分布 Temperate Asia	29	21	5 (23.8)	6 (28.6)	10 (47.6)
地中海区、西亚至中亚分布 Mediterranean & West to Central Asia	1	1	0 (0.0)	0 (0.0)	1 (100.0)
中亚分布 Central Asia	21	17	3 (17.6)	6 (35.3)	8 (47.1)
东亚分布 East Asia	150	101	20 (19.8)	36 (35.6)	45 (44.6)
中国特有分布 Endemic to China	31	24	2 (8.3)	9 (37.5)	13 (54.2)
小计 Subtotal	1,491	1,001	230 (23.0)	338 (33.7)	433 (43.3)
生活型 Life form
一年生草本 Annual herb	139	96	19 (19.8)	18 (18.8)	59 (61.5)
多年生草本 Perennial herb	1,208	796	182 (22.9)	289 (36.3)	325 (40.8)
木本 Woody	144	109	29 (26.6)	31 (28.4)	49 (45.0)
小计 Subtotal	1,491	1,001	230 (23.0)	338 (33.7)	433 (43.3)
海拔 Altitude
<1,000 m	51	38	13 (34.2)	7 (18.4)	18 (47.4)
1,000-2,000 m	145	101	28 (27.7)	25 (24.8)	48 (47.5)
2,000-3,000 m	222	156	24 (15.4)	77 (49.4)	55 (35.3)
3,000-4,000 m	411	255	65 (25.5)	102 (40)	88 (34.5)
4,000-5,000 m	196	122	27 (22.1)	35 (28.7)	60 (49.2)
> 5,000 m	9	6	3 (50.0)	1 (16.7)	2 (33.3)
小计 Subtotal	1,034	678	160 (23.6)	247 (36.4)	271 (40.0)

	报道条数 No. of counts	种数 No. of species (%)
	报道条数 No. of counts	总数 Total	新多倍体 Neopolyploids	古多倍体 Paleopolyploids	二倍体 Diploids
分布型 Areal-type
世界分布 Cosmopolitan	177	139	38 (27.3)	21 (15.1)	80 (57.6)
泛热带分布 Pantropic	37	33	3 (9.1)	15 (45.5)	15 (45.5)
东亚(热带、亚热带)及热带南美间断 East Asia (Tropical & Subtropical) & Tropical South America disjuncted	1	1	0 (0.0)	1 (100)	0 (0.0)
旧世界热带分布 Old World Tropics	111	83	26 (31.3)	28 (33.7)	29 (34.9)
旧世界热带分布 Old World Tropics	111	83	26 (31.3)	28 (33.7)	29 (34.9)	热带亚洲至热带大洋洲分布 Tropical Asia to Tropical Australasia Oceania	5	5	2 (40.0)	2 (40.0)	1 (20.0)
热带亚洲至热带非洲分布 Tropical Asia to Tropical Africa	18	11	2 (18.2)	4 (36.4)	5 (45.5)	热带亚洲至热带大洋洲分布 Tropical Asia to Tropical Australasia Oceania	5	5	2 (40.0)	2 (40.0)	1 (20.0)
热带亚洲(印度-马来西亚)分布 Tropical Southeast Asia to Indo-Malaya & Tropical Southwest Pacific Islands	43	26	7 (26.9)	16 (61.5)	3 (11.5)
北温带分布 North Temperate	838	516	118 (22.9)	185 (35.8)	213 (41.3)
东亚和北美洲间断分布 East Asia & North America disjuncted	18	12	2 (16.7)	6 (50)	4 (33.3)
旧世界温带 Old World Temperate	11	11	2 (18.2)	3 (27.3)	6 (54.5)
温带亚洲分布 Temperate Asia	29	21	5 (23.8)	6 (28.6)	10 (47.6)
地中海区、西亚至中亚分布 Mediterranean & West to Central Asia	1	1	0 (0.0)	0 (0.0)	1 (100.0)
中亚分布 Central Asia	21	17	3 (17.6)	6 (35.3)	8 (47.1)
东亚分布 East Asia	150	101	20 (19.8)	36 (35.6)	45 (44.6)
中国特有分布 Endemic to China	31	24	2 (8.3)	9 (37.5)	13 (54.2)
小计 Subtotal	1,491	1,001	230 (23.0)	338 (33.7)	433 (43.3)
生活型 Life form
一年生草本 Annual herb	139	96	19 (19.8)	18 (18.8)	59 (61.5)
多年生草本 Perennial herb	1,208	796	182 (22.9)	289 (36.3)	325 (40.8)
木本 Woody	144	109	29 (26.6)	31 (28.4)	49 (45.0)
小计 Subtotal	1,491	1,001	230 (23.0)	338 (33.7)	433 (43.3)
海拔 Altitude
<1,000 m	51	38	13 (34.2)	7 (18.4)	18 (47.4)
1,000-2,000 m	145	101	28 (27.7)	25 (24.8)	48 (47.5)
2,000-3,000 m	222	156	24 (15.4)	77 (49.4)	55 (35.3)
3,000-4,000 m	411	255	65 (25.5)	102 (40)	88 (34.5)
4,000-5,000 m	196	122	27 (22.1)	35 (28.7)	60 (49.2)
> 5,000 m	9	6	3 (50.0)	1 (16.7)	2 (33.3)
小计 Subtotal	1,034	678	160 (23.6)	247 (36.4)	271 (40.0)

科名 Family	属名 Genus	报道条数 No. of counts	种数 No. of species (%)
科名 Family	属名 Genus	报道条数 No. of counts	总数 Total	新多倍体 Neopolyploids	古多倍体 Paleopolyploids	二倍体 Diploids
伞形科 Apiaceae	柴胡属 Bupleurum	15	12	1 (8.3)	0 (0.0)	11 (91.7)
伞形科 Apiaceae	独活属 Heracleum	34	16	2 (12.5)	14 (87.5)	0 (0.0)
伞形科 Apiaceae	藁本属 Ligusticum	12	11	2 (18.2)	9 (81.8)	0 (0.0)
天门冬科 Asparagaceae	黄精属 Polygonatum	27	18	3 (16.7)	14 (77.8)	1 (5.6)
菊科 Asteraceae	香青属 Anaphalis	37	18	15 (83.3)	3 (16.7)	0 (0.0)
菊科 Asteraceae	火绒草属 Leontopodium	20	13	7 (53.8)	6 (46.2)	0 (0.0)
菊科 Asteraceae	橐吾属 Ligularia	20	17	1 (5.9)	16 (94.1)	0 (0.0)
菊科 Asteraceae	风毛菊属 Saussurea	50	41	2 (4.9)	39 (95.1)	0 (0.0)
紫葳科 Bignoniaceae	角蒿属 Incarvillea	15	12	0 (0.0)	12 (100.0)	0 (0.0)
桔梗科 Campanulaceae	蓝钟花属 Cyananthus	23	14	2 (14.3)	0 (0.0)	12 (85.7)
豆科 Fabaceae	黄耆属 Astragalus	12	12	2 (16.7)	0 (0.0)	10 (83.3)
龙胆科 Gentianaceae	龙胆属 Gentiana	55	43	7 (16.3)	18 (41.9)	18 (41.9)
鸢尾科 Iridaceae	鸢尾属 Iris	13	10	0 (0.0)	10 (100.0)	0 (0.0)
百合科 Liliaceae	葱属 Allium	91	35	14 (40.0)	3 (8.6)	18 (51.4)
百合科 Liliaceae	百合属 Lilium	69	25	2 (8.0)	23 (92.0)	0 (0.0)
百合科 Liliaceae	豹子花属 Nomocharis	23	11	1 (9.1)	10 (90.9)	0 (0.0)
百合科 Liliaceae	重楼属 Paris	27	18	1 (5.6)	0 (0.0)	17 (94.4)
列当科 Orobanchaceae	马先蒿属 Pedicularis	21	21	0 (0.0)	0 (0.0)	21 (100.0)
禾本科 Poaceae	鹅观草属 Roegneria	26	13	13 (100.0)	0 (0.0)	0 (0.0)
报春花科 Primulaceae	报春花属 Primula	25	18	0 (0.0)	13 (72.2)	5 (27.8)
毛茛科 Ranunculaceae	乌头属 Aconitum	71	45	9 (20.0)	0 (0.0)	36 (80.0)
毛茛科 Ranunculaceae	银莲花属 Anemone	11	10	1 (10.0)	0 (0.0)	9 (90.0)
毛茛科 Ranunculaceae	翠雀属 Delphinium	100	56	2 (3.6)	0 (0.0)	54 (96.4)
毛茛科 Ranunculaceae	毛茛属 Ranunculus	17	15	10 (66.7)	0 (0.0)	5 (33.3)
蔷薇科 Rosaceae	苹果属 Malus	22	11	6 (54.5)	5 (45.5)	0 (0.0)
蔷薇科 Rosaceae	悬钩子属 Rubus	29	27	11 (40.7)	0 (0.0)	16 (59.3)
虎耳草科 Saxifragaceae	虎耳草属 Saxifraga	10	10	4 (40.0)	2 (20.0)	4 (40.0)

科名 Family	属名 Genus	报道条数 No. of counts	种数 No. of species (%)
科名 Family	属名 Genus	报道条数 No. of counts	总数 Total	新多倍体 Neopolyploids	古多倍体 Paleopolyploids	二倍体 Diploids
伞形科 Apiaceae	柴胡属 Bupleurum	15	12	1 (8.3)	0 (0.0)	11 (91.7)
伞形科 Apiaceae	独活属 Heracleum	34	16	2 (12.5)	14 (87.5)	0 (0.0)
伞形科 Apiaceae	藁本属 Ligusticum	12	11	2 (18.2)	9 (81.8)	0 (0.0)
天门冬科 Asparagaceae	黄精属 Polygonatum	27	18	3 (16.7)	14 (77.8)	1 (5.6)
菊科 Asteraceae	香青属 Anaphalis	37	18	15 (83.3)	3 (16.7)	0 (0.0)
菊科 Asteraceae	火绒草属 Leontopodium	20	13	7 (53.8)	6 (46.2)	0 (0.0)
菊科 Asteraceae	橐吾属 Ligularia	20	17	1 (5.9)	16 (94.1)	0 (0.0)
菊科 Asteraceae	风毛菊属 Saussurea	50	41	2 (4.9)	39 (95.1)	0 (0.0)
紫葳科 Bignoniaceae	角蒿属 Incarvillea	15	12	0 (0.0)	12 (100.0)	0 (0.0)
桔梗科 Campanulaceae	蓝钟花属 Cyananthus	23	14	2 (14.3)	0 (0.0)	12 (85.7)
豆科 Fabaceae	黄耆属 Astragalus	12	12	2 (16.7)	0 (0.0)	10 (83.3)
龙胆科 Gentianaceae	龙胆属 Gentiana	55	43	7 (16.3)	18 (41.9)	18 (41.9)
鸢尾科 Iridaceae	鸢尾属 Iris	13	10	0 (0.0)	10 (100.0)	0 (0.0)
百合科 Liliaceae	葱属 Allium	91	35	14 (40.0)	3 (8.6)	18 (51.4)
百合科 Liliaceae	百合属 Lilium	69	25	2 (8.0)	23 (92.0)	0 (0.0)
百合科 Liliaceae	豹子花属 Nomocharis	23	11	1 (9.1)	10 (90.9)	0 (0.0)
百合科 Liliaceae	重楼属 Paris	27	18	1 (5.6)	0 (0.0)	17 (94.4)
列当科 Orobanchaceae	马先蒿属 Pedicularis	21	21	0 (0.0)	0 (0.0)	21 (100.0)
禾本科 Poaceae	鹅观草属 Roegneria	26	13	13 (100.0)	0 (0.0)	0 (0.0)
报春花科 Primulaceae	报春花属 Primula	25	18	0 (0.0)	13 (72.2)	5 (27.8)
毛茛科 Ranunculaceae	乌头属 Aconitum	71	45	9 (20.0)	0 (0.0)	36 (80.0)
毛茛科 Ranunculaceae	银莲花属 Anemone	11	10	1 (10.0)	0 (0.0)	9 (90.0)
毛茛科 Ranunculaceae	翠雀属 Delphinium	100	56	2 (3.6)	0 (0.0)	54 (96.4)
毛茛科 Ranunculaceae	毛茛属 Ranunculus	17	15	10 (66.7)	0 (0.0)	5 (33.3)
蔷薇科 Rosaceae	苹果属 Malus	22	11	6 (54.5)	5 (45.5)	0 (0.0)
蔷薇科 Rosaceae	悬钩子属 Rubus	29	27	11 (40.7)	0 (0.0)	16 (59.3)
虎耳草科 Saxifragaceae	虎耳草属 Saxifraga	10	10	4 (40.0)	2 (20.0)	4 (40.0)

科名 Family	属数 No. of genus	报道条数 No. of counts	种数 No. of species (%)
科名 Family	属数 No. of genus	报道条数 No. of counts	总数 Total	新多倍体 Neopolyploids	古多倍体 Paleopolyploids	二倍体 Diploids
伞形科 Apiaceae	13	100	67	7 (10.4)	40 (59.7)	20 (29.9)
天南星科 Araceae	6	19	15	7 (46.7)	7 (46.7)	1 (6.7)
天门冬科 Asparagaceae	4	54	30	5 (16.7)	24 (80.0)	1 (3.3)
菊科 Asteraceae	28	200	149	35 (23.5)	75 (50.3)	39 (26.2)
紫葳科 Bignoniaceae	1	15	12	0 (0.0)	12 (100.0)	0 (0.0)
十字花科 Brassicaceae	12	29	22	3 (13.6)	0 (0.0)	19 (86.4)
桔梗科 Campanulaceae	3	28	18	3 (16.7)	2 (11.1)	13 (72.2)
豆科 Fabaceae	14	67	58	4 (6.9)	3 (5.2)	51 (87.9)
龙胆科 Gentianaceae	7	70	58	11 (19.0)	21 (36.2)	26 (44.8)
鸢尾科 Iridaceae	1	13	10	0 (0.0)	10 (100.0)	0 (0.0)
百合科 Liliaceae	16	251	114	27 (23.7)	49 (43.0)	38 (33.3)
兰科 Orchidaceae	12	37	26	7 (26.9)	19 (73.1)	0 (0.0)
列当科 Orobanchaceae	1	21	21	0 (0.0)	0 (0.0)	21 (100.0)
禾本科 Poaceae	6	47	27	25 (92.6)	1 (3.7)	1 (3.7)
蓼科 Polygonaceae	2	17	12	3 (25.0)	6 (50.0)	3 (25.0)
报春花科 Primulaceae	3	40	24	2 (8.3)	13 (54.2)	9 (37.5)
毛茛科 Ranunculaceae	22	247	159	29 (18.2)	1 (0.6)	129 (81.1)
蔷薇科 Rosaceae	6	61	47	19 (40.4)	8 (17.0)	20 (42.6)
虎耳草科 Saxifragaceae	5	25	23	10 (43.5)	3 (13.0)	10 (43.5)
茄科 Solanaceae	7	14	11	9 (81.8)	1 (9.1)	1 (9.1)