辣椒的起源、进化与栽培历史

doi:10.16420/j.issn.0513-353x.2021-0853

摘要/Abstract

摘要：

辣椒起源于玻利维亚中南部年降雨量不到500 mm的半干旱区,属亚热带无霜区,最初原始野生种为多年生草本植物。野生辣椒的利用可追溯到8 000 ~ 7 500年以前,早期辣椒依靠飞鸟传播种子,生长区域从发源地玻利维亚逐渐扩大到南美洲、中美洲,再到北美洲西南部,在不同生态区进化产生10多个栽培种的近缘野生种和约20个非近缘野生种。辣椒栽培种由共同祖先Capsicum chacoense进化而来,紫花祖先迁移到安第斯高地,进化产生了绒毛辣椒（Capsicum pubescens）;白花祖先迁移到玻利维亚南部相对干燥的地区进化产生下垂辣椒（Capsicum baccatum）,继续迁移到潮湿的亚马逊盆地,进化产生了一年生辣椒、灌木辣椒和中国辣椒的共同祖先。共同祖先继续向外迁移,在墨西哥和中美洲北部进化产生了一年生辣椒,在加勒比地区进化产生了灌木辣椒,在亚马逊河流域北部谷地进化产生了中国辣椒。辣椒的驯化是将野生种从原产地移出进行人工栽培开始,将易脱落、果实小、色泽单一、果实朝上的野生种,改变成不易脱落、果实朝下、肥大化及形状、颜色多样化、经济效益好的栽培种。一年生辣椒在墨西哥和中美洲最早进化,已有6 000多年的栽培历史,其他4个栽培种至少有4 000年的栽培历史,是美洲最古老的栽培植物之一。

关键词: 辣椒, 起源, 进化, 驯化, 野生种, 栽培种, 栽培历史

Abstract:

The Pepper originated in the semi-arid area in central and southern Bolivia where the annual rainfall is less than 500 mm. This is a subtropical frost-free area. The original wild species are perennial herbs. According to the analysis and study of the starch granules,the Capsicum starch microfossils have been found dating from 8 000-7 500 years ago. Over thousands of years,Capsicum seeds spread out from Bolivia with the help of birds and indigenous peoples,thus spreading throughout South and Central America,then to the southwest of North America. In different ecological regions,more than 10 wild relatives and about 20 wild non-relatives of cultivated species have evolved. Capsicum cultivars evolved from the common ancestor Capsicum chacoense. The purple flower ancestors migrated to the Andes highlands and evolved to Capsicum pubescens. And the white flower ancestors migrated to the relatively dry area of southern Bolivia to evolve to Capsicum baccatum,continuing to migrate to the humid Amazon basin and evolve to the common ancestors of Capsicum annuum,Capsicum frutescens L. and Capsicum chinense Jacquin. They continued to migrate outward,evolving Capsicum annuum in Mexico and northern Central America,Capsicum frutescens L. in the Caribbean,and Capsicum chinense Jacquin in the northern valleys of the Amazon Basin. The pepper domestication started by removing the wild species from the original area for artificial cultivation. The wild species with the fruit small,single-color,upward-facing and easily falling off were domesticated into cultivated species with the fruit succulent,various-shaped,colorful,downward-facing,and not easy falling off,i.e. good economic benefits. Capsicum annuum was the first cultivated species domesticated in Mexico and Central America more than 6 000 years ago,and the other four cultivated species had been cultivated for at least 4 000 years,which are the oldest cultivated plants in America.

Key words: pepper, origin, evolution, domestication, wild species, cultivated species, cultivation

中图分类号:

S641.3

邹学校, 朱凡. 辣椒的起源、进化与栽培历史[J]. 园艺学报, 2022, 49(6): 1371-1381.

ZOU Xuexiao, ZHU Fan. Origin,Evolution and Cultivation History of the Pepper[J]. Acta Horticulturae Sinica, 2022, 49(6): 1371-1381.

图/表 1

参考文献 50

[1]	Bailey L H. 1923. The cultivated evergreens. London:Macmillan:434.
[2]	Bosland P W. 1996. Capsicums:innovative uses of an ancient crop// Janick J. Progress in new crops. Arlington, VA: ASHS Press,479-487.
[3]	Dai Xiong-ze. 2008. Essay on the origin and spread of pepper. Journal of China Capsicum,(3):48-49. (in Chinese)
	戴雄泽. 2008. 漫话辣椒的起源和传播. 辣椒杂志,(3):48-49.
[4]	de Candolle A. 1886. Origin of cultivated plants. New York: Hafner Public Co 1964 reprint.
[5]	Dickau R. 2005. Resource use,crop dispersals,and the transition to agriculture in prehistoric Panama:Evidence from starch grains and macroremains. Romanian Journal of Morphology and Embryology: 46 (2):113-116.
[6]	Eshbaugh W H. 1977. The taxonomy of the genus Capsicum- Solanaceae// Pochard E. Capsicum 77. Comptes Rendus 3me Congrès EUCARPIA Piment,Avignon-Montfavet,France: 13-26.
[7]	Eshbaugh W H. 2012. The taxonomy of the genus Capsicum// Russo V M. Peppers:botany,production and uses. CAB Internaional:14-28.
[8]	Guo Jia-zhen, Mao Sheng-li, Huang San-wen. 2008. Pepper// Zhu De-wei. Crops and their wild relatives in China. Beijing: China Agriculture Press:704-746. (in Chinese)
	郭家珍, 毛胜利, 黄三文. 2008. 辣椒//朱德蔚. 中国作物及其野生近缘植物(上). 北京: 中国农业出版社:704-746.
[9]	Heiser C B, Smith P G. 1953. The cultivated capsicum peppers. Economic Botany, 7 (3):214-227. doi: 10.1007/BF02984948 URL
[10]	Heiser C. 1971. Notes on some species of Solarium(sect. Leptostemonum)in Latin America. Baileya, 18:59-65.
[11]	Hoshikawa Kiyochika. 1981. The origin and spread of cultivated plants. Duan Chuang-de,Ding Fa-yuan trans trans. Zhengzhou: Henan Science and Technology Press:136-137. (in Chinese)
	星川清亲. 1981. 栽培植物的起源与传播. 段传德,丁法元,译. 郑州: 河南科学技术出版社:136-137.
[12]	International Board for Plant Genetic Pesources (IBPGR). 1985. Forages for mediterranean and adjacent arid/semi-arid areas.
[13]	Jensen R J, McLeod M J, Eshbaugh W H, Gutiman S I. 1979. Numerical taxonomic analyses of allozymic variation in Capsicum (Solanaceae). Taxon, 28:315-327. doi: 10.2307/1219739 URL
[14]	Kim S, Park M, Yeom S I, Kim Y M, Lee J M, Lee H A, Seo E, Choi J, Cheong K, Kim K T, Jung K, Lee G W, Oh S K, Bae C, Kim S B, Lee H Y, Kim S Y, Kim M S, Kang B C, Jo Y D, Yang H B, Jeong H J, Kang W H, Kwon J K, Shin C, Lim J Y, Park J H, Huh J H, Kim J S, Kim B D, Cohen O, Paran I, Suh M C, Lee S B, Kim Y K, Shin Y, Noh S J, Park J, Seo Y S, Kwon S Y, Kim H A, Park J M, Kim H J, Choi S B, Bosland P W, Reeves G, Jo S H, Lee B W, Cho H T, Choi H S, Lee M S, Yu Y, Choi Y D, Park B S, Deynze A, Ashrafi H, Hill T, Kim W T, Pai H S, Ahn H K, Yeam I, Giovannoni J J, Rose J K, Sørensen I, Lee S J, Kim R W, Choi I Y, Choi B S, Lim J S, Lee Y H, Choi D. 2014. Genome sequence of the hot pepper provides insights into the evolution of pungency in Capsicum species. Nature Genetics, 46:270-278. doi: 10.1038/ng.2877 URL
[15]	Lentz D L, Beaudry-Corbett M P. 1996. Foodstuffs,forests,fields,and shelter:a paleoethnobotanical analysis of vessel contents from the ceren site,El salvador. Latin American Antiquity, 7 (3):247-262. doi: 10.2307/971577 URL
[16]	Linne C. 1753. Species Plantarum. Holmiae:Impensis Laurentii Salvii.
[17]	Liu Su. 2018. Pepper:the origin is unkown. China Science & Technology Education,(3):68-69. (in Chinese)
	刘夙. 2018. 辣椒:你不了解我的起源. 中国科技教育,(3):68-69.
[18]	Liu Wen-ming, An Zhi-xing, Jing Li-jun, Huang Guo-qing. 2005. Species within,origins of the capsicum species and its spread to the world. Journal of China Capsicum,(4):17-18. (in Chinese)
	刘文明, 安志信, 井立军, 黄国青. 2005. 辣椒的种类、起源和传播. 辣椒杂志,(4):17-18.
[19]	Martínez O, Arce-Rodríguez M L, Hernández-Godínez F, Escoto-Sandoval C, Cervantes-Hernández F, Hayano-Kanashiro C. 2021. Transcriptome analyses throughout chili pepper fruit development reveal novel insights into domestication process. Plants,(10):585.
[20]	McLeod M J, Eshbaugh W H, Guttman S I. 1979. A preliminary biochemical systematic study of the genus Capsicum Solanaceae// Hawkes J G,Lester R N,Skelding A D. The biology and taxonomy of the Solanaceae. New York: Academic Press:701-713.
[21]	McLeod M J, Guttman S I, Eshbaugh W H. 1982. Early evolution of chili peppers(Capsicum). Economic Botany, 36 (4):361-368. doi: 10.1007/BF02862689 URL
[22]	McLeod M J, Guttman S I, Eshbaugh W H, Rayle R E. 1983. An electrophoretic study of evolution in Capsicum(Solanaceae). Evolution, 37 (3):562-574. doi: 10.1111/j.1558-5646.1983.tb05573.x URL
[23]	Nee M, Bohs L, Knapp S. 2006. New species of Solanum and Capsicum(Solanaceae)from Bolivia with clarification of nomenclature in some Bolivian Solanum. Brittonia, 58:322-356. doi: 10.1663/0007-196X(2006)58[322:NSOSAC]2.0.CO;2 URL
[24]	Ou L J, Zhang Z Q, Dai X Z, Zou X X. 2012. Seed micromorphology and germination characteristics of wild and cultivated pepper strains. Plant Systematics and Evolution, 298 (6):1211-1216. doi: 10.1007/s00606-012-0622-8 URL
[25]	Ou L J, Zou X X. 2012. Inter simple sequence repeat analysis of genetic diversity of five cultivated pepper species. African Journal of Biotechnology, 11 (4):752-757. doi: 10.5897/AJB10.2551 URL
[26]	Pearsall D M. 2003. Plant food resources of the ecuadorian formative: an overview and comparison to the central andes// Raymond J S,Burger R L. Archaeology of formative ecuador. a symposium at Dumbarton Oaks,7 and 8 October 1995. Washington D C:Dumbarton Oaks Research Library and Collection:213-257.
[27]	Pearsall D M, Chandler-Ezell K, Zeidler J A. 2004. Maize in ancient Ecuador:results of residue analysis of stone tools from the Real Alto site. Journal of Archaeological Science, 31 (4):423-442. doi: 10.1016/j.jas.2003.09.010 URL
[28]	Perry L, Dickau R, Zarrillo S, Holst I, Pearsall D M, Piperno D R, Berman M J, Cooke R G, Rademaker K, Ranere A J, Raymond J S, Sandweiss D H, Scaramelli F, Tarble Kay, Zeidler J A. 2007. Starch fossils and the domestication and dispersal of chili peppers(Capsicum spp L.)in the Americas. Science, 315 (5814):986-988. doi: 10.1126/science.1136914 URL
[29]	Pickersgill B. 1969. The archaeological record of chili peppers(Capsicum spp.)and the sequence of plant domestication in peru. American Antiquity, 34 (1):54-61. doi: 10.2307/278313 URL
[30]	Pickersgill B. 1981. Biosystematics of crop-weed complexes. Die Kulturpflanze, 29:377-388. doi: 10.1007/BF02014766 URL
[31]	Pickersgill B. 1984. Migrations of chilli peppers,Capsicum spp,in the Americas// Stone D. Pre-columbian plant migration. Papers of the peabody museum of archaeology and ethnology. Cambridge: Harvard University Press:105-123.
[32]	Pickersgill B. 1988. The genus Capsicum: a multidisciplinary approach to the taxonomy of cultivated and wild plants. Biologisches Zentralblatt, 107 (4):381-389.
[33]	Piperno D R, Pearsall D M. 1998. The origins of agriculture in the lowland Neotropics. Academic Press: San Diego, CA,USA,400.
[34]	Piperno D R, Ranere A J, Holst I, Hansell P. 2000. Starch grains reveal early root crop horticulture in the Panamanian tropical forest. Nature, 407 (6806):894-897. doi: 10.1038/35038055 URL
[35]	Pozzobon M T, Schifino-wittman M T, Bianchetti L de bem. 2006. Chromosome numbers in wild and semi-domesticated Brazilian Capsicum L.(Solanaceae)species:do x = 12 and x = 13 represent two evolutionary lines? Botanical Journal of the Linnaean Society, 151:259-269. doi: 10.1111/j.1095-8339.2006.00503.x URL
[36]	Qin C, Yu C S, Shen Y O, Fang X D, Chen L, Min J M, Cheng J W, Zhao S C, Xu M, Luo Y, Yang Y L, Wu Z M, Mao L K, Wu H Y, Ling-Hu C Y, Zhou H K, Lin H J, González-Morales S, Trejo-Saavedra D L, Tian H, Tang X, Zhao M J, Huang Z Y, Zhou A W, Yao X M, Cui J J, Li W Q, Chen Z, Feng Y Q, Niu Y C, Bi S M, Yang X W, Li W P, Cai H M, Luo X R, Montes-Hernández S, Leyva-González M A, Xiong Z Q, He X J, Bai L J, Tan S, Tang X Q, Liu D, Liu J W, Zhang S X, Chen M S, Zhang L, Zhang L, Zhang Y C, Liao W Q, Zhang Y, Wang M, Lv X D, Wen B, Liu H J, Luan H M, Zhang Y G, Yang S, Wang X D, Xu J H, Li X Q, Li S C, Wang J Y, Palloix A, Bosland P W, Li Y R, Krogh A, Rivera-Bustamante R F, Herrera-Estrella L, Yin Y, Yu J P, Hu K L, Zhang Z M. 2014. Whole-genome sequencing of cultivated and wild peppers provides insights into Capsicum domestication and specialization. Proceedings of the National Academy of Sciences of the United States of America, 111 (14):5135-5140.
[37]	Smith C E. 1967. Plant remains// Byes D S. The Prehistory of the Tehuacan Valley,Vol. 1. Environment and Subsistence. Andover MA:RS Peabody Foundation:220-255.
[38]	Smith P G, Heiser C B. 1957a. Taxonomy of Capsicum sinense Jacq. and the geographic distribution of the cultivated Capsicum species. Bulletin of the Torrey Botanical Club, 84 (6):413-420. doi: 10.2307/2482971 URL
[39]	Smith P G, Heiser C B. 1957b. Breeding behavior of the cultivated peppers. Journal of the American Society for Horticultural Science, 70:286-290.
[40]	Stebbins G L, Major J. 1965. Endemism and speciation in the California flora. Ecological Monographs, 35:1-35. doi: 10.2307/1942216 URL
[41]	Stebbins G L. 1972. Ecological distribution of centers of major adaptive radiation in angiosperms// ValentineD H. Taxonomy,Phytogeography,and Evolution. New York Academic Press:7-34.
[42]	Stuart W. 2020. The devil's dinner:A gastronomic and cultural history of chili peppers. Beijing: Social Sciences Aca Demic Press:11-14. (in Chinese)
	斯图尔特沃尔顿. 2020. 艾栗斯,译. 魔鬼的晚餐:改变世界的辣椒和辣椒文化. 北京: 社会科学文献出版社:11-14.
[43]	Tang B Q, Xie L L, Li X F, Yang H P, Liu Y, Cheng X, Liang C L, Chen J, Zhou S D, Wang D H, Zheng J Y, Dai X Z, Zou X X, Liu F. 2021. Novel structural annotation and functional expression analysis of GTP_EFTU conserved genes in pepper based on the PacBio sequencing data. Horticultural Plant Journal, 7 (5):443-456. doi: 10.1016/j.hpj.2020.09.007 URL
[44]	Unzueta O Q. 1975. Mapa ecológica de Bolivia:memoria explicativa. Ministerio de Asuntos Campesinos y Agropecuarios,División de Suelos,Riegos e Ingeniera,La Paz,Bolivia. 309.
[45]	Yoshimoto Yamamoto. 2018. World history of pepper:a journey across Europe,Asia,and Africa,a hot table revolution Chen Xian-ruo trans trans. Taiwan, China: Marco Polo Press:51-75,203-243. (in Chinese)
	山本纪夫. 2018. 辣椒的世界史:橫跨欧亚非的寻味旅程, 一场热辣过瘾的餐桌革命. 陈娴若,译. 中国台湾: 马可孛罗出版社:51-75,203-243.
[46]	Zou Xue-xiao. 2002. Chinese pepper. Beijing: China Agriculture Press:1-21. (in Chinese)
	邹学校. 2002. 中国辣椒. 北京: 中国农业出版社:1-21.
[47]	Zou Xue-xiao. 2009. Pepper genetics and breeding. Beijing: Science Press:9-32. (in Chinese)
	邹学校. 2009. 辣椒遗传育种学. 北京: 科学出版社:9-32.
[48]	Zou Xue-xiao. 2017. Pepper breeding// Fang Zhi-yuan. Vegetable breeding in China. Beijing: China Agriculture Press:1113-1157. (in Chinese)
	邹学校. 2017. 辣椒育种// 方智远. 中国蔬菜育种学. 北京: 中国农业出版社:1113-1157.
[49]	Zou Xuexiao, Ma Yanqing, Dai Xiongze, Li Xuefeng, Yang Sha. 2020. Spread and industry development of pepper in China. Acta Horticulturae Simica, 47 (9):1715-1726. (in Chinese)
	邹学校, 马艳青, 戴雄泽, 李雪峰, 杨莎. 2020. 辣椒在中国的传播与产业发展. 园艺学报, 47 (9):1715-1726.
[50]	Zou Xue-xiao, Zhu Fan. 2020. The path of pepper introduction into China and its spreading route in China. Journal of Hunan Agricultural University(Natural Sciences), 46 (6):629-640. (in Chinese)
	邹学校, 朱凡. 2020. 辣椒传入中国的途径与传播路径. 湖南农业大学学报(自然科学版), 46 (6):629-640.

辣椒属物种 Capsicum	美洲分布 South America distribution	染色体数2n Chromosome number 2n
C. annuum L.	哥伦比亚北部到美国南部 Colombia north to southern USA	24,48
C. baccatum L.	阿根廷、玻利维亚、巴西、巴拉圭、秘鲁Argentina,Bolivia,Brazil,Paraguay,Peru	24
C. buforum Hunz.	巴西(e) Brazil (e)	26
C. caballeroi M. Nee	玻利维亚(e) Bolivia (e)
C. campylopodium Sendtn.	巴西南部、巴拉圭(e) Southern Brazil,Paraguay (e)	26
C. cardenasii Heiser & Smith	玻利维亚(e) Bolivia (e)	24
C. ceratocaylx M. Nee	玻利维亚(e) Bolivia (e)
C. chacoense Hunz.	阿根廷、玻利维亚Argentina,Bolivia	24
C. chinense Jacq.	美国南部、亚马逊、加勒比海South America,Amazonas,Caribbean	24
C. coccineum (Rusby) Hunz.	玻利维亚、秘鲁Bolivia,Peru
C. comutum (Hiem) Hunz.	巴西南部(e) Southern Brazil (e)	26
C. dimorphum (Miers) Kuntze	哥伦比亚Colombia
C. usenii Bitter	巴西东南部(e) Southeastern Brazil (e)
C. eshbaughii Barboza	玻利维亚(e) Bolivia (e)	24
C. eximium Hunz.	阿根廷、玻利维亚Argentina,Bolivia	24
C. flexuosum Sendtn.	巴西、巴拉圭Brazil,Paraguay	24
C. friburgense Bianchetti & Barboza	巴西南部(e) Southern Brazil (e)	26
C. frutescens L.	美国南部、亚马逊、加勒比海 South America,Amazonas,Caribbean	24
C. galapagoenese Hunz.	厄瓜多尔(e) Ecuador (e)	24
C. geminifolium (Dammer) Hunz.	哥伦比亚、厄瓜多尔Colombia,Ecuador
C. hookenanum (Miers) Kuntze	厄瓜多尔(e) Ecuador (e)
C. hunzikerianum Barboza & Bianchetti	巴西南部(e) Southern Brazil (e)
C. lanceolatum (Greenm.) Morton & Standi.	墨西哥、危地马拉、洪都拉斯Mexico,Guatemala,Honduras	26
C. leptopodum (Dunal) Kuntze	巴西(e) Brazil (e)
C. minuntifJorum (Rusby) Hunz.	阿根廷、玻利维亚、巴拉圭Argentina,Bolivia,Paraguay
C. mirabile Mart.	巴西南部 (e) Southern Brazil (e)	26
C. parvifolium Sendtn.	哥伦比亚、巴西北部,委内瑞拉Colombia,northern Brazil,Venezuela	24
C. pereirae Barboza & Bianchetti	巴西南部(e) Southern Brazil (e)	26
C. praetermissum Heiser & Smith	巴西南部(e) Southern Brazil (e)	24
C. pubescens Ruiz & Pav.	安第斯、美国南部、危地马拉、墨西哥Andean,South America,Guatemala,Mexico	24
C. recurvatum Witasek	巴西南部(e) Southern Brazil (e)
C. rhomboideum (Humb.,Bonp., & Kuntze) Kuntze	墨西哥南部到秘鲁南部和委内瑞拉Southern Mexico to s Peru and Venezuela	26
C. chottianum Sendtn.	阿根廷、巴西南部、巴拉圭东南部Argentina,southern Brazil,se Paraguay	26
C. scclnikianum Hunz.	秘鲁(e) Peru (e)
C. tovarii Eshbaugh,Smith & NicKrent	秘鲁(e) Peru (e)	24
C. villosum Sendtn.	巴西南部(e) Southern Brazil (e)	26

辣椒属物种 Capsicum	美洲分布 South America distribution	染色体数2n Chromosome number 2n
C. annuum L.	哥伦比亚北部到美国南部 Colombia north to southern USA	24,48
C. baccatum L.	阿根廷、玻利维亚、巴西、巴拉圭、秘鲁Argentina,Bolivia,Brazil,Paraguay,Peru	24
C. buforum Hunz.	巴西(e) Brazil (e)	26
C. caballeroi M. Nee	玻利维亚(e) Bolivia (e)
C. campylopodium Sendtn.	巴西南部、巴拉圭(e) Southern Brazil,Paraguay (e)	26
C. cardenasii Heiser & Smith	玻利维亚(e) Bolivia (e)	24
C. ceratocaylx M. Nee	玻利维亚(e) Bolivia (e)
C. chacoense Hunz.	阿根廷、玻利维亚Argentina,Bolivia	24
C. chinense Jacq.	美国南部、亚马逊、加勒比海South America,Amazonas,Caribbean	24
C. coccineum (Rusby) Hunz.	玻利维亚、秘鲁Bolivia,Peru
C. comutum (Hiem) Hunz.	巴西南部(e) Southern Brazil (e)	26
C. dimorphum (Miers) Kuntze	哥伦比亚Colombia
C. usenii Bitter	巴西东南部(e) Southeastern Brazil (e)
C. eshbaughii Barboza	玻利维亚(e) Bolivia (e)	24
C. eximium Hunz.	阿根廷、玻利维亚Argentina,Bolivia	24
C. flexuosum Sendtn.	巴西、巴拉圭Brazil,Paraguay	24
C. friburgense Bianchetti & Barboza	巴西南部(e) Southern Brazil (e)	26
C. frutescens L.	美国南部、亚马逊、加勒比海 South America,Amazonas,Caribbean	24
C. galapagoenese Hunz.	厄瓜多尔(e) Ecuador (e)	24
C. geminifolium (Dammer) Hunz.	哥伦比亚、厄瓜多尔Colombia,Ecuador
C. hookenanum (Miers) Kuntze	厄瓜多尔(e) Ecuador (e)
C. hunzikerianum Barboza & Bianchetti	巴西南部(e) Southern Brazil (e)
C. lanceolatum (Greenm.) Morton & Standi.	墨西哥、危地马拉、洪都拉斯Mexico,Guatemala,Honduras	26
C. leptopodum (Dunal) Kuntze	巴西(e) Brazil (e)
C. minuntifJorum (Rusby) Hunz.	阿根廷、玻利维亚、巴拉圭Argentina,Bolivia,Paraguay
C. mirabile Mart.	巴西南部 (e) Southern Brazil (e)	26
C. parvifolium Sendtn.	哥伦比亚、巴西北部,委内瑞拉Colombia,northern Brazil,Venezuela	24
C. pereirae Barboza & Bianchetti	巴西南部(e) Southern Brazil (e)	26
C. praetermissum Heiser & Smith	巴西南部(e) Southern Brazil (e)	24
C. pubescens Ruiz & Pav.	安第斯、美国南部、危地马拉、墨西哥Andean,South America,Guatemala,Mexico	24
C. recurvatum Witasek	巴西南部(e) Southern Brazil (e)
C. rhomboideum (Humb.,Bonp., & Kuntze) Kuntze	墨西哥南部到秘鲁南部和委内瑞拉Southern Mexico to s Peru and Venezuela	26
C. chottianum Sendtn.	阿根廷、巴西南部、巴拉圭东南部Argentina,southern Brazil,se Paraguay	26
C. scclnikianum Hunz.	秘鲁(e) Peru (e)
C. tovarii Eshbaugh,Smith & NicKrent	秘鲁(e) Peru (e)	24
C. villosum Sendtn.	巴西南部(e) Southern Brazil (e)	26