洪欣 KEENEJeremy 單皖粵 DOVanTruong 溫放

摘 要：? ?原唇柱苣苔屬（ Chirita? Buch.-Ham. ex D. Don）為一個人為界定的屬， 2011年在分子系統(tǒng)學(xué)研究的基礎(chǔ)上對該屬及其近緣屬開展了系統(tǒng)發(fā)育重建工作，其中絕大部分的原唇柱苣苔屬唇柱苣苔組（Sect.? Gibbosaccus? C. B. Clark）的物種被并入了廣義報春苣苔屬（ Primulina? Hance）。然而，由于歷史原因和早期經(jīng)典分類學(xué)在研究方法上的局限性以及對現(xiàn)報春苣苔屬部分物種的營養(yǎng)器官與生殖器官的認知不夠，該屬下一些物種的分類仍存在一些問題，亟待深入研究。比如，在對中國和越南分布的苦苣苔科植物開展研究的過程中，作者發(fā)現(xiàn)兩個報春苣苔屬的物種——廣布于中國西南和華南直至中南半島中部的鐘冠報春苣苔[ Primulina swinglei? （Merr.） Mich. MOller & A. Weber]命名人和原被認為是中國與廣西特有種的疏花報春苣苔[ P. laxiflora （W. T. Wang） Yin. Z. Wang]之間的鑒定存在分類學(xué)問題，需要進一步厘清兩者之間的關(guān)系。該文對這兩個物種進行了形態(tài)比較，同時通過對這兩種植物的原始描述對比、植物標本檢查、栽培觀察以及野外實地觀察，確定疏花報春苣苔是鐘冠報春苣苔的異名。此外，還明確了鐘冠報春苣苔的后選指定模式標本。

關(guān)鍵詞：? 唇柱苣苔屬， 后選模式指定， 新異名， 疏花報春苣苔， 分類學(xué)

1 Introduction

Primulina? Hance （1883： 169） is a Sino-Vietnamese genus of the family Gesneriaceae consisting of at least? 200 species of perennial rosette herbs （MOller， 2019; Wen et al.， 2019）. The species grows mainly in wet， shaded， mountainous regions of South and Southwest China （especially Guangxi， Southeast Yunnan， Guizhou， North Guangdong and South Hunan） and North Vietnam （Wei et al.， 2010; Weber et al.， 2011; Wang et al.， 2011; MOller et al.， 2016） respectively revised the former genus? Chirita? Buch.-Ham.ex D. Don （1822： 83） and allied genera by utilizing molecular data. These studies showed that? Chirita? was a polyphyletic group and needed to be reorganized. This work then led to the synonymization of? Chirita? within an enlarged? Primulina? genus， due to naming priority. Before this reorganization， numerous new species of former? Chirita? were described by Wang （1984a， b， 1985， 1990） and Wang et al. （1998）. Due to the limitation of research methods， field observations， and insufficient knowledge of flowers and fruits for certain species of former? Chirita ， the taxonomy of some species continues to be problematic.

During a review of Gesneriaceae from the Sino-Vietnam area， we found that specimens labeled as? Primulina swinglei? （Merrill 1918： 156） Mich. MOller & A. Weber in Weber et al. （2011： 785） and? P. laxiflora? （W. T. Wang 1985： 21） Yin Z. Wang in Wang et al. （2011： 61） are widely confused with each other. Our field investigations and examinations of type materials demonstrated that? P. laxiflora? is conspecific with? P. swinglei ， a species widely distributed in Southwest to South China to North Vietnam. We proposed that? P. laxiflora? should be reduced to a synonym of? P. swinglei . The lectotype for? P. swinglei? is also designated here under the International Code of Botanical Nomenclature （Shenzhen Code） （Turland & Wiersema 2017; Turland et al.， 2018）. During our research on type specimens of? P. swinglei ， we found that two or more collections or two specimens were simultaneously designated as types， without indicating the holotype in the protologue. The lectotype for? P. swinglei? are also named here since this taxon was published based on syntypes， without mentioning the holotype.

2 Materials and Methods

2.1 Field investigation and specimen examination

We have conducted fieldwork in Guangxi and Guangdong of China （including the type localities of? Primulina laxiflora? and? P. swinglei ） and North Vietnam since 2012. During the floristic field surveys， we observed living plants and collected specimens. Numerous new populations of? P. swinglei? were discovered during these field trips. Plants from different populations have been regularly monitored in the field since their discovery. Furthermore， some plants were collected to carry out cultivation experiments （Groot et al.， 2018） in the nursery of the Gesneriad Conservation Center of China （GCCC） in Guangxi Institute of Botany by the authors over the past six years. We also checked specimens （including type specimens） from the following herbaria： A， E， GH， HN， IBK， IBSC， K， MO， PE， UC and VMN （abbreviations follow Thiers 2015）. All morphological characters were studied using a dissecting microscope （SZX16， Olympus， Tokyo， Japan）. Additionally， photos of the whole plants and flowers were taken with a digital camera （X-H1， Fuji， Sendai， Japan）. Characters were described using the terminology presented by Wang et al. （1998）.

2.2 Specimens examined

Primulina swinglei? ：? CHINA.? ?Guangdong Province：? ?Huizhou City， Boluo County， Luofu Mountain， 16 August 1917，? C. O. Levine 1538? （IBSC; GH）; Boluo County， Luofu Mountain， 28 July 1930，? N. Q. Chen 41431? （IBSC）; Boluo County， Luofu Mountain， 13 Junuary 1935，? L. Deng 331? （IBSC）; Ruyuan County， Wuyang Town， 18 October 1936，? Y. Li 2014? （IBSC）; no specific locality， no exact collection time，? X. R. Liang 69669? （IBK）.? ?Guangxi Zhuang Autonomous Region? ： Qinzhou City， 8 July 1912，? K. K. Tsoong 1980? （IBSC）; Fangchenggang City， Naliang Town， 7 July 1917，? K. K. Chung s. n.? （IBSC）; no specific locality， 15 May 1924，? Kwangsi Mus. 254? （IBSC）; Fangchenggang City， Shiwandashan Mt.， 14 July 1937，? X. R. Liang 69669? （IBSC）; Wuming County， Matou Town， 1 September 1958，? Y.C. Chen 785? （IBK）; Nanning， Medicine Hort. Garden， 28 May 1964，? X. C. Huang 00085? （GXMI）; Wuming County， Matou Town， 25 June 1965，? X. C. Huang， H. R. Zhu， C. Y. Xie 3814? （GXMI， PE）; Nanning City， Medicine Hort. Garden， 21 April 1975，? G. Y. Yang 7007? （GXMG）; Lingshan County， Taiping Town， 11 July 1977，? Lingshanzu 1-4041? （GXMI）; no specific locality， 25 June 1978，? Chengjunzu 00624? （GXMI）; Shanglin County， Dafeng Town， 2 July 1978，? Shanglindui 2-725? （GXMI）; Nanning City， Medicine Hort. Garden， 4 July 1979，? Z. Y. Ni， M. L. Chen 8028? （GXMG）; Nanning City， Medicine Hort. Garden， 1 September 1981，? X. H. Lu 9032? （GXMG）; Nanning City， Medicine Hort. Garden， 22 October 1981，? Z. Y. Ni 9035? （GXMG）; Fangchenggang City， Fulong Town， 9 July 2010，? Shiwandashancaijidui 2672? （IBK）; Fangchenggang City， Fulong Town， 10 July 2010，? ?Shiwandashancaijidui 2723? （IBK）; Fangchenggang City， Fulong Town， 11 July 2010，? Shiwandashancaijidui 2791? （IBK）; Fangchenggang City， Dongzhong Town， 19 July 2010，? Shiwandashancaijidui 3191? （IBK）;? Wuming County， Daming Mountain， 9 August 2010，? L. Wu， R. H. Jiang， et al. D0839? （IBK）; Wuming County， Daming Mountain， 10 August 2010，? L. Wu， R. H. Jiang， et al. D0919? （IBK）; Wuming County， Daming Mountain， 7 July 2011，? L. Wu， S. L. Jin D2566? （IBK）; Fangchenggang City， Nasuo Town， 1 August 2012，? W. B. Xu， Y. S. Huang 11713? （IBK）; Rong County， 24 March 2015，? W. B. Xu 12124? （IBK）.? ?VIETNAM.? ?Ton Kin， Mont. Bavi， près de Van-Maou， sur les roches moussues， 22 July 1886，? Balansa 4294? （P）; Ton Kin， forêts du Mont. Bavi， sur les bords ombragés et rocheux des torrents， July 1887，? Balansa 4287? （Kew， P）; Ton Kin， July 1887，? Balansa， Benedict 4287? （E）; Ton Kin， 1 September 1939，? W. T. Tsang 29473? （IBSC）; Ton Kin， Ha Coi， Tong Fa， Taai Wong Mo Shan， 11-23 September 1939，? 3 rd? Indo-China Expedition W. T. Tsang 09475， 29473? （E）; Ton Kin， de Sontay， Mont Bavi， April 1940，? 7.223? （E）; Ton Kin， Dam Ha， Sai Wong Mo Shan， July to September 1940，? 4 th? Indo-China Expedition W. T. Tsang 30186? （E）; Cao Bang， Tra Linh， Quoc Toan， 25 May 1997，? L. Averyanov， N. T. Hiep VH4903? （E）; Huong Son District， Ngam Thep， 23 May 1998，? N. T. Hiep， P. K. Loc， N. Q. Binh， L. Xiem 847? （E）; Quan Binh， Ninh Hoa， Hoa Tien， 30 April 2011，? L. A. Averyanov， P. K. Loc， N. Q. Hieu， P. V. The， N. T. Vinh CPC 2547? （E）.

Primulina? ? ? ?laxiflora? ?：? ?CHINA? .? ?Guangdong Province? ： Dapu County， Gucun Town， 13 June 1957，? L. Deng 5161? （IBSC）.? ?Guangxi Zhuang Autonomous Region? ： Heng County， Zhenlong Distrition， 8 May 1957，? Z. Z. Chen 50480? （IBSC）; Lingyun County， 17 June 1961，? S. W. Chen 19753? （IBK）; Longzhou County， Nonggang， Minqiang， 19 July 2009，? R. H. Jiang， W. H. Wu， D. X. Nong H09212? （IBK）; Longzhou County， Nonggang， Xiangshui， 29 July 2009，? Y. S. Huang， X. X. Ye， L. Wu H09835? （IBK）; Longzhou County， Nonggang， Liaowangtai， 3 March 2011，? Y. S. Huang y0067? （IBK）; Longzhou County， Nonggang， Xiangshui， 25 September 2011，? Y. Liu， R. C. Peng JRH2485? （IBK）; Fengshan County， Jiangzhou Town， 24 July 2014，? X. Y. Huang， Y. D. Peng， J. H. Li 451223140724022LY? （GXMG）; no specific locality， no exact collection time，? L. Q. Chen 90123? （IBSC）.? ?VIETNAM.? ?Xuan Truong， Bao Lac， no exact collection time，? V. T. Do， F. Wen， L. F. Fu VNM-CN 801? （VMNM）; Dak Rong Natural Reserve， Quang Tri Prov， no exact collection time，? V. T. Do， F. Wen， Y. G. Wei， Z. B. Xin VMN-CN 964? （VMNM）; Bavi National Park， no exact collection time，? V. T. Do， F. Wen， Y. G. Wei， Z. B. Xin VMN-CN 1138? （VMNM）; Ha Giang， Bac Me， Bac Me Nature Reserve， 16 June 2012，? V. T. Do， F. Wen VMN-CN229? （VMN， IBK）.

3 Results and Discussion

E.D. Merrill from the Bureau of Science of Manila spent his annual leave from 9 to 27 August， 1917 exploring the Luofu Mountain in Guangdong Province， China， for botanical field collection with C. O. Levine from the Canton Christian College. On this trip， Merrill made a collection aggregating 544 numbers. Levine also made an extensive collection in the same period and the same locality. In 1918，? Didymocarpus swinglei? Merr. （1918： 156） was first described by Merrill based on the type specimens： E. D. Merrill 10692 and the additional collections， C. O. Levine 1538 （Vitek et al.， 2000）. It was placed in the genus? Didymocarpus? Wallich （1819： 378） because of the unilobed linguiform shape of the stigma （Merrill， 1918）. The species is named after Walter T. Swingle from the United States Department of Agriculture， who made Merrills trip to China possible. However， in the protologue， Merrill did not indicate which of the two specimens he cited was the holotype.

When revising the family， Wang established the Series? Swinglei? W. T. Wang （1981： 62）， but he did not have the opportunity to check the type specimens. Wang referenced the specimens C. O. Levine 1538 （ [IBSC， barcode no. 0549123， 0649578]， L. O. Levine in Wangs paper is a mistake）， which were collected at the same locality by Levine on August 16th， 1917， and indicated it as the paratype （Wang， 1981， 1985）. Thus， almost all duplicates of C. O. Levine 1538 stored in different herbaria were indicated as type specimens of? ?Primulina swinglei? （i.e.? IBSC0549123，? IBSC0649578，? and GH00015873），? and were respectively annotated as paratype， syntype and isoparatype on the sheets. In these articles， Wang （1981， 1985） also referenced specimen Chen 41431，mistakenly identified as topotype on the sheet? [IBSC， barcode no. 0004889， 0549115] which was collected at the same locality in? 28 July， 1930， and he also referenced Liang 69669? [IBK， barcode no. 00054451] collected in the Shiwandashan Mountain， Guangxi Zhuang Autonomous Region. The latter was identified as? Chirita pellegriniana? Burtt （1960： 98） by Wood （1974）， which is a synonym of? Primulina swinglei? （Wang， 1990; Wang et al.， 1998; Weber et al.， 2011）.

In Vietnam， Didymocarpus balansae? Pellegrin （1926： 415） was described based on B. Balansas collections from Mont Bavi （Tonkin） in 1886-1887 and was recognized as an endemic species （Pellegrin & Lecomte， 1930）. Both specimens （Balansa 4287 and Balansa 4294） were simultaneously designated as types （Pellegrin， 1926）. It was regarded as a synonym of? Chirita pellegriniana? by Burtt （Burtt， 1960）. Later， it was considered to be conspecific with? Primulina swinglei? （Wang， 1985; Wang et al.， 1998; Weber et al.， 2011）. Therefore， these types were designated as the syntypes of? P. swinglei .

Primulina laxiflora? was described on the basis of a single collection， S. W. Chen 19753? [PE00030667 （Fig. 1）]， and recognized as an endemic from Lingyun County， NW Guangxi， China （Wang et al.， 1998; Li & Wang， 2005; Wei et al.， 2010）. Since its publication， this taxon has been generally considered to be very similar to? P. swinglei ， but to differ by a smaller leaf blade， 6-9.6 cm long （ vs . 6- 6 cm long）， smaller corolla， ca. 1.7 cm long （ vs . 2.4-4.2 cm long）， tube nearly tubular （ vs . campanulate to funnelform） and abaxial coralla lip 0.6-1.5 cm long （ vs . 1-3 cm long） （Wang， 1985; Wang et al.， 1998）. While? P. swinglei? has a wide geographic distribution， predominantly from East Guangdong through Guangxi to North Vietnam， where it grows in a variety of different habitats （Karst and granite landscapes）， with most populations occurring in karst landscapes （Do et al.， 2013）. We notably observed different individuals from the type locality of? P. swinglei ， Luofu Mountain of Guangdong， China， and found that many small individuals can get to? bloom. After consulting the original and relevant literature? （Merrill， 1918; Pellegrin， 1926; Burtt，? 1960; Weber et al.， 2011）， conducting field observations

4 Taxonomic Treatment

Primulina swinglei? （Merr.） Mich. MOller & A. Weber? ?in Taxon 60： 785. 2011;? ?—? ?Didymocarpus swinglei? Merrill， in Philipp. J. Sci. 13： 156. 1918? —? ?Chirita swinglei? （Merr.） W. T. Wang， in Bull. Bot. Res.， Harbin 1（4）： 62. 1981;? —? ?Chirita pellegriniana? B. L. Burtt， in Notes Roy. Bot. Gard. Edinb. 23： 98. 1960;? ?—? ?Didymocarpus balansae? Pellegrin， in Bull. Soc. Bot. France 73： 415. 1926.

Lectotype （designated here）：? ?CHINA. Guangdong Province： Huizhou City， Boluo County， Luofu Mountain， 9 to 27 August 1917， E. D. Merrill 10692 （lectotype， NY Herb. No. 63230? [Fig. 3]; isolectotypes： IBSC Herb. No. 0649577; UC Herb. No. 301080）.

=? ?Primulina? ? ? ?laxiflora? ? （W. T. Wang） Yin Z. Wang? ?syn. nov.? ?Type：? CHINA. Guangxi Zhuang Autonomous Region ： Lingyun County， rocks in limestone hills， 17 June 1961， S. W. Chen 19753 （holotype， PE Herb. No. 00030667）.

For a full description of? Primulina swinglei? see Wang et al. （1998） and Li & Wang （2005）.

Distribution and habitat：? ?South China （Guangdong Province， Guangxi Zhuang Autonomous Region）， North Vietnam （Hà Giang Province， Cao Bng Province， Ninh Bình Province， Ba Vì mountain range）

Conservation status：? ?During our fieldwork， we found that farmlands and rubber plantations were expanding in these areas， which would result in deforestation， habitat loss， and fragmentation of this species. However， due to the fact that the populations found in Guangxi and Guangdong of China and from Vietnam are composed of numerous individuals， and populations regenerate well; we estimate that this? species will not suddenly become extinct. Furthermore， the individuals and populations of? Primulina swinglei? are abundant in the two countries. Thus， following the IUCN （2017） red list categories and criteria， the conservation status of this species is of Least Concern （LC）.

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（ 責(zé)任編輯 李 莉 ）