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The Seed Plant Flora of the Mount Jinggangshan Region, Southeastern China

  • Lei Wang,

    Affiliation College of Resource Environment and Tourism, Capital Normal University, Haidian District, Beijing, P. R. China

  • Wenbo Liao ,

    lsslwb@mail.sysu.edu.cn

    Affiliation State Key Laboratory of Biocontrol and Guangdong Key Laboratory of Plant Resources, School of Life Sciences, Sun Yat-sen University, Guangzhou, Guangdong, P. R. China

  • Chunquan Chen,

    Affiliation Jinggangshan Administration of Jiangxi Province, Jinggangshan, Jiangxi, P. R. China

  • Qiang Fan

    Affiliation State Key Laboratory of Biocontrol and Guangdong Key Laboratory of Plant Resources, School of Life Sciences, Sun Yat-sen University, Guangzhou, Guangdong, P. R. China

Abstract

The Mount Jinggangshan region is located between Jiangxi and Hunan provinces in southeastern China in the central section of the Luoxiao Mountains. A detailed investigation of Mount Jinggangshan region shows that the seed plant flora comprises 2,958 species in 1,003 genera and 210 families (Engler’s system adjusted according to Zhengyi Wu’s concept). Among them, 23 species of gymnospermae belong to 17 genera and 9 families, and 2,935 species of angiosperms are in 986 genera and 201 families. Moreover, they can also be sorted into woody plants (350 genera and 1,295 species) and herbaceous plants (653 genera and 1,663 species). The dominant families are mainly Fagaceae, Lauraceae, Theaceae, Hamamelidaceae, Magnoliaceae, Ericaceae, Styracaceae, Aquifoliaceae, Elaeocarpaceae, Aceraceae, Rosaceae, Corylaceae, Daphniphyllaceae, Symplocaceae, Euphorbiaceae, Pinaceae, Taxodiaceae, Cupressaceae and Taxaceae. Ancient and relic taxa include Ginkgo biloba, Fokienia hodginsii, Amentotaxus argotaenia, Disanthus cercidifolia subsp. longipes, Hamamelis mollis, Manglietia fordiana, Magnolia officinalis, Tsoongiodendron odorum, Fortunearia sinensis, Cyclocarya paliurus, Eucommia ulmoides, Sargentodoxa cuneata, Bretschneidera sinensis, Camptotheca acuminata, Tapiscia sinensis, etc. The flora of Mount Jinggangshan region includes 79 cosmopolitan genera and 924 non-cosmopolitan genera, which are 7.88% and 92.12% of all genera. The latter includes 452 tropical genera (48.92%) and 472 temperate genera (51.08%). The temperate elements include 44 genera endemic to China, accounting for 4.76% of all genera. Among 1,003 genera, 465 have only a single species and 401 are oligotypic genera (with 2-5 species). These genera account for 86.34% of all genera. The floristic analysis indicates that the flora of Mount Jinggangshan region is closely related to the flora of Mount Wuyishan region in southeastern China. The flora of Mount Jinggangshan region also contains many elements of central and southern China. Mount Jinggangshan region is an important north-south floristic passageway and is also a boundary between the floras of eastern, central and south China.

Introduction

The Mount Jinggangshan region is located in the center of the Luoxiao Mountains, an important, large, north-south trending mountain range in southeastern China. Compared with other larger mountain ranges, such as Mount Wuyishan, Nanling Mountains, Wuling Range, Mount Emeishan, Qinling Mountains, Hainan Mountains and Hengduan Mountains, the natural resources and biodiversity of the Luoxiao Mountains have been less well studied.

The Mount Jinggangshan region has four natural reserves: the Mount Jinggangshan National Nature Reserve, the Mount Qixiling National Nature Reserve and the Mount Nanfengmian National Nature Reserve, all of Jiangxi Province, and the Mount Taoyuandong National Natural Reserve of Hunan Province. During 1983-1984, the government of Jiangxi Province organized a comprehensive natural resource survey on the Mount Jinggangshan Nature Reserve [1]. Later, several institutions and scientists developed surveys and research projects on the biodiversity of the other three nature reserves [2-4]. Interestingly, the four reserves have the boundaries which are connected to each other, forming an integrated region. Based on a detailed research, it is stated that the flora of the Mount Jinggangshan region is the core of the Luoxiao Mountains’. Therefore, it is significant to comprehensively analyze the flora of the Mount Jinggangshan region.

Based on the field surveys, the specimens collected in the Mount Jinggangshan region from 2009 to 2012 and deposited in herbaria, and relevant literature, we compiled a checklist of the seed plant in the Mount Jinggangshan region [5]. Additionally, we analyzed the composition of the flora of the Mount Jinggangshan region in order to discuss its origin, evolution and floristic status.

Geography

Geographic Limits

The Mount Jinggangshan region, in the central section of the Luoxiao Mountains, has four, linked, nature reserves. It lies between 26° 13'04″-26°52'30″ N and 113° 56′30″-114°22'00″E and covers an area of 480.05 km2 (Figure 1). The Luoxiao Mountains is a large north-south trending mountain range that forms the boundary between Jiangxi and Hunan provinces in southeastern China.

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Figure 1. Map showing the locations of Mount Jinggangshan region and other twelve mounts in China.

Notes: JG: Mount Jinggangshan region; QY: Mount Qiyunshan; LS: Mount Lushan; NL: Nanling Mountains; WY: Mount Wuyishan region; WL: Wuling Range; SNJ: Mount Shennongxia; EM, Mount Emeishan; TW: Taiwan Mountains; HN: Hainan Mountains; TB: Mount Taibeishan; XSBN: Mount Xishuangbangna; GLG: Mount Gaoligongshan.

https://doi.org/10.1371/journal.pone.0075834.g001

Topography

The Mount Jinggangshan region is deeply cut by deep, V-shaped valleys, resulting in a steep topography. There are two main peaks, Nanfengmian (2,120.4 m above sea level) in Jiangxi Province and Lingfeng (2,122 m) in Hunan Province. The lowest point is 200 m above sea level. The Mount Jinggangshan region has more than 300 peaks above 1,000 m. The relative elevation ranges from 500 to 1,300 m.

Climate

The Mount Jinggangshan region is under the influence of the monsoon climate and characterized by four distinct seasons and abundant water and heat. Based on the data gathered by the Mount Jinggangshan Weather Bureau from 1971 to 2000, the annual mean temperature is 14.2°C. The hottest month is July, with a mean temperature of 23.9°C and an extreme high temperature of 36.7°C. The coldest month is January, with a mean temperature of 3.4°C and an extreme lowest temperature of −11.0°C. The annual mean sum of radiation ranges from 85 to 105 kcal/cm. The annual cumulative temperature (the sum of daily temperature mean >10°C) is 4,224°C. On average, there are 247.5 fog-free days per year. The annual mean precipitation is 1,889.8 mm with the greatest precipitation of 2,878.8 mm and the lowest precipitation of 1,297.4 mm. The annual mean evaporation capacity is 978.8 mm. The relative humidity is about 85%.

Soils

There are four major soil types in Mount Jinggangshan region: mountain red, yellow, dark yellow-brown and meadow soil. Among them, mountain yellow soil has the largest distribution area, occurring between 800 and 1,200 m. The second most extensive is mountain red soil, which is often at altitudes below 850 m. Mountain dark yellow-brown soil is mainly between 1,200 and 2,100 m. Mountain meadow soil is mainly between 1,600 m and 1,800 m [1].

Vegetation

Based on plot investigations and the Vegetation Classification System of China [6], the natural vegetation of Mount Jinggangshan region is divided into 12 types, 82 association groups and 167 associations [5]. Among them, the warm coniferous and broadleaved mixed forests, including Pinus massoniana mixed forests, Cunninghamia lanceolata mixed forests, Amentotaxus argotaenia mixed forests and Nageia nagi mixed forests, dominate with 28%-30% of the total natural vegetation. Typical evergreen broadleaved forests are also diverse. They include 15 association groups and 39 associations and account for 21.6% of the total natural vegetation, with the main dominant species including Schima superba, Schima argentea, Castanopsis sclerophylla, Castanopsis eyrei, Castanopsis lamontii, Cyclobalanopsis glauca, Cyclobalanopsis sessilifolia, Lithocarpus glaber, Cyclobalanopsis multinervis, Phoebe bournei, Phoebe hunanensis, Machilus leptophylla, Michelia maudiae, Michelia foveolata, Daphniphyllum glaucescens, Ternstroemia gymnanthera, Sloanea sinensis, etc.

The next largest type is the ravine or lowland monsoon evergreen broadleaved forests, including 12 association groups and 26 associations. The dominant species comprise Castanopsis fargesii, Castanopsis tibetana, Castanopsis fabri, Castanopsis carlesii, Cyclobalanopsis fleuryi, Altingia chinensis, Altingia gracilipes, Elaeocarpus japonicus, Michelia chapensis, Manglietia fordiana, Machilus thunbergii Exbucklandia tongkinensis, etc. Generally, these forests are evergreen broadleaved forests that include tree ferns, epiphyllous liverworts, vines and lianas, pteridophytes, epiphytic orchids, trees with buttress roots and strangler vines.

Geological History

Located in the central section of the Luoxiao Mountains, southeastern Eurasia and southeast of the junction zone of the Yangzi and Cathaysian paleo-plates, Mount Jinggangshan region has experienced a long geological history and undergone complicated evolutionary processes. The region was a basin in the South China Ocean during the Caledonian orogeny of the Neoproterozoic to Eopaleozoic, the margin of the sea during the late Paleozoic, Indosinian orogeny, geological tectonism of Yan mountain and the Himalayan mountain tectonic movement. Cambrian, Ordovician, Devonian and Quaternary strata occur within the Mount Jinggangshan region. The area has been above the sea for more than 20 million years. Through ecological succession, a diversity of life forms have evolved and the distinct vegetation of Mount Jinggangshan region has been formed [1].

Methods

Firstly, a checklist of the seed plant in Mount Jinggangshan region was prepared. We conducted field surveys and collected specimens in Mount Jinggangshan region from 2009 to 2012, with the permissions of four Nature Reserve Administration. No endangered or protected species were involved. Moreover, we consulted the specimens deposited in the herbarium of the Mount Jinggangshan Natural Reserve and the relevant literatures, such as Flora of China [7], Flora of Jiangxi [8,9], List of Jiangxi Seed Plant [10] and Species Checklist of Five Provinces in East China [11].

Secondly, we analyzed the composition and characteristics of the seed plant flora of Mount Jinggangshan region and compared our findings with information from other main mounts in China. Similarity coefficients were calculated according to shared families, genera and species using the formula S = 2c/(a+b) (S is the similarity coefficient between two regions, %; c equals the number of shared taxa; a equals the number of taxa within the region; and b equals the number of taxa within the region). The mounts selected for comparison included Mount Wuyishan region [12-14], Mount Qiyunshan [15], Mount Lushan [16], Wuling Range [17], Mount Emeishan [18], Mount Taibaishan [19], Nanling Mountains [20], Mount Shennongjia [21], Taiwan Mountains [22], Hainan Mountains [23], Mount Xishuangbanna [24] and Mount Gaoligongshan [25].

The Mount Wuyishan region in this paper includes two natural reserves: the Mount Wuyishan Natural Reserve of Fujian Province and the Mount Wuyishan Natural Reserve of Jiangxi Province. The former is situated between 27° 32'36″-27°55'15″ N and 117° 24'12″-118°02'50″ E and covers an area of 565.27 km2 [12]. It was listed as a World Natural Heritage site in December, 1999. The latter is situated between 27° 48'11″-28° 00’35″N and 117° 39'30″-117°55'47″ E and covers an area of 160.07 km2 [14]. The two natural reserves adjoin each other and share the same main peak, Huanggangshan, therefore, they are regarded as one integrated region. The Mount Wuyishan region is located between 27° 32'36″-28° 00’35″N and 117° 24'12″-118°02'50″ E and covers an area of 725.34 km2. The statistics on the seed plants in Mount Jinggangshan region and Mount Wuyishan region were given in Table 1.

Regions/TaxaFamiliesGeneraSpeciesIncluding infraspecies*References
Mount Jinggangshan19695118001855 [1]
Mount Qixiling18265114271464 [3]
Mount Nanfengmian17867614921533 [2]
Mount Taoyuandong18772815661618 [4]
Total (Mount Jinggangshan region)210100329583187In this paper
Mount Wuyishan in Fujian Province18978418061882 [11]
Mount Wuyishan in Jiangxi Province19380519332025 [12]
Total (Mount Wuyishan region)20189123312484In this paper

Table 1. Statistics of the seed plants in Mount Jinggangshan region and Mount Wuyishan region.

Notes:The figures in this table may differ from published reports. We have edited the checklists of those mountains to make them uniform at the family, genus and species level based on Zhengyi Wu [26-30]. Infraspecific taxa are considered to be equal to species for statistical purposes.
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Results

Floristic Composition

Table 2 shows the composition of the seed plant flora of Mount Jinggangshan region. Among them, woody plants are from 350 genera and 1295 species, while herbaceous plants are from 653 genera and 1663 species. Table 3, Table 4, Table 5, Table 6 rank all 210 families in Mount Jinggangshan region based on numbers of genera and species, and contain some information of families, such as the numbers of genera and species in Mount Jinggangshan region [5], in China [31] and in the word [32,33], the numbers of endemic general and endemic species, and Areal-types. The families can be divided into 5 grades according to the numbers of species included (Table 7). The seed plant flora of Mount Jinggangshan region is mainly made up of oligotypic families (100 families) and mesotypic families (57), which account for 74.76% of all families, 55.63% of all genera and 62.43% of all species. Compared with other mounts in rich biodiversity, Mount Jinggangshan region has less single-species families (40), such as Mount Emeishan 47 families, Nanling Mountains 59, Mount Dinghushan 58, Mount Lushan 49, Taiwan Mountain 56, so it goes to show that the flora of Mount Jinggangshan region obtain obvious differentiation.

TaxaMount Jinggangshan region [6]Jiangxi Province [8-10]China [31]
familygenusspeciesfamilygenusspeciesfamilygenusspecies
Gymnosperms91723923351234230
Angiosperms201986293521913094085325316626046
Dicotyledoneae1707622399
Monocotyledoneae31224536
Total (Percent%)21010032958228 (92.1)1332 (75.3)4120 (71.8)337 (62.3)3200 (31.3)26276 (11.3)

Table 2. Composition of the seed plant flora of Mount Jinggangshan region.

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FamiliesNumber of genera1GE2Number of species3SE4A5FamiliesNumber of generaGENumber of speciesSEA
Poaceae99/244/6685199/1375/10025311Asclepiadaceae6/44/180123/220/2200122
Compositae73/239/16201188/2477/22750291Hamamelidaceae12/19/30222/75/95168
Rosaceae29/56/95147/1119/2830701Melastomataceae8/26/18822/154/5005172
Cyperaceae15/37/98141/673/435091Caesalpiniaceae7/17/16522/110/260062
Papilionaceae46/127/480131/1100/12000342Symplocaceae1/1/122/25/32042
Labiatae33/97/236288/758/7175371Cucurbitaceae10/31/11821/165/84562
Orchidaceae35/177/880184/1037/21950142Solanaceae8/19/10221/99/246022
Lauraceae9/25/5083/437/2500502Primulaceae3/13/2021/580/800148
Rubiaceae20/80/60071/602/1000192Smilacaceae2/2/221/67/31562
Theaceae10/15/22160/373/610432Gesneriaceae8/57/147120/413/3870172
Fagaceae6/6/758/324/670302Magnoliaceae6/11/15120/97/250119
Ranunculaceae10/40/6255/739/2525258Ulmaceae7/8/16119/47/230102
Scrophulariaceae20/61/6554/669/1700131Acanthaceae14/63/22918/175/350032
Polygonaceae6/16/4350/229/110068Hydrangeaceae7/11/1718/131/190118
Liliaceae23/50/175148//335/2000101Styracaceae6/10/1117/84/160112
Apiacae24/101/434247/522/3780138Tiliaceae5/10/5016/79/45073
Euphorbiaceae13/66/222147/366/5970112Crassulaceae3/12/3416/247/137041
Aquifoliaceae1/1/147/118/405272Polygalaceae2/4/1816/48/104561
Ericaceae5/14/12643/720/3995308Araceae6/28/10615/193/402552
Urticaceae11/25/5442/238/262572Amaranthaceae4/14/17415/38/205001
Caprifoliaceae4/12/1542/195/42038Menispermaceae8/19/7014/79/42052
Vitaceae7/8/1439/143/850212Apocynaceae6/38/41514/169/455562
Celastraceae5/13/69138/213/1300212Malvaceae6/17/24314/68/422532
Rutaceae12/28/161134/134/1815112Aristolochiaceae3/5/7114/101/350122
Verbenaceae7/18/3034/170/1100132Onagraceae3/15/2414/58/65008
Araliaceae11//25/43132/181/1450162Elaeagnaceae1/2/314/44/4598
Aceraceae1/2/331/152/200188Lardizabalaceae4/7/913/33/3668
Cruciferae15/102/321129/417/340018Thymelaeaceae3/11/4813/90/89161
Rhamnaceae6/16/5029/137/900171Meliosmaceae1/1/113/35/5073
Myrsinaceae5/6/4128/129/143564Gentianaceae3/15/8012/351/90038
Moraceae6/9/3827/161/110052Dioscoreaceae1/1/312/80/80532
Violaceae1/4/2026/125/80081Boraginaceae7/47/148311/186/274058
Caryophyllaceae13/28/8624/294/220021Loranthaceae6/8/6811/50/95052
Oleaceae7/13/2424/180/615182Campanulaceae5/15/8411/136/238048
Actinidiaceae2/2/3124/53/3551814Saxifragaceae4/13/2911/281/63058

Table 3. Ranking of families (with >10 species) in Mount Jinggangshan region based on numbers of genera and species [5,32,33].

Notes: 1 Number of genera in Mount Jinggangshan region/number in China /number in the world; 2 Number of genera endemic to China; 3 Number of species in Mount Jinggangshan region/number in China /number in the world; 4 Number of species endemic to China; 5 Areal-types of families of the seed plants in Mount Jinggangshan region.
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FamiliesNumber of genera1GE2Number of species3SE4A5FamiliesNumber of generaGENumber of speciesSEA
Convolvulaceae7/21/5710/115/160001Utriculariaceae1/2/37/19/32011
Lythraceae4/11/2510/48/55011Hydrocharitaceae5/9/186/22/11602
Anacardiaceae3/15/7010/51/98532Juglandaceae5/7/916/27/5028
Buxaceae3/3/410/22/7082Sterculiaceae4/20/686/80/110012
Cornaceae3/5/1210/51/10068Flacourtiaceae3/10/8016/21/50012
Corylaceae2/4/410/30/6958Chenopodiaceae2/49/1006/198/140001
Schisandraceae2/2/310/29/9279Geraniaceae2/2/76/68/80501
Balsaminaceae1/2/210/191/100082Pinaceae4/10/1116/87/21058
Hypericaceae1/4/1010/54/30021Podophyllaceae2/2/26/7/759
Commelinaceae6/13/409/53/65202Trilliaceae2/2/46/10/5019
Berberidaceae2/2/149/245/70199Eriocaulaceae1/1/106/45/116012
Begoniaceae1/1/29/90/140182Potamogetonaceae1/1/26/30/10201
Fumariaceae1/7/169/215/45018Taxaceae4/4/615/13/2328
Chloranthaceae2/3/48/16/7542Lemnaceae3/3/65/7/3001
Ebenaceae1/1/48/58/54842Meliaceae3/6/525/10/26112
Sabiaceae1/1/38/26/10077Staphyleaceae3/3/35/15/4523
Vacciniaceae1/2/228/87/40071Zingiberaceae3/20/495/145/118817
Naucleaceae5/8/117/28/14304Lobeliaceae2/3/205/24/60012
Alismataceae2/5/127/12/8101Valerianaceae2/4/175/40/31518
Elaeocarpaceae2/2/127/52/62512Alangiaceae1/1/15/8/1722
Juncaceae2/2/77/80/43001Clethraceae1/1/25/16/7613
Salicaceae2/2/557/228/101078Oxalidaceae1/2/65/13/77001
Pittosporaceae1/1/97/34/20024Piperaceae1/4/55/54/360032

Table 4. Ranking of families (with 5-10 species) in Mount Jinggangshan region based on numbers of genera and species.

Notes: 1 Number of genera in Mount Jinggangshan region/number in China /number in the world; 2 Number of genera endemic to China; 3 Number of species in Mount Jinggangshan region/number in China /number in the world; 4 Number of species endemic to China; 5 Areal-types of families of the seed plants in Mount Jinggangshan region.
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FamiliesNumber of genera1GE2Number of species3SE4A5FamiliesNumber of generaGENumber of speciesSEA
Cupressaceae4/8/294/31/14028Iteaceae1/1/23/12/1819
Papaveraceae3/11/4114/58/76028Myrtaceae1/10/1313/91/462522
Haloragaceae2/2/84/7/14501Pyrolaceae1/4/143/31/4018
Iridaceae2/5/674/44/180012Stilaginaceae1/1/13/17/17014
Loganiaceae2/5/134/15/42022Trapaceae1/1/13/5/30010
Annonaceae1/22/1294/122/222024Guttiferae2/5/272/210/45002
Aucubaceae1/1/14/11/11114Hypoxidaceae2/3/82/8/16002
Balanophoraceae1/2/74/18/5012Nyssaceae2/2/512/7/2219
Cephalotaxaceae1/1/14/7/28314Portulacaceae2/3/362/7/39501
Droseraceae1/2/34/7/11501Santalaceae2/7/442/32/93502
Illiciaceae1/1/14/26/5049Saururaceae2/3/42/4/609
Philadelphaceae1/1/14/15/7549Acoraceae1/1/12/4/408
Phytolaccaceae1/2/184/7/6512Amaryllidaceae1/5/592/23/80002
Plantaginaceae1/1/474/13/135001Callitrichaceae1/1/12/4/2511
Viscaceae1/1/24/11/7011Ceratophyllaceae1/1/12/5/701
Sapindaceae3/25/13513/51/158022Cleomaceae1/1/102/3/30002
Simarubaceae3/4/193/12/9512Dipsacaceae1/4/112/20/290010
Betulaceae2/2/63/37/14518Myricaceae1/1/32/4/5701
Mimosaceae2/11/563/65/280002Olacaceae1/4/142/8/10312
Monotropaceae2/4/123/5/2108Orobanchaceae1/10/992/40/206008
Pontederiaceae2/2/93/5/3302Parnassiaceae1/1/22/36/5108
Buddlejaceae1/1/73/29/10012Podocarpaceae1/4/172/20/12508
Calycanthaceae1/2/53/4/1129Proteaceae1/2/802/21/160012
Cuscutaceae1/1/13/10/17001Sambucaceae1/1/12/5/2018
Daphniphyllaceae1/1/13/12/2507Stachyuraceae1/1/12/8/10114
Ehretiaceae1/1/13/12/5002Stemonaceae1/1/42/8/2715
Helwingiaceae1/1/13/5/8014Typhaceae1/1/12/10/1101

Table 5. Ranking of families (with 2-4 species) in Mount Jinggangshan region based on numbers of genera and species.

Notes: 1 Number of genera in Mount Jinggangshan region/number in China /number in the world; 2 Number of genera endemic to China; 3 Number of species in Mount Jinggangshan region/number in China /number in the world; 4 Number of species endemic to China; 5 Areal-types of families of the seed plants in Mount Jinggangshan region.
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FamiliesNumber of genera1GE2Number of species3SE4A5FamiliesNumber of generaGENumber of speciesSEA
Alliaceae1/3/131/112/79501Nageiaceae1/1/11/5/507
Asparagaceae1/1/11/24/30005Najadaceae1/1/11/4/5001
Bignoniaceae1/14/1101/43/80004Nandinaceae1/1/11/1/1014
Bischofiaceae1/1/11/2/217Nelumbonaceae1/1/11/1/209
Bretschneideraceae1/1/111/1/1115Nymphaeaceae1/2/51/8/4801
Cabombaceae1/1/21/1/601Palmae1/19/1891/95/236102
Cannabaceae1/2/21/3/308Passifloraceae1/2/161/22/60512
Capparaceae1/4/161/30/48002Penthoraceae1/1/11/1/209
Ellisiophyllaceae1/1/11/1/107Periplocaceae1/1/11/4/1212
Erythroxylaceae1/1/41/3/24002Phrymataceae1/1/11/1/109
Eucommiaceae1/1/111/1/1115Pistaciaceae1/1/11/3/10112
Euryalaceae1/1/51/1/95014Samydaceae1/2/171/18/40002
Ginkgoaceae1/1/111/1/1115Sargentodoxaceae1/1/111/1/1115
Gnetaceae1/1/11/7/3002Sparganiaceae1/1/11/5/1408
Grossulariaceae1/1/11/45/15008Spigeliaceae1/2/21/6/802
Hippocastanaceae1/1/21/8/3218Taccaceae1/2/21/4/1202
Hydrocotylaceae1/1/11/2/202Tapisciaceae1/1/211/2/5115
Menyanthaceae1/2/51/4/5801Taxodiaceae1/5/3011/7/13009
Molluginaceae1/2/91/6/8702Zannichelliaceae1/1/61/1/2001
Musaceae1/3/31/7/3504Zygophyllaceae1/5/261/27/28501

Table 6. Ranking of families (with 1 species) in Mount Jinggangshan region based on numbers of genera and species.

Notes: 1 Number of genera in Mount Jinggangshan region/number in China /number in the world; 2 Number of genera endemic to China; 3 Number of species in Mount Jinggangshan region/number in China /number in the world; 4 Number of species endemic to China; 5 Areal-types of families of the seed plants in Mount Jinggangshan region.
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TaxaSingle-species families (1 species)Oligotypic families (2-10 species)Mesotypic families (11-50 species)Pluritypic families (51-100 species)Macrotypic families (>100 species)
Gymnosperms4 (4:4)15 (14:21)
Angiosperms36 (36:36)95 (173:462)57 (371:1361)8 (143:553)5 (262:806)
Total40 (40:40)100 (187:483)57 (371:1361)8 (143:553)5 (262:806)
Percent of total (%)19.05/3.99/1.35247.62/18.64/16.3327.14/36.99/46.013.81/14.26/18.702.38/26.12/27.25

Table 7. Statistics of the family size from the seed plant flora of Mount Jinggangshan region.

Notes: 1 Number of families (number of genera included in the families : number of species included in the families); 2 Percent of all 210 families (percent of all 1003 genera : percent of all 2958 species).
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The typical families in Mount Jinggangshan region are Fagaceae, Lauraceae, Theaceae, Hamamelidaceae, Magnoliaceae, Ericaceae, Styracaceae, Aquifoliaceae, Elaeocarpaceae, Aceraceae, Daphniphyllaceae, Hydrangeaceae, Rosaceae, Symplocaceae, Euphorbiaceae, Pinaceae, Taxodiaceae, Cupressaceae and Taxaceae. Other families with biogeographical implication are Bretschneideraceae, Tapisciaceae, Sargentodoxaceae, Eucommiaceae, Ginkgoaceae, Nyssaceae, Stachyuraceae, Helwingiaceae, Aucubaceae, Berberidaceae, Corylaceae, Cephalotaxaceae, Actinidiaceae and Schisandraceae, etc.

Floristic Geographic Elements

According to the concept of families proposed by Zhengyi Wu [28,29,30], the family areal-types of spermatophyte flora in Mount Jinggangshan region can be divided into 12 types (Table 8, Table 3, Table 4, Table 5, Table 6), which are further grouped into three categories: Cosmopolitan, Tropical and Temperate.

Areal-types of familiesNumber of families in JGR / in ChinaPercent of total non-cosmopolitan families in JGR (%)
1. Cosmopolitan42/50-—
2. Pantropic81/12048.21
3. Tropical Asian and tropical American disjunct5/112.98
4. Old World Tropic7/174.17
5. Tropical Asia to Tropical Australia2/101.19
6. Tropical Asia to tropical Africa0 / 70
7. Tropical Asia (Indo-Malaysia)6/203.57
8. North temperate38/4722.62
9. East Asian and North American disjunct14/178.33
10. Old World temperate2/61.19
11. Temperate Asia0 / 00
12. Mediterranean region, western to central Asia1/80.60
13. Central Asia0 / 10
14. East Asia7/184.17
15. Endemic to China5/82.98
Total210/337100

Table 8. Areal-types of families from the seed plant flora of Mount Jinggangshan region (JGR).

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The genera can be divided into 5 categories according to size (Table 9). Based on the generic distribution concept proposed by Zhengyi Wu [26,27], the 1,003 seed plant genera can be divided into 14 types and 17 sub-types (Table 10). Those types and sub-types can be further sorted into three groups: cosmopolitan, 79 genera (7.88% of total, including 456 species); tropical, 452 genera (48.92% of non-cosmopolitan genera, including 1168 species); and temperate, 472 genera (51.08% of non-cosmopolitan genera, including 1278 species). Temperate genera contain 44 genera endemics to China (4.76% of non-cosmopolitan genera, including 56 species). The larger cosmopolitan genera include Cyperus (including 48 species), Rubus (45), Carex (42), Polygonum (37), Viola (26), Lysimachia (19), Clematis (24), Polygala (14), Salvia (11), Rhamnus (10) and Ranunculus (9). Among no-cosmopolitan genera, 51 genera with more than 10 species include 971 species, accounting for 32.83% of all species. For example, Ilex has 47 species, Rhododendron 31, Acer 31, Symplocos 22, Euonymus 21, Camellia 20, Eurya 19, Litsea 17, Cinnamomum 14, Machilus 13, Neolitsea 11, Lithocarpus 15, Cyclobalanopsis 14, Castanopsis 13, Ficus 16, Callicarpa 15, Photinia 14, Meliosma 13 and Styrax 11. The genera including the dominant species in arborous layer, in shrub layer and in herbaceous layer in Mount Jinggangshan region are listed in Table 11.

TaxaSingle-species genera1 (1 species)Oligotypic genera (2-5 species)Mesotypic genera (6-10 species)Pluritypic genera (11-20 species)Macrotypic genera (>21 species)
Gymnosperms13 (13)25 (12)
Angiosperms452 (452)396 (1155)86 (640)37 (529)14 (442)
Total465 (465)401 (1167)86 (640)37 (529)14 (442)

Table 9. Statistics of the genus size from the seed plant flora of Mount Jinaggangshan region.

Notes: 1 Single-species genera include monotypic genus containing only one species in the world. There are 63 monotypic genera in the Mount Jinggangshan region; 2 Number of genera (number of species included in the genera).
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Areal-types of generaNumber of genera in JGR / in ChinaPercent of the total non-cosmopolitan genera in JGR (%)
1. Cosmopolitan79/107-—
2. Pantropic164/30417.75
2-1.Tropical Asia, Australasia (to New Zealand) and Central to South America (or Mexico) disjunct10/211.08
2-2.Tropical Asia, Africa and Central to South America disjunct8/320.87
3. Tropical Asian and tropical American disjunct17/781.84
4. Old World Tropical54/1505.84
4-1.Tropical Asia, Africa (or East Africa, Madagascar) and Australasia disjunct12/271.30
5. Tropical Asia to Tropical Australia42/1544.55
5-1.Chinese (southwest) Subtropical and New Zealand disjunct1/20.11
6. Tropical Asia to tropical Africa29/1453.14
6-2.Tropical Asia and East Africa or Madagascar disjunct3/80.32
7. Tropical Asia (Indo-Malaysia)80/4608.66
7-1.Java (or Sumatra), Himalaya to south and southwest China disjunct or dispersed12/311.30
7-2.Tropical India to south China (particularly southern Yunnan)6/530.65
7-3.Burma, Thailand to southwest China4/360.43
7-4.Vietnam (or Indochina) to south China (or southwest China)10/661.08
8. North temperate113/19312.23
8-4.North Temperate and south temperate disjunct31/783.35
8-5.Eurasia and South America temperate disjunct1/80.11
9. East Asia and North America disjunct75/1228.12
9-1.East Asia and Mexicao disjunct1/20.11
10.Old World temperate41/1194.44
10-1.Mediterranean, western Asia (or central Asia) and East Asia disjunct9/300.97
10-2.Mediterranean and Himalayan disjunct1/70.11
10-3.Eurasia and southern Africa (sometimes Australasia also) disjunct4/180.43
11. Temperate Asia14/641.52
12. Mediterranean region, western to central Asia3/1330.32
12-3.Mediterranen to temperate-tropical Asia, Australasia and South America disjunct2/70.22
13. Central Asia1/730.11
14. East Asia60/756.49
14-1.Sino-Himalayan (SH)21/1422.27
14-2.Sino-Japanese (SJ)51/1005.52
15. Endemic to China*44/2514.76
Total1003/3200100

Table 10. Areal-types of genera from the seed plant flora of Mount Jinggangshan region (JGR).

Notes: * The genera endemic to China include Ginkgo, Cunninghamia, Pseudotaxus, Parakmeria, Tsoogiodendron, Sargentodoxa, Saruma, Eomencon, Yinshania, Poliothyrsis, Tutcheria, Clematoclethra, Speranskia, Chimonanthus, Fortunearia, Semiliquidambar, Eucommia, Pteroceltis, Monimopetalum, Poncirus, Eurycorymbus, Bretschneidera, Tapiscia, Cyclocarya, Camptotheca, Metapanax, Tetrapanax, Changium, Dickinsia, Melliodendron, Biondia, Sheareria, Sinojohnstonia, Thyrocarpus, Schnabelia, Bostrychanthera, Hanceola, Speirantha, Changnienia, Indocalamus, Gelidocalamus, Oligostachyum, Omphalotrigonotis, Emmenopterys.
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generaarborous layershrub layerherbaceous layer
TropicalCyclobalanopsis, Lithocarpus, Ternstroemia, Cleyera, Schima, Camellia, Cinnamomum, Machilus, Phoebe, Neolitsea, Exbucklandia, Sycopsis, Distylium, Michelia, Manglietia, Styrax, Alniphyllum, Elaeocarpus, Symplocos, Ficus, Dendropanax, Sapium, Fokienia, AmentotaxusEurya, Maesa, Ardisia, Turpinia, Sageretia, Tarenna, Smilax, Clethra, Lasianthus, Psychotria, Bambusa, Helicteres, Mallotus, Embelia, Gardenia, Smithia, Melastoma, Myrsine, Adina, Toddalia, Lindera, KadsuraImpatiens, Osbeckia, Phyllagathis, Elatostema, Pellionia, Didymocarpus, Ophiorrhiza, Elephantopus, Sesbania, Eupatorium, Pollia, Lycianthes, Alpinia, Paraphlomis, Floscopa, Brachiaria, Cyrtococcum, Heteropogon, Ichnanthus, Miscanthus, Thysanolaena, Pennisetum, Sporobolus, Coelogyne, Bulbostylis, Neyraudia, Fimbristylis
TemperateQuercus, Fagus, Castanopsis, Castanea, Rhododendron, Lyonia, Cerasus, Sorbus, Carpinus, Dendrobenthamia, Cornus, Acer, Alnus, Osmanthus, Magnolia, Illicium, Halesia, Stewartia, Liquidambar, Pterocarya, Pinus, Abies, Taxus, TsugaViburnum, Elaeagnus, Rosa, Berberis, Fraxinus, Vaccinium, Sambucus, Hydrangea, Toxicodendron, Lyonia, Pieris, Actinidia, Bredia, Deutzia, Enkianthus, Holboellia, PleioblastusCimicifuga, Eomecon, Draba, Sedum, Saxifraga, Fagopyrum, Circaea, Bredia, Geum, Habenaria, Oenanthe, Valeriana, Artemisia, Inula, Kalimeris, Swertia, Trigonotis, Bothriospermum, Pedicularis, Hosta, Veronicastrum, Oreocharis, Prunella, Aspidistra, Liriope, Acorus, Arundinella, Aster, Deyeuxia, Spodiopogon, Triarrhena

Table 11. The genera including the dominant species in arborous layer, in shrub layer and in herbaceous layer of the vegetation of Mount Jinggangshan region.

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Comparison with Other Twelve Main Mounts in China

Table 12 shows similarity coefficients between Mount Jinggangshan region and other twelve main mounts in China. Among them, the flora of Mount Jinggangshan region is most similar to Mount Wuyishan region’s, as one would expect from their close proximity. The similarities at family, genus and species level are 93.4%, 85.1% and 71.6%, respectively. Next are Mount Qiyunshan (family 89.8% /genus 81.4% /species 67.5%) and Mount Lushan (92.5%/81.3%/63.3%). The above mountains are all in eastern China. The third most similar regions are the Nanling Mountains (south of Mount Jinggangshan region) (90.4%/77.6%/61.6%) and the Wuling Range (west of Mount Jinggangshan) (93.1%/81.0%/57.6%).

MountJGWYQYLSNLWLSNJEMTWHNTBXSBNGLG
Area (km2)48072511029256310000705154359903300056324184046
F1210201182192219213181202246244156236215
G210038917907579969647808691251124763312171025
S32958233121521732271330402155241133783369175937673798
WYF4192/93.4
G5809/85.4
S61814/68.6
QYF176/89.8170/88.8
G730/81.4668/74.5
S1724/67.51409/62.9
LSF186/92.5179/91.1164/87.7
G715/81.3645/78.3580/75.0
S1484/63.31244/61.21056/54.4
NLF194/90.4188/89.5179/89.3176/85.6
G776/77.6704/74.6720/80.6591/67.4
S1746/61.61427/56.61560/64.1993/44.7
WLF197/93.1197/95.2174/88.1184/90.9190/88.0
G797/81.0724/78.1649/74.0645/75.0713/72.8
S1726/57.61352/50.31241/47.81126/47.21337/46.5
SNJF168/85.9167/87.4151/83.2159/85.3165/82.5179/90.9
G618/69.3576/68.9497/63.3533/69.4537/60.5658/75.5
S1079/42.2873/38.9730/33.9821/42.2751/30.91346/51.8
EMF182/88.4180/89.3164/85.4172/87.3181/86.0159/76.6169/88.3
G677/72.3629/71.5580/69.9555/68.3627/67.2715/78.0602/73.0
S1073/40.0839/35.4802/35.2731/35.3869/33.91353/49.61024/44.9
TWF196/86.0186/83.2171/79.9178/81.3199/85.6192/83.7164/76.8182/81.3
G752/66.7677/63.2624/61.1586/58.4726/64.6689/62.2528/52.0632/59.6
S949/30.0781/27.4707/25.6633/24.8829/27.2714/22.2416/15.0535/18.5
HNF175/77.1167/75.1168/78.9159/72.9190/82.1140/61.3144/67.8161/72.2207/84.5
G600/53.3553/51.7547/53.7450/44.9664/59.2526/47.6358/35.3470/44.4799/64.0
S905/28.6747/26.2834/30.2482/18.91103/36.3605/18.9281/10.2444/15.4899/26.6
TBF142/77.6139/77.9125/74.0140/80.5136/72.5133/72.1144/85.5145/81.0138/68.7120/60.0
G468/57.2428/56.2362/50.9421/60.6374/45.9473/59.2524/74.2445/59.3399/42.4247/26.3
S653/27.7518/25.3414/21.2553/31.7401/17.9748/31.2951/48.6605/29.0251/9.8162/6.3
XSBNF183/82.1173/79.2171/81.8165/77.1196/81.3147/65.5150/71.9169/77.2203/84.2212/88.3128/65.3
G611/55.0548/52.0541/53.9443/44.9659/59.6569/52.2399/40.0526/50.4755/61.2863/70.0279/30.2
S804/23.9564/18.5674/22.8464/16.9882/27.2699/20.5340/11.5604/19.6646/18.11304/36.5189/6.8
GLGF185/87.1178/85.6168/84.6170/83.5186/85.7155/72.4167/84.3187/89.7193/83.7175/76.3143/77.1182/80.7
G649/64.0588/61.4556/61.3517/58.0638/63.1678/68.2566/62.7660/69.7671/59.0557/49.0433/52.2661/59.0
S741/21.9562/18.3554/18.6474/17.1656/20.2910/26.6659/22.1947/30.5510/14.2525/14.7456/16.41110/29.3

Table 12. Similarity coefficients at family, genus and species between Mount Jinggangshan region and other twelve main mounts in China.

Notes: 1 Number of families in the mount; 2 Number of genera in the mount; 3 Number of species in the mount; 4 Number of shared families/coefficient of similarity (%); 5 Number of shared genera/coefficient of similarity (%); 6 Number of shared species/coefficient of similarity (%). JG: Mount Jinggangshan region; WY: Mount Wuyishan region; QY: Mount Qiyunshan; LS: Mount Lushan; NL: Nanling Mountains; WL: Wuling Range; SNJ: Mount Shennongxia; EM, Mount Emeishan; TW: Taiwan Mountains; HN: Hainan Mountains; TB: Mount Taibeishan; XSBN: Mount Xishuangbangna; GLG: Mount Gaoligongshan.
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Mount Jinggangshan region is at the northern distribution limit of Gnetaceae and Samydaceae. They occur in Mount Jinggangshan region and Nanling Mountains, but not in Mount Wuyishan region, Wuling Range or Mount Lushan. The Nageiaceae, Capparaceae and Erythroxylaceae, mainly in southern China, are also in Mount Jinggangshan region and Mount Wuyishan region, but not Mount Lushan and Wuling Range. The Zygophyllaceae, mainly in northern dry areas, is in Mount Jinggangshan, but not Mount Qiyunshan, Nanling Mountains or Mount Wuyishan region. Mount Jinggangshan region marks the southernmost distribution of Zygophyllaceae.

Some important tropical or south tropical genera such as Tsoongiodendron, Exbucklandia, Altingia, Disanthus, Semiliquidambar, Cryptocarya, Anneslea, Garcinia, Pithecellobium, Heteropanax, Psychotria, Passiflora, Blastus and Bredia are mainly in southern China. They occur in Mount Jinggangshan region, but not Mount Lushan or Wuling Range.

Some typical genera in central China, such as Saruma, Clematoclethra and Dickinsia [34], occur in Mount Jinggangshan region, but not Mount Wuyishan region, Mount Qiyunshan or Nanling Mountains. Some genera endemic to central and eastern China, including Eucommia, Bretschneidera, Sargentodoxa, Pseudotaxus, Cyclocarya, Changnienia, Eomecon, Changium, Fortunearia, Sheareria, are also in Mount Jinggangshan. The following genera are mainly in Mount Jinggangshan region and eastern China: Ginkgo, Cunninghamia, Bretschneidera, Monimopetalum, Hilliella, Gelidocalamus, Speirantha.

From Figure 1 and Figure 2, it can be seen that tropical genera (Tr) increase and temperate genera (Tm) decrease from north to south and from west to east. The rate of change is expressed by Tr/Tm value. For example, Mount Jinggangshan region has 452 tropical genera and slightly less than 472 temperate genera, so Tr/Tm=452/472=0.96. The Mount Wuyishan region has 400 tropical genera and slight less than 425 temperate genera, so Tr/Tm=0.94.

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Figure 2. Numbers and ratios of the tropical genera and the temperate genera in Mount Jinggangshan region and other twelve mounts in China.

https://doi.org/10.1371/journal.pone.0075834.g002

Discussion and Conclusions

The flora of the Mount Jinggangshan region includes abundant primitive gymnosperms and angiosperms. Among them are 9 families of gymnosperms with 17 genera and 23 species. They account for 75.0%, 50.0% and 10.0% of wild gymnosperms (12 families, 34 genera and 230 species) in China respectively, and 69.2%, 23.9% and 2.9% of all gymnosperms (13 families, 71 genera and 800 species) in the world. Ginkgo biloba can be traced back to the Permian. Amentotaxus argotaenia, Fokienia hodginsii and Podocarpus macrophyllus were widely distributed in Cretaceous. Abies beshanzuensis var. ziyuanensis, endemic to China and critically endangered (<500 individuals in 2013), is looked as an evidence that Abies migrated from north to south and from high elevation to low elevation during the Quaternary ice age and withdraw to north and to higher elevation in post glacial [35]. The Mount Jinggangshan region is the largest distribution area of Abies beshanzuensis var. ziyuanensis (<350 individuals in 2013) and the lowest distribution area of Abies at the same latitude in the world.

To angiosperms, there are plentiful primitive representatives of the ancient families, such as Tsoongiodendron odorum, Manglietia fordiana and Magnolia officinalis, belonging to Magnoliaceae, and Exbucklandia tonkinensis, Hamamelis mollis and Fortunearia sinensis, belonging to Hamamelidaceae. The fossils of Exbucklandia (in the Paleocene in North America and China), Fortunearia (in the Oligocene in German and Pliocene in Japan), and Hamamelis (in Cretaceous in Sweden and Eocene in China) have been discovered [32,36-38]. Other primitive taxa include Disanthus cercidifolia subsp. longipes, Eucommia ulmoides, Cyclocarya paliurus, Sargentodoxa cuneata, Bretschneidera sinensis, Camptotheca acuminata, and Eurycorymbus cavaleriei.

As a refugium, there are many communities consisting of relict species as the dominant species, such as Amentotaxus argotaenia, Cunninghamia lanceolata, Fokienia hodginsii, Nageia nagi, Pseudotaxus chienii, Taxus wallichiana var. mairei, Tsuga chinensis, Exbucklandia tonkinensis and Disanthus cercidifolia subsp. longipes.

The seed plant flora of Mount Jinggangshan region includes many endemics genera and species. There are 44 genera endemics to China and 1146 species endemics to China (in 424 genera and 131 families), accounting for 17.53% and 39.52% of all in China respectively. The number of endemics genera are equivalent to other mounts in rich biodiversity, such as Mount Shennongjia (43 endemic genera), Mount Wuyishan region (38), Wuling Range (37) and Mount Emeishan (39). The important families with more endemic species include Rosaceae, Lauraceae, Theaceae, Labiatae, Papilionaceae, Fagaceae and Ericaceae.

There are 17 local endemic species in Mount Jinggangshan region, a number similar to Mount of Wuyishan region (20 species), such as Rhododendron jinggangshanicum, Rhododendron strigosum, Rhododendron xiaoxidongense, Neillia jinggngahsnensis, Rubus glandulosocarpus, Actinidia chinensis var. jinggangshanensis, Acer cordatum var. jinggangshanensis, Vitis jinggangensis, Trichosanthes jinggangshanica, Impatiens jinggangensis, Impatiens jinggangensis var. pauciflora, Hemiboea subacaulis var. jiangxiensis, which are restricted to the Mount Jinggangshan region. Rhododendron kiangsiense, Rhododendron hypoblematosum, Rhododendron crassistylum, Elaeagnus jiangxiensis and Gelidocalamus stellatus are centered in the Mount Jinggangshan region, but also occur in the Luoxiao Mountains.

The greater part of the Chinese flora was usually considered to belong to the Holarctic Kingdom, with a small part in the south belonging to the Paleotropic kingdom in the early days [39]. Hungta Chang [40] and Zhengyi Wu [41] successively proposed that the Chinese flora should be a separate floristic kingdom, the East Asian Kingdom, based on its primitiveness, origin and integrity. Based on Wu’s concept, the flora of Mount Jinggangshan region belongs to the East Asian Kingdom, the Sino-Japanese subkingdom, east China Region and south Jiangxi-east Hunan sub-region. As discussed above, the flora of Mount Jinggangshan region is closely related to the flora of Mount Wuyishan region. They share endemic genera and other elements, therefore both of them should be typical of the flora of eastern China.

However, the flora of Mount Jinggangshan region has obvious transitional properties. Many endemic genera are characteristic of either eastern or central China. The Mount Jinggangshan region is also the southeastern boundary for some typical genera of central China. Furthermore, the Mount Jinggangshan region is a corridor between northern and southern China. Renlin Liu implied that the Mount Jinggangshan region has many tropical elements, especially characterized by Fagaceae, Lauraceae, Theaceae, Hamamelidaceae, and Magnoliaceae [42]. These families make up the dominant elements of the evergreen broadleaved forests as in Nanling Mountains.

In conclusion, Mount Jinggangshan region is one of the richest regions in biodiversity in southeastern China and an important refugium for Tertiary relicts. The region in the central section of the Luoxiao Mountains is an important north-south floristic passageway and is also a boundary between the floras of eastern, central and south China.

Acknowledgments

We are deeply grateful to David E. Boufford of the Harvard University Herbaria for his good revisions to this paper. We also thank to Zhen Li, Pengyuan Li, Huijuan Jing, Shi Shi, Zifa Huang, Xiangming Zeng, and Jian Zhao for their fieldwork in the Mount Jinggangshan region from 2009 to 2011.

Author Contributions

Conceived and designed the experiments: WL. Performed the experiments: LW. Analyzed the data: WL. Contributed reagents/materials/analysis tools: QF CC. Wrote the manuscript: LW. Field works: LW WL CC QF.

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