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Research Article
A new synonym of Enkianthus perulatus (Ericaceae) in East Asia, based on morphological and molecular evidence
expand article infoHua Liang‡, Lu Jiang‡, Danqi Li‡§, Yi Yang‡, Dengmei Fan‡, Zhiyong Zhang‡
‡ Jiangxi Agricultural University, Nanchang, China
§ Chinese Academy of Sciences, Jiujiang, China

Corresponding author: Dengmei Fan ( dmf.625@163.com )

Corresponding author: Zhiyong Zhang ( zhangzy@mail.jxau.edu.cn )

Academic editor: Hugo de Boer
© 2022 Hua Liang, Lu Jiang, Danqi Li, Yi Yang, Dengmei Fan, Zhiyong Zhang.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Liang H, Jiang L, Li D, Yang Y, Fan D, Zhang Z (2022) A new synonym of Enkianthus perulatus (Ericaceae) in East Asia, based on morphological and molecular evidence. PhytoKeys 214: 61-74. https://doi.org/10.3897/phytokeys.214.94294
Open Access

Abstract

Enkianthus calophyllus was once treated as a synonym of E. serrulatus. However, field observations indicate that E. calophyllus is distinct from E. serrulatus but resembles E. perulatus in flowers, leaves, fruits and seeds. Hence, a taxonomic revision of these species was conducted based on morphological comparisons of flowers, leaves, fruits and seeds, as well as molecular analyses of nuclear ribosomal internal transcribed spacer (nrITS) and six plastid DNA markers (psbA-trnH, rpl32-trnL, trnL-trnF, rps16-trnQ, psbJ-petA and matK). The morphological and molecular results reject the synonymization of E. calophyllus with E. serrulatus, and instead show it to be placed in a clade with E. perulatus. Based on molecular evidence and a reassessment of the morphology we synonymize E. calophyllus with the older name E. perulatus.

Keywords

Enkianthus calophyllus, Enkianthus perulatus, morphology, phylogeny, synonym

Introduction

Enkianthus Lour. is a small genus in Ericaceae with about 12–17 species (Anderberg 1994; Kron et al. 2002; Fang and Stevens 2005). It is only distributed in East Asia, and most of its component species are in China and Japan. Enkianthus is ornamentally important for its elegant flowers and ecologically valuable because it always dominates in subtropical montane elfin forests (Hsu 1982). Accumulating molecular phylogenetic evidence suggests that Enkianthus is the first diverging lineage of Ericaceae (Kron and Chase 1993; Kron 1996; Morton et al. 1996; Anderberg et al. 2002; Kron et al. 2002), indicating that this genus is key to understand the evolution of Ericaceae.

Species in Enkianthus are shrubs or small trees, leaves blade serrate or subentire, inflorescences often umbels and racemes, corollas broadly campanulate to urceolate, capsule loculicidal, seeds often lamellate-winged (Fang and Stevens 2005). The species of Enkianthus vary in leaf texture, inflorescences structure, corolla shape, and anther morphology, pollen and seed (Cheng and Lai 1988; Anderberg 1994; Kron et al. 2002; Sarwar and Takahashi 2006). Infrageneric relationships of Enkianthus were studied by Anderberg (1994), who proposed a classification comprising four sections (sect. Enkianthus, sect. Andromedina, sect. Enkiantella and sect. Meisteria). Among them, sect. Enkianthus is monophyletic according to phylogenetic analyses (Tsutsumi and Hirayama 2012). However, due to the variable morphology in this genus (Hsu 1982), classification of some species, especially those with wide distribution range, remains controversial.

During the past years, we have found several unique Enkianthus populations in montane areas of Zhejiang and Jiangxi Province in China (Fig. 1). These plants are 1–3 m tall, white urceolate flowers with distinct basal gibbosities, rhombic-elliptic leaves and erect capsule. After scrutinizing the protologue and type specimens, we found that our collections matched the description of E. calophyllus T.Z. Hsu exactly (Fig. 2, Suppl. material 1: fig. S1B; Hsu 1985). When Fang and Stevens (2005) treated E. calophyllus as a synonym of E. serrulatus (E.H. Wils.) C.K. Schneid. (Fig. 3, Suppl. material 1: fig. S1C) in Flora of China (FOC), flowering specimens of E. calophyllus were lacking (Fang and Stevens 2005). However, our collections and E. calophyllus differ from E. serrulatus by urceolate corollas with distinct basal gibbosities, margin with ciliate, smaller fruits (0.5–0.7 cm × 0.3–0.4 cm) and seeds without distinct wings (Figs 2, 3; Schneider 1911; Hsu 1985), and such characters were described for E. perulatus C.K. Schneid. from Japan (Fig. 4, Suppl. material 1: fig. S1A; Schneider 1911). These observations raise a question about the taxonomic status of E. calophyllus and the identity of our collections. To identify our new collections and clarify the taxonomic status of E. calophyllus, morphological comparisons and molecular phylogenetic analyses were performed to study the taxonomic relationships amongst E. calophyllus, E. perulatus, E. serrulatus and our new collections.

Figure 1. 

Distribution of Enkianthus perulatus based on specimen records and our field investigation. Black triangles indicate E. perulatus; red triangles indicate E. calophyllus (= E. perulatus).

Figure 2. 

Morphology of Enkianthus calophyllus (= E. perulatus) A flowering branch B fruiting branch C leaves D flowers E seeds F leaf margin. A–F photographed by H. Liang.

Figure 3. 

Morphology of Enkianthus serrulatus A flowering branch B fruiting branch C leaves D flowers E seeds F leaf margin. A–F photographed by H. Liang.

Figure 4. 

Morphology of Enkianthus perulatus A flowering branch (Hooker 1870) B fruiting branch (USF 121400) C leaves (K-000780276) D flowers (Hooker 1870) E seeds F leaf margin (K-000780276). E photographed by H. Liang.

Materials and methods

Morphological studies

Our collections and digital images of Enkianthus perulatus, E. calophyllus and E. serrulatus from the Chinese Virtual Herbarium (http://www.cvh.ac.cn/), the web of Plants of Taiwan (http://tai2.ntu.edu.tw), the Kingdonia (http://kun.kingdonia.org/) and the Global Biodiversity Information Facility (https://www.gbif.org/) were examined to test whether there are significant differences in leaves and fruits of the three species. The high-resolution images of those Enkianthus species were taken from specimens deposited at CDBI, CSFI, CSH, GA, GXMI, GZTM, JIU, JXAU, K, KUN, L, LGB, NAS, NTUF, P, PE, SYS, TAI, USF, and ZY. We randomly selected three well-preserved leaves and/or fruits from each specimen (28 specimens of E. serrulatus, 18 of E. calophyllus and 15 of E. perulatus). Among them, nine specimens of E. calophyllus and seven of E. serrulatus were collected by ourselves. Eight morphological characters, i.e., leaf length, leaf width, ratio of leaf length/width, pedicel length, fruit length, fruit width, ratio of fruit length/width and carpopodium length, were measured in this study. ImageJ (Rasband 1997) was applied to the examination of the focal characters.

ANOVA was performed to test the significance of pairwise difference of eight characters using SPSS 26. Principal Component Analysis (PCA) was performed in ORIGIN 2021 to investigate the morphological variations among E. perulatus, E. calophyllus and E. serrulatus. Morphological analysis was not carried out for flowers and seeds, because there were only a few specimens available for analysis.

Sample collection, DNA extraction, PCR amplification, and sequencing

We collected 19 samples from 13 populations (1–3 individuals per population) of five Enkianthus species (Suppl. material 1: table S1). Of these, 17 samples belonged to species of sect. Enkianthus, i.e., six of E. calophyllus, four of E. serrulatus, four of E. perulatus, and three of E. quinqueflorus Lour. (Suppl. material 1: table S1). Two species of sect. Enkiantella, E. chinensis Franch. and E. deflexus (Griff.) Schneid. were also collected. Based on previous studies (Tsutsumi and Hirayama 2012), we downloaded nuclear ribosomal internal transcribed spacer (nrITS) and plastid DNA markers of other Enkianthus species from the National Center for Biotechnology Information’s (NCBI; http://www.ncbi.nlm.nih.gov/) nucleotide database (Suppl. material 1: table S1). In addition, we selected species of genera Rhododendron, Vaccinium and Clethra as outgroups to carry out the phylogenetic analysis of sect. Enkianthus according to previous studies (Kron et al. 2002; Liu et al. 2014). Voucher specimens are deposited in the Herbarium of Jiangxi Agricultural University (JXAU). Total genomic DNA were extracted from the silica-dried leaves using a modified cetyltrimethylammonium bromide method (Doyle and Doyle 1987). Six chloroplast DNA (cpDNA) regions (psbA-trnH, rpL32-trnL, trnL-trnF, rps16-trnQ, psbJ-petA and matK) (Taberlet et al. 1991; Sang et al. 1997; Shaw et al. 2007) and nrITS (Sun et al. 1994) in 19 individuals from 13 populations of Enkianthus, were PCR amplified and sequenced. The PCR amplification protocols followed Cheng et al. (2021), and primers are listed in supplementary (Suppl. material 1: table S2). Newly generated sequences in this study are deposited in GenBank (Suppl. material 1: table S1).

Phylogenetic analysis with cpDNA and nrDNA sequence data

The matrices of DNA sequences were aligned using MAFFT v.7 (Katoh and Standley 2013), and improved manually using BioEdit 7.0.9 (Hall 1999). Bayesian inference (BI) and maximum likelihood (ML) were used for phylogenetic analysis on the CIPRES Science Gateway 3.3 (www.phylo.org; Miller et al. 2015) with the best-fit model of DNA substitution estimated by jModelTest v.2.1.4 (Darriba et al. 2012). The alignments of nrITS, matK and the concatenated plastid DNA (psbA-trnH + rpL32-trnL + trnL-trnF + rps16-trnQ + psbJ-petA + matK) were analyzed with GTR + G, GTR and GTR + G + I model, separately. We reconstructed a matK phylogeny of Enkianthus because the other five chloroplast DNA regions were sequenced only in a subset of species. Bayesian analysis was constructed using MrBayes v.3.2.7 (Ronquist et al. 2012). We performed two independent BI runs with one cold and three heated chains for 10,000,000 Markov chain Monte Carlo generations. We sampled trees every 1,000 generations and discarded the first 25% generations as burn-in. ML analysis was conducted by RAxML-HPC (Stamatakis 2014) with 1000 bootstrap replications.

Results and discussion

Morphological analyses

Principal Component Analysis (PCA) showed that our collections of Enkianthus calophyllus clustered with their type specimens and the 95% confidence ellipse of E. calophyllus intersected marginally that of E. serrulatus; however, the ellipse of E. calophyllus almost overlapped with that of E. perulatus (Fig. 5). In addition, pairwise comparisons of the eight morphological traits among E. perulatus, E. calophyllus and E. serrulatus (ANOVA analysis) showed there were significant differences between E. calophyllus and E. serrulatus (Fig. 6), but no significant difference between E. calophyllus and E. perulatus in all compared traits except for leaf width and ratio of leaf length/width (Fig. 6). Last but not least, morphological observation also found that E. calophyllus was almost the same as E. perulatus, but differs from E. serrulatus in flowers and seeds (Table 1). These results suggest that E. calophyllus should be conspecific with E. perulatus rather than E. serrulatus. The morphological description and comparison are elaborated in Table 1.

Figure 5. 

The Principal Component Analysis (PCA) plot for the morphological variations amongst Enkianthus perulatus, E. calophyllus (= E. perulatus) and E. serrulatus. Red, gray and blue triangles represent E. perulatus, E. calophyllus and E. serrulatus, respectively. Yellow circles indicate type specimens. The confidence ellipse level is 95%.

Figure 6. 

Box plots of the character comparisons amongst Enkianthus perulatus (EP), E. calophyllus (= E. perulatus; EC) and E. serrulatus (ES) A leaf length B leaf width C ratio of leaf length/width D pedicel length E fruit length F fruit width G ratio of fruit length/width H carpopodium length. The different lowercases on the top of the vertical line indicate significant differences (P < 0.05) between the three taxa.

Table 1.
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Comparisons of morphological characters amongst Enkianthus perulatus, E. calophyllus and E. serrulatus.

Characters E. perulatus E. calophyllus E. serrulatus
Habit shrubs deciduous, 1–2 m tall shrubs deciduous, 1–3 m tall shrubs or small tree, deciduous, 3–6 m tall
Texture of leaf Papery Papery papery or thickly papery
Petiole 0.5–1.2 cm 0.5–1.4 cm 0.7–2.1 cm
Leaf blade oblong, obovate-oblong; 2–4 (–5) cm × 0.8–2.0 cm rhombic-elliptic, elliptic; 2.5–5.0 cm × 1.5–3.0 cm elliptic, oblong-elliptic or obovate-elliptic; 6–9 (–11) cm × 3–4 (–5) cm
Leaf margin margin with ciliate margin with ciliate margin without ciliate
Inflorescence umbellate, 1–5-flowered umbellate, 1–5-flowered umbellate, 2–6 (–9)-flowered
Corolla urceolate with distinct basal gibbosities; white; 0.6–0.7 cm × 0.5–0.7 cm urceolate with distinct basal gibbosities; white; 0.6–0.8 cm × 0.5–0.8 cm oblong-urceolate without distinct basal gibbosities; greenish-white; 1.3–1.6 cm × 0.8–1.0 cm
Length/width of Corolla 1.0–1.3 1.0–1.4 1.5–2.0
Pedicel 1–2 cm 1.4–2.2 cm 2–3 cm
Fruit capsule erect, oblong, 0.6–0.7 cm × 0.3–0.4 cm capsule erect, oblong, 0.5–0.7 cm × 0.3–0.4 cm capsule erect, oblong, 0.8–1.2 cm × 0.5–0.8 cm
Seed without distinct wings without distinct wings with distinct wings
Distribution Taiwan China, Japan (Honshu, Shikoku and Kyushu) China (Zhejiang, Fujian, Jiangxi) China (Jiangxi, Hubei, Hunan, Guangdong, Guangxi, Guizhou, Chongqing, Sichuan, Yunnan)
Altitude 200–1600 m 600–1200 m 800–1800 m

Phylogenetic relationships

Alignment length of nrITS sequences based on 10 species (E. calophyllus = E. perulatus) of Enkianthus (approx. 83% species of Enkianthus, Fang and Stevens 2005) is 595 bp, including 75 variable sites and 60 parsimony informative sites. Alignments of matK consisting of the same 10 species contain 755 constant sites, 24 variable sites and 9 parsimony informative sites. The concatenated length for six plastid DNA fragments based on five species of Enkianthus is 4,768 bp, and the matrix contains 124 variable sites and 85 parsimony informative sites in total. Phylogenetic analyses based on nrITS or matK supported that sect. Enkianthus was a monophyletic clade. Although the nrITS tree showed that six accessions of E. calophyllus form a monophyletic clade (bootstrap value, BS = 88, Bayesian posterior probability, PP = 0.86; Fig. 7A), this clade was nested within E. perulatus (BS = 100, PP = 1; Fig. 7A). Notably, E. calophyllus intermingled with E. perulatus, forming a highly supported clade in the matK tree (BS = 97, PP = 0.97; Suppl. material 1: fig. S2). Furthermore, the six plastid DNA tree supported that the monophyly of E. perulatus and E. calophyllus was recovered again (BS = 100, PP = 1; Fig. 7B), and E. calophyllus is paraphyletic with respect to E. perulatus (Fig. 7B). In all the trees, E. serrulatus clustered with E. quinqueflorus rather than with E. perulatus.

Taken together, we propose that E. calophyllus should be recognized as a new synonym of E. perulatus rather than the synonym of E. serrulatus as suggested by Fang and Stevens (2005). In addition, E. serrulatus together with E. quinqueflorus may represent a well differentiated lineage relative to E. perulatus.

Figure 7. 

Phylogenetic relationships of Enkianthus A phylogenetic tree based on nuclear DNA loci (ITS) B phylogenetic tree based on six plastid markers (psbA-trnH, rpl32-trnL, trnL-trnF, rps16-trnQ, psbJ-petA and matK). Numbers and asterisksa above branches are Maximum Likelihood bootstrap values / Bayesian posterior probability (> 50%). Asterisks indicate that the support values are 100%. The phylogenetic positions of E. perulatus, E. calophyllus (= E. perulatus) and E. serrulatus are highlighted in red, grey and blue, respectively.

Taxonomic treatments

Enkianthus perulatus (Miq.) C.K. Schneid

Andromeda perulata Miq., Ann. Mus. Bot. Lugduno-Batavi 1: 31. 1863. Basionym. Type: Japan. W. Botanicus 57 (holotype: L-0007044!, Suppl. material 1: fig. S1A).

Enkianthus japonicus Hook. f., Bot. Mag. 96: 5822. 1870. Type: Japan. 1860, R. Alcock s.n. (holotype: K-000780276!). ≡ Enkianthus perulatus var. japonicus (Hook. f.) Nakai, J. Jap. Bot. 12(12): 896. 1936. ≡ Enkianthus perulatus f. japonicus (Hook. f.) Kitam., Acta Phytotax. Geobot. 25: 36. 1972.

—Enkianthus taiwanianus S.S. Ying, Quart. J. Chin. Forest. 9: 145. 1976. Type: China. Taoyuan City, Peichiatienshan, 1976, Ying 5301 (holotype: NTUF-F00006903!; isotype: NTUF-F00008372, NTUF-F00008372, NTUF-F00008372, NTUF-F00008372!). ≡ Enkianthus perulatus var. taiwanianus (S.S. Ying) Y.C. Liu, Tr. Taiwan 514. 1988.

—Enkianthus calophyllus T.Z. Hsu, Acta Bot. Yunnan. 7(2): 151–152. 1985. syn. nov. Type: China. Zhejiang: Jingning County, 16 May 1959, S.Y. Zhang 5286 (holotype: PE-00005629!, Suppl. material 1: fig. S1B; isotype: NAS-00063024!, KUN-0001289!).

Description

Deciduous shrubs up to 3 m tall. Branchlets terete, glabrous. Leaves alternate on young shoots, generally 4–6 crowded at apex of branchlets; petioles 0.5–1.4 cm long; blades papery, oblong, obovate-oblong, rhombic-elliptic or elliptic, 2–5 cm long, 0.8–3.0 cm wide, apex acuminate, mucronate, base attenuate-cuneate or cuneate, margin ciliate, upper surface glabrous except sparsely pubescent on midrib, lower surface villous along lower part of midrib. Inflorescences terminal, umbellate, 1–5-flowered, pendulous. Pedicels 1.0–2.2 cm long, glabrous, reflexed at flowering, straight at fruiting. Bracts lanceolate, white or green, 0.8–1.8 cm long, caducous. Calyx lobes triangular, 2–4 mm long, glabrous, deeply 5-lobed. Corollas urceolate, with distinct basal gibbosities, white, 6–8 mm long, 5–8 mm wide, shallowly 5-lobcd; lobes widely ovate, obtuse, much reflexed. Stamens 10, ca. 2/3 as long as corolla; filaments villous at base, anthers with 2 awns on upper dorsal side. Ovaries glabrous. Capsules erect, oblong, 5-ridged, 5–7 mm long, 3–4 mm wide. Seeds compressed, narrowly oblong, ca. 4 mm long, 1 mm wide, with marginal-like ridges, without distinct wings.

Phenology

Flowering from April to May; fruiting from May to November.

Distribution and habitat

Enkianthus perulatus is distributed in China (Zhejiang Fujian, Jiangxi and Taiwan) and Japan (Honshu, Shikoku and Kyushu) (Fig. 1). It grows on open rocky slopes, mountain slopes, cliffs, serpentine area, by roadsides or at forest margins at altitudes of 200–1600 m.

Additional specimens examined

China. Zhejiang: Yueqing City, Yandangshan, 6 April 2015, X.Y. Ye 2015040609 (CSH); ibidem, 20 May 2019, H. Liang LSBZ-259 (JXAU). Fujian: Taining County, Xinqiao Town, 16 June 1978, G.L. Cai 445 (KUN). Jiangxi: Jinggangshan City, Jinggang Mountains, 15 July 1965, S.K. Lai et al. 4466 (LBG); Lichuan County, Huixianfeng, 20 October 1985, S.K. Lai & D.F. Huang 473 (LBG); ibidem, 16 November 2021, H. Liang 088 (JXAU). Taiwan: Taibei City, Tunlu, 14 April 1935, T. Suzuki 19235 (TAI); Taibei City, Lupeishan, 16 April 1991, Y.B. Cheng & T.S. Hsieh 1202 (TAI); Taoyuan City, Peichatienshan, 28 September 1984, R.T. Li 3532 (TAI); Chiayi City, Alishan, 5 April 1982, Y.F. Chen 4604 (TAI), ibidem, 17 May 1982, Y.F. Chen 4902 (TAI). Japan. Honshu: Wakayama-Prefecture, Ohdaigahara Mountains, 23 May 1925, S. Saito (PE); Shizuoka-Prefecture, Tagata-gun, Sanagi Mountains, 24 April 1952, M. Furuse 24733 (PE); Nagano-Prefecture, Iida-shi, 27 April 1962, M. Furuse 39658 (PE); Aichi-Prefecture, Minami-shitara-gun, 22 April 1978, M. Furuse 12532 (PE); Aichi-Prefecture, Shinshiro-shi, 6 August 1978, M. Furuse 13039 (PE).

Additional specimens of Enkianthus serrulatus examined

China. Guangxi: Debao County, 25 April 1977, D. Fang et al. 3-219 (GXMI); Longsheng County, 24 August 2018, H. Liang LSBZ-218 (JXAU); Longlin County, 17 May 1977, T.H. Wei 3-0606 (GXMI); Xingan County, 26 July 1997, G.Z. Li 15137 (PE). Guizhou: Chishui City, 24 May 2020, H. Liang LSBZ-297 (JXAU); Leishan County, 13 June 2020, H. Liang LSBZ-323 (JXAU); Songtao County, 22 July 1959, T.P. Zhu et al. 1592 (KUN); Suiyang County, 11 May 2010, Y.F. Zhou KKS101197 (ZY); Zhengan County, 14 October 2014, H.W. Zhang 520324141014031LY (GZTM). Hubei: Lichuan County, 7 October 1980, B. Bartholomew et al. 2014 (PE); Tongshanx County, 14 May 2017, H.Y. Zhan et al. LXP5905 (LBG); Yichang City, 19 March 2017, D.G. Zhang et al. ZCJ170319117 (JIU). Hunan: Sangzhi County, 4 August 2017, Z.Y. Zhang et al. LSBZ-142 (JXAU); Xinning City, 5 July 2017, Z.K. Liu LSBZ-135 (JXAU); Yongshun County, 3 August 2017, Z.Y. Zhang et al. LSBZ-138 (JXAU); Zhangjiajie City, 11 September 2015, H. Zhou & D.S. Zhou 15091113 (CSFI). Jiangxi: Jinggangshan City, 27 August 2020, Y.F. Liu LSBZ-365 (JXAU); Luxi County, 23 June 1984, M.X. Nie 113 (LBG); Suichuan County, 24 June 2016, Z.C. Liu et al. Lxp-13-18312 (SYS); Wuning County, 22 May 2014, Y.H. Zhan et al. LXP0912 (LBG). Sichuan: Hechuan City, 1934, D.J. Yu 3112 (PE); Leibo County, 3 July 1983, Q.S. Zhao & Z.J. Zhao 121212 (PE); Xuyong County, 17 September 2013, W.B. Ju & H.N. Deng HGX13668 (CDBI). Yunnan: Maguan County, 31 July 1961, S.G. Wu 3597 (KUN); Suijiang County, 4 May 1973, B.X. Sun 0112 (KUN). Chongqing: Fengjie County, 28 June 1958, M.Y. Fang 24515 (NAS); Shizhu County, 2 June 1978, W.H. Wang 1571 (CDBI).

Acknowledgements

We are grateful to the curators of the herbaria of CDBI, CSFI, CSH, GA, GXMI, GZTM, JIU, JXAU, K, KUN, L, LGB, NAS, NTUF, P, PE, SYS, TAI, USF and ZY for access to the specimens for our study. This study is supported by the National Natural Science Foundation of China (No. 31660059, No. 31960049) and the Training Plan for Academic and Technical Leaders (leading talents) of major disciplines in Jiangxi, China (No. 20213BCJ22006).

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Supplementary material

Supplementary material 1 

Supplementary data

Hua Liang, Lu Jiang, Danqi Li, Yi Yang, Dengmei Fan, Zhiyong Zhang

Data type: figures and tables (word document)

Explanation note: Type specimen of Enkianthus perulatus, E. calophyllus and E. serrulatus. Phylogenetic relationships of Enkianthus based on matK. Sample collection information and GenBank accessions in this study. Sequences of primers used for PCR amplification and sequencing.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
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