Research Article
Research Article
Paraphlomis hsiwenii (Lamiaceae), a new species from the limestone area of Guangxi, China
expand article infoYa-Ping Chen, Jin-Fei Xiao, Chun-Lei Xiang, Xiong Li§
‡ Kunming Institute of Botany, Chinese Academy of Sciences, Kunming, China
§ Guangxi Forestry Inventory and Planning Institute, Nanning, China
Open Access


The indumentum of nutlets is shown to be of phylogenetic importance in previous molecular phylogenetic studies of Paraphlomis, a genus of Lamiaceae with approximately 30 species distributed mainly in southern China and Southeast Asia. Nearly half the species of Paraphlomis are known from limestone areas. In this study, we described and illustrated a new species, P. hsiwenii, from the karst mountain forests in Guangxi Zhuang Autonomous Region, China. Our molecular phylogenetic analyses revealed that P. hsiwenii is recovered in a clade consisting of species with hairy nutlets. The new species is morphologically most similar to P. pagantha from the same clade, but they differ in the morphology of lamina bases, length of pedicels and calyces, as well as the morphology of upper corolla lips.


Ajugoides, karst, Matsumurella, nutlet, Paraphlomideae


In a recently updated phylogenetic and taxonomic study of Lamioideae, the tribe Paraphlomideae was established to accommodate three genera, Ajugoides Makino, Matsumurella Makino, and Paraphlomis (Prain) Prain (Bendiksby et al. 2011). Ajugoides is a monotypic genus endemic to Japan and Matsumurella comprises five species distributed in East Asia (Harley et al. 2004; Bendiksby et al. 2011). As the largest genus of Paraphlomideae, Paraphlomis consists of ca. 30 species occurring mainly in China (especially in the south of the Yangtze River) and Southeast Asia (Wu and Li 1977; Li and Hedge 1994; Harley et al. 2004; Chen et al. 2021).

Morphologically, Paraphlomis can be distinguished by its herbaceous habit with stoloniferous stems and simple hairs, two to many-flowered verticillasters, actinomorphic and tubular to obconical calyces, 2-lipped (1/3) corollas, and apically truncate ovaries (Wu and Li 1977; Li and Hedge 1994; Harley et al. 2004; Chen et al. 2021). Two sections and five series were recognized in a previous infrageneric classification of Paraphlomis from China, divided based on the shape of calyx (tubular or obconical) and calyx teeth (e.g., conspicuous or inconspicuous, broadly triangular or subulate) (Li 1965; Wu and Li 1977). However, the most recent molecular phylogenetic study of Paraphlomis indicated that nutlet morphology (e.g., glabrous or hairy, obviously inflated or not) rather than the above-mentioned calyx characters is of phylogenetic value for the subdivision of the genus.

Species of Paraphlomis are mostly accustomed to shady and moist places in tropical and subtropical evergreen and mixed forests, and nearly half of the species are karst-adapted (Wu and Li 1977; Li and Hedge 1994; Zhang et al. 2020; Chen et al. 2022a). During our recent field investigations in the limestone area of Diding Natural Reserve in Guangxi Zhuang Autonomous Region, China, we found a putative new species of Paraphlomis which is characterized by hairy nutlets. We further confirmed its specific status as a new species and placement within the genus based on molecular phylogenetic and morphological evidence, and named it P. hsiwenii Y.P.Chen & XiongLi.

Materials and methods

Molecular phylogenetic analyses

We sampled a total of 34 accessions representing 19 species and four varieties/subspecies of Paraphlomis and two species of Matsumurella as the ingroups, and included two taxa, Phlomis fruticosa L. and Phlomoides dentosa var. glabrescens (Danguy) C.L. Xiang & H. Peng from tribe Phlomideae as the outgroups. Only two accessions of the new species and one accession of Paraphlomis pagantha Doan were newly sampled and sequenced here, while sequences of the remaining accessions were all retrieved from our previous studies (Chen et al. 2021, 2022a, b).

Total genomic DNA was extracted from silica-gel-dried leaf material using the modified CTAB method (Doyle and Doyle 1987). We selected five DNA markers for the phylogenetic reconstruction following previous studies (Chen et al. 2021, 2022a, b), i.e. the nuclear ribosomal internal and external transcribed spacers (ITS and ETS) and three plastid DNA regions (rpl32-trnL, rps16, and trnL-trnF). Primers used for the polymerase chain reaction (PCR) amplification and sequencing of the five regions, as well as the PCR mixtures and procedures, were the same as those described in Chen et al. (2021). Voucher information and GenBank accession numbers for all sequences are listed in Appendix 1.

Previous phylogenetic studies of Paraphlomis revealed significant topological incongruences between the nuclear and plastid trees (Chen et al. 2021, 2022a, b), therefore, we performed partitioned maximum likelihood (ML) and partitioned Bayesian inference (BI) analyses for the combined nuclear data set and combined plastid data set separately. Both the ML and BI analyses were conducted on the Cyberinfrastructure for Phylogenetic Research Science (CIPRES) Gateway (; Miller et al. 2010), using RAxML-HPC2 (Stamatakis 2014) and MrBayes (Ronquist et al. 2012), respectively. Detailed settings for the two analyses followed those described in Chen et al. (2019). TreeGraph 2 (Stover and Müller 2010) was employed to visualize and annotate the resulting trees.

Taxonomic studies

Type specimens and protologues for all species of Paraphlomis were collated. Specimens of the genus from 21 public herbaria (BM, CDBI, E, GNNU, GXMI, HAST, HIB, IBK, IBSC, JIU, JJF, K, KUN, KYO, MW, NAS, PE, SM, SZ, TI, and WUK; abbreviations follow Thiers 2022) were also checked for the morphological comparison of P. hsiwenii with other species of Paraphlomis. Living plants of some species of the genus were observed and collected during our field investigation, and these specimens were further used for the morphological comparison of the new species. Other taxonomic and floristic literature related to Paraphlomis was reviewed, and the terminology used by Li and Hedge (1994) was adopted here for the morphological description of the new species.

Results and discussion

A total of 15 sequences (i.e., the five DNA regions of the two accessions of P. hsiwenii and one accession of P. pagantha) were newly generated in the present study. The aligned length of the combined nuclear data set and combined plastid data set was 1251 bp (808 bp for ITS, 443 bp for ETS) and 2479 bp (850 bp for rpl32-trnL, 812 bp for rps16, 817 bp for trnL-trnF), respectively. The topologies of the BI and ML trees were largely consistent with each other, but the BI trees are slightly better resolved. Thus, only the Bayesian 50% majority-rule consensus trees of the two combined data sets are presented, the posterior probabilities (PP) and Bootstrap support (BS) values being superimposed on the nodes (Figs 1, 2).

Figure 1. 

Bayesian 50% majority-rule consensus tree of Paraphlomis based on combined nuclear (ITS and ETS) data set. Support values ≥ 0.50 PP or 50% BS are displayed above the branches (an “*” indicates a support value = 1.00 PP or 100% BS and a “-” indicates a support value < 50% BS). Multiple accessions of the same species are numbered according to Appendix 1.

Figure 2. 

Bayesian 50% majority-rule consensus tree of Paraphlomis based on combined plastid (rpl32-trnL, rps16, and trnL-trnF) data set. Support values ≥ 0.50 PP or 50% BS are displayed above the branches (an “*” indicates a support value = 1.00 PP or 100% BS). Multiple accessions of the same species are numbered according to Appendix 1.

Both the nuclear and plastid data sets recovered species of Matsumurella within the Paraphlomis clade (Fig. 1: PP = 1.00/BS = 100%; Fig. 2: PP = 1.00/BS = 98%), indicating that neither of the two genera is monophyletic. As for relationships within the Paraphlomis-Matsumurella clade, some deep nodes in the nuclear tree (Fig. 1) and most shallow nodes in the plastid tree (Fig. 2) are poorly resolved, and topological conflicts between the two trees can be immediately recognized, especially at the placements of P. javanica var. pteropoda D. Fang & K.J. Yan, P. albiflora (Hemsl.) Hand.-Mazz., and P. nana Y.P. Chen, C. Xiong & C.L. Xiang. The backbone topologies of Paraphlomis recovered in the present study are largely consistent with those of previous studies (Chen et al. 2021, 2022a, b), and the intergeneric relationships within Paraphlomideae and phylogenetic relationships within Paraphlomis have been discussed in Chen et al. (2021). Therefore, our following discussion will focus on the placement of the new species.

The two accessions of P. hsiwenii group together and form a strongly supported clade (Fig. 1: PP = 1.00/BS = 100%; Fig. 2: PP = 1.00/BS = 100%). In the plastid tree, relationships between P. hsiwenii and other species of Paraphlomis are not resolved (Fig. 2). In the nuclear tree, the new species is sister to P. jiangyongensis X.L. Yu & A. Liu but with weak support values (Fig. 1: PP = 0.66/BS = 54%). The two species are further placed within a robustly supported clade (Fig. 1: PP = 1.00/BS = 91%) together with P. albida Hand.-Mazz., P. gracilis (Hemsl.) Kudô, P. kwangtungensis C.Y. Wu & H.W. Li, P. pagantha, and Matsumurella yangsoensis (Y.Z. Sun) Bendiksby. This clade is corresponding to “Clade III” in Chen et al. (2021) and all its members have hairy nutlets/ovaries. The densely hispid and glandular nutlets of P. hsiwenii and its recovery within Clade III further support that nutlet morphology might be of phylogenetic significance for the infrageneric classification of Paraphlomis (Chen et al. 2021).

Morphologically, P. hsiwenii is most similar to P. pagantha, when comparing it with other species with hairy nutlets. For example, most species of Clade III are characterized by densely hispid or strigose stems and laminas, whereas both P. hsiwenii and P. pagantha have densely hispidulous stems and subglabrous to glabrous laminas (Fig. 3). They also share ovate laminas with serrate margins and triangular calyx teeth (Figs 3, 4). The two species differ mainly in the morphology of lamina bases, which are not decurrent in P. hsiwenii (Fig. 3) but obviously decurrent in P. pagantha. Another difference is that the length of pedicels and calyces is much longer in the new species (Fig. 4). Moreover, the upper corolla lips are ca. 6 mm wide with emarginate apices in P. hsiwenii (Fig. 4), but much narrower (ca. 3 mm wide) with entire apices in P. pagantha. Other morphological differences between the two species can be found in Table 1. Geographically, the new species is now only discovered from the limestone area in Diding Natural Reserve at the Sino-Vietnamese border, whereas P. pagantha usually grows in the evergreen forests in northern Vietnam and Hainan Province, China (Fig. 5). Notably, P. pagantha was treated as a synonym of P. lancidentata Y.Z. Sun by Suddee and Paton (2006). However, the nutlets of P. lancidentata are glabrous and the two species are recovered within different clades in the phylogenetic trees (Figs 1, 2).

Table 1.

Morphological comparisons between Paraphlomis hsiwenii and P. pagantha.

Characters P. hsiwenii P. pagantha
Lamina Papery, ovate, base not decurrent, glabrous above Papery to membranous, ovate to oblong, base decurrent, sparsely strigose above
Pedicel 2–3 mm long Approximately 1 mm long
Calyx Approximately 6 mm long, teeth ca. 2 mm long Approximately 4 mm long, teeth ca. 1 mm long
Corolla Tube purplish-red at upper part; upper lip greenish-yellow, ca. 8 × 6 mm, apex emarginate; lower lip ca. 8 × 10 mm, lateral lobes greenish-yellow Tube with purplish-red spots at upper part; upper lip yellow to white, ca. 7 × 3 mm, apex entire; lower lip ca. 6 × 6 mm, lateral lobes yellow to white
Figure 3. 

Morphology of Paraphlomis hsiwenii from the type locality A, B habitat C–E habit F leaves G stolons. Scale bar: 5 cm (A, C photographed by Xiao-Lei Ma B, D, E photographed by Xiong Li F photographed by Ya-Ping Chen G photographed by Jin-Fei Xiao).

Figure 4. 

Floral traits of Paraphlomis hsiwenii. A frontal view of flowers B dorsal view of flowers C lateral view of flowers D lateral view of calyces E frontal view of calyces F dissected calyx G dissected corolla and lateral view of corolla H lateral and frontal view of ovary I lateral view of nutlet. Scale bars: 2 mm (F); 4 mm (G); 500 μm (H) (A–C, G, H photographed by Jin-Fei Xiao D–F photographed by Ya-Ping Chen I photographed by Xiong Li).

Taxonomic treatment

Paraphlomis hsiwenii Y.P.Chen & XiongLi, sp. nov.

Figs 3, 4


China, Guangxi, Jingxi City, Nanpo Town, Longting, Diding Natural Reserve, among shrubs in forests of limestone area, 23°3'29.55"N, 105°57'13.82"E, alt. 1181 m, 25 Jun 2022, J.F. Xiao & X.L. Ma XJF095 (holotype: KUN!; isotypes: K!, KUN!, MO!, PE!).


Paraphlomis hsiwenii is morphologically most similar to P. pagantha, but differs in having laminas glabrous above (vs. sparsely strigose above), bases of laminas not decurrent (vs. decurrent), calyces ca. 6 mm long (vs. ca. 4 mm long) with teeth ca. 2 mm long (vs. ca. 1 mm long), and upper corolla lips emarginate at apex (vs. entire at apex).

Perennial herbs 50–120 cm tall, stoloniferous. Stems erect, simple or branched, obtusely 4-angled, densely retrorse hispidulous and glandular. Leaves opposite; lamina ovate, papery, 3–11 × 2.5–5 cm, apex acute to acuminate, margin serrate, base cuneate to broadly cuneate, adaxially green, glabrous, abaxially light green, purplish-green, or purple, densely glandular, lateral veins 3–5-paired; petioles 0.5–2 cm long, densely retrorse hispidulous and glandular. Verticillasters 2–14-flowered; bracteoles subulate, ca. 0.5 mm long, early deciduous; pedicels 2–3 mm long, densely retrorse hispidulous and glandular. Calyx green to yellowish-green, campanulate, ca. 6 mm long, densely hispidulous and glandular outside; teeth 5, subequal, triangular, reflexed, ca. 2 mm long, sparsely hispidulous inside, apex acute. Corolla ca. 1.4 cm long; tube ca. 6 mm long, ca. 1.5 mm wide, purplish-red at upper part; 2-lipped, upper lip oblong, yellowish-green, erect, concave, ca. 8 mm long, ca. 6 mm wide, sparsely pubescent outside, apex emarginate, lower lip ca. 8 mm long, ca. 1 cm wide, sparsely pubescent outside, 3-lobed, medium lob largest, white, dotted with purplish-red spots, subcircular, concave, apex emarginate, ca. 6 mm long, ca. 6 mm wide, lateral lobes ovate, yellowish-green, dotted with purplish-red spots, reflexed, ca. 4 mm long, ca. 2 mm wide. Stamens 4, straight, included, filaments hispid at base, anther cells 2, divergent. Style included, glabrous, apex subequally 2-lobed, lobes subulate. Ovary truncate at apex, densely hispid and glandular. Nutlets yellowish-brown, triquetrous-oblong, ca. 3.5 mm long, apex hispid and glandular.


Flowering from June to July, fruiting from July to August.

Distribution and habitat

Paraphlomis hsiwenii is currently only known from Diding Natural Reserve in Guangxi, China (Fig. 5). It occurs in shady places in evergreen broad-leaved forests or among shrubs in limestone mountains at an altitude of ca. 1200 m.

Figure 5. 

Distribution of Paraphlomis hsiwenii (star) and P. pagantha (circle).


The new species is named after the Chinese taxonomist Hsi-Wen Li, who passed away in 2021 and had contributed tremendously to the taxonomy of Lamiaceae from China.

Chinese name

(assigned here). xī wén jiǎ cāo sū (锡文假糙苏).

Additional specimens examined

China. Guangxi: Jingxi City, Nanpo Town, Longdingtun, Diding Natural Reserve, alt. 1231 m, 19 Aug 2020, W.H. Wu et al. DD426 (KUN).

Specimens of P. pagantha examined

China. Hainan: Danzhou City, Shaposhan (Mt. Shamaoling), 29 Aug 1927, W.T. Tsang 672-16171 (IBSC0718446, IBSC0585106, PE00834801); Ledong County, Chang’e Village, Mt. Chang’eling, 16 Jun 1936, S.K. Lau 27154 (IBK00059945, IBSC0585103, KUN0274797, NAS00224460, PE00834800); Ledong County, Mt. Jianfengling, alt. 750 m, 16 Jun 1959, Z.F. Wei 122572 (IBSC0585104); Qionghai City, Huishan Natural Reserve, 21 Apr 2021, L.X. Yuan et al. s.n. (KUN); Wanning City, Xinglong Town, Langmingtian Village, Wutiaosang, 17 Jul 1935, F.C. How 73217 (IBK00059942, IBSC0585105, PE00834802); Wanning City, Xinglong Town, Mt. Niuguling, 16 Apr 1935, F.C. How 71953 (BM, IBK00059943, IBSC0585102); Wanning City, mountain behind Nanlin Nongchang, 5 May 1984, Z.X. Li et al. 1661 (IBSC0585107); Wenchang City, Tongguling Natural Reserve, 4 Aug 2021, L.X. Yuan et al. s.n. (KUN). VIETNAM. Tonkin (Hanoi): Mont. Bavi, Aug 1887, B. Balansa 2914 (Type: K000928198); Ba Vi, Son Tay, alt. 800 m, 14 Jun 1962, Ban 6893 (IBSC0616357).


We would like to thank the staff of the following herbaria for their kind help in research facilities: BM, CDBI, E, GXMI, HIB, IBK, IBSC, K, KUN, KYO, MW, NAS, PE, SZ, TI. Thanks are also given to Mr. Xiao-Lei Ma for his help in field collection, and to the reviewers for their valuable suggestions that improved our manuscript. This work was supported by the National Natural Science Foundation of China (Grant No. 31800168) and the Yunnan Fundamental Research Projects (Grant Nos. 202101AU070067 & 202101AT070159) to YPC. Thanks are also given to Mr. Xiao-Lei Ma for his help in field collection, and to the reviewers for their valuable suggestions that improved our manuscript.


  • Bendiksby M, Thorbek L, Scheen AC, Charlotte L, Olof R (2011) An updated phylogeny and classification of Lamiaceae subfamily Lamioideae. Taxon 60(2): 471–484.
  • Chen YP, Liu A, Yu XL, Xiang CL (2021) A preliminary phylogenetic study of Paraphlomis (Lamiaceae) based on molecular and morphological evidence. Plant Diversity 43(3): 206–215.
  • Chen YP, Sun ZP, Xiao JF, Yan KJ, Xiang CL (2022a) Paraphlomis longicalyx (Lamiaceae), a new species from the limestone area of Guangxi and Guizhou Provinces, southern China. Systematic Botany 47(1): 251–258.
  • Chen YP, Xiong C, Zhou HL, Chen F, Xiang CL (2022b) Paraphlomis nana (Lamiaceae), a new species from Chongqing, China. Turkish Journal of Botany 46(2): 176–182.
  • Doyle JJ, Doyle JD (1987) A rapid DNA isolation procedure for small quantities of fresh leaf tissue. Phytochemical Bulletin 19: 11–15.
  • Harley RM, Atkins S, Budantsev AL, Cantino PD, Conn BJ, Grayer R, Harley MM, de Kok R, Krestovskaja T, Morales R, Paton AJ, Ryding O, Upson T (2004) Labiatae. In: Kubitzki K, Kadereit JW (Eds) The families and genera of vascular plants, vol. 7. Springer, Berlin and Heidelberg, 167–275.
  • Li HW (1965) Revisio generis Paraphlomis Labiatarum Sinensium. Acta Phytotaxonomica Sinica 10: 57–74.
  • Li HW, Hedge IC (1994) Lamiaceae. In: Wu CY, Raven PH (Eds) Flora of China, vol. 17. Science Press, Beijing and Missouri Botanical Garden Press, St. Louis, 269–291.
  • Miller MA, Pfeiffer W, Schwartz T (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. Proceedings of the Gateway Computing Environments Workshop (GCE). New Orleans, LA, 1–8.
  • Ronquist F, Teslenko M, van der Mark P, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61(3): 539–542.
  • Suddee S, Paton A (2006) Validation of Lamiaceae names. Kew Bulletin 61: 619–621.
  • Thiers B (2022) Index Herbariorum: a global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium. [Accessed Jul 2022]
  • Wu CY, Li HW (1977) Paraphlomis Prain. In: Wu CY, Li HW (Eds) Flora Reipublicae Popularis Sinicae, vol. 65(2). Science Press, Beijing, 545–572.
  • Zhang RB, Deng T, Dou QL, Wei RX, He L, Ma CB, Zhao S, Hu S (2020) Paraphlomis kuankuoshuiensis (Lamiaceae), a new species from the limestone areas of northern Guizhou, China. PhytoKeys 139: 13–20.

Appendix 1

Table A1.

Sequence information for all samples used in present study. A “/” indicates a missing sequence. Herbarium abbreviations are listed after the vouchers. The accession numbers marked in bold represent sequences newly generated.

Taxon Voucher Country ITS ETS rpl32-trnL rps16 trnL-trnF
Matsumurella chinensis (Benth.) Bendiksby 1 Y. Yang OYY00316 (KUN) Pingxiang, Jiangxi, China MW602147 MW602117 MW602021 MW602053 MW602084
Matsumurella chinensis (Benth.) Bendiksby 2 Y. Yang OYY00131 (KUN) Guilin, Guangxi, China MW602148 MW602118 MW602022 MW602054 MW602085
Matsumurella yangsoensis (Y.Z. Sun) Bendiksby L. Wu & W.B. Xu 10965 (IBK) Yangshuo, Guangxi, China MW602142 MW602112 / / /
Paraphlomis albida Hand.-Mazz. var. albida A. Liu et al. LK0841 (CSFI) Ningyuan, Hunan, China MW602124 MW602091 MW601996 MW602028 MW602060
Paraphlomis albida var. brevidens Hand.-Mazz. Y.P. Chen EM312 (KUN) Hezhou, Guangxi, China MW602130 MW602098 MW602003 MW602035 MW602067
Paraphlomis albiflora (Hemsl.) Hand.-Mazz. C.M. Tan et al. 1806393 (JJF) Jiujiang, Jiangxi, China / MW602101 MW602006 MW602038 MW602069
Paraphlomis coronata (Vaniot) Y.P. Chen & C.L. Xiang 1 E.D. Liu et al. 3043 (KUN) Emeishan, Sichuan, China MW602137 MW602107 MW602012 MW602044 MW602075
Paraphlomis coronata (Vaniot) Y.P. Chen & C.L. Xiang 2 C.L. Xiang 358 (KUN) Jiangkou, Guizhou, China MW602123 MW602090 MW601995 MW602027 MW602059
Paraphlomis foliata (Dunn) C.Y. Wu & H.W. Li subsp. foliata S.P. Chen s.n. (KUN) Jiangle, Fujian, China / MW602097 MW602002 MW602034 MW602066
Paraphlomis foliata subsp. montigena X.H. Guo & S.B. Zhou Y.C. Dai s.n. (KUN) Hangzhou, Zhejiang, China OM836064 OM884453 OM884456 OM884459 OM884462
Paraphlomis gracilis (Hemsl.) Kudô var. gracilis 1 A. Liu LK0931 (CSFI) Changsha, Hunan, China MW602134 MW602104 MW602009 MW602041 MW602072
Paraphlomis gracilis (Hemsl.) Kudô var. gracilis 2 C.L. Xiang XCL1315 (KUN) Chongqing, China MW602141 MW602111 MW602016 MW602048 MW602079
Paraphlomis gracilis var. lutienensis (Y.Z. Sun) C.Y. Wu C.L. Xiang XCL881 (KUN) Shibing, Guizhou, China MW602131 MW602099 MW602004 MW602036 MW602068
Paraphlomis hispida C.Y. Wu X. Li LX200702 (GXF) Napo, Guangxi, China MW602132 MW602102 MW602007 MW602039 MW602070
Paraphlomis hsiwenii Y.P.Chen & XiongLi 1 W.H. Wu et al. DD426 (KUN) Jingxi, Guangxi, China OP605346 OP609841 OP609848 OP609855 OP609862
Paraphlomis hsiwenii Y.P.Chen & XiongLi 2 W.H. Wu et al. DD426 (KUN) Jingxi, Guangxi, China OP605347 OP609842 OP609849 OP609856 OP609863
Paraphlomis intermedia C.Y. Wu & H.W. Li X. Zhong et al. ZX16823 (CSH) Suichang, Zhejiang, China MW602135 MW602105 MW602010 MW602042 MW602073
Paraphlomis javanica (Blume) Prain var. javanica 1 Y.P. Chen s.n. (KUN) Kunming, Yunnan, China MW602121 MW602088 MW601993 MW602025 MW602057
Paraphlomis javanica (Blume) Prain var. javanica 2 L.B. Jia et al. JLB0029 (KUN) Maguan, Yunnan, China MW602143 MW602113 MW602017 MW602049 MW602080
Paraphlomis javanica var. pteropoda D. Fang & K.J. Yan X. Li 2020090501 (GXF) Jingxi, Guangxi, China MW602140 MW602110 MW602015 MW602047 MW602078
Paraphlomis jiangyongensis X.L. Yu & A. Liu 1 A. Liu et al. LK1104 (CSFI) Jiangyong, Hunan, China MW602128 MW602095 MW602000 MW602032 MW602064
Paraphlomis jiangyongensis X.L. Yu & A. Liu 2 A. Liu et al. LK1104 (CSFI) Jiangyong, Hunan, China MW602129 MW602096 MW602001 MW602033 MW602065
Paraphlomis kwangtungensis C.Y. Wu & H.W. Li Y.P. Chen & Y. Zhao EM1391 (KUN) Huaiji, Guangdong, China MW602126 MW602093 MW601998 MW602030 MW602062
Paraphlomis lanceolata Hand.-Mazz. 1 C.Z. Huang s.n. (KUN) Guidong, Hunan, China MW602145 MW602115 MW602019 MW602051 MW602082
Paraphlomis lanceolata Hand.-Mazz. 2 A. Liu et al. LK0825 (CSFI) Ningyuan, Hunan, China MW602146 MW602116 MW602020 MW602052 MW602083
Paraphlomis lancidentata Y.Z. Sun X. Zhong et al. ZX16824 (CSH) Suichang, Zhejiang, China MW602136 MW602106 MW602011 MW602043 MW602074
Paraphlomis longicalyx Y.P. Chen & C.L. Xiang Y.P. Chen et al. EM583 (KUN) Huanjiang, Guangxi, China OK104771 OK104774 OK104778 OK104780 OK104783
Paraphlomis membranacea C.Y. Wu & H.W. Li M.S. Nuraliev 1057 (MW) Thanh Son, Phu Tho, Vietnam / MW602100 MW602005 MW602037 /
Paraphlomis nana Y.P. Chen, C. Xiong & C.L. Xiang 1 C. Xiong XC21097 (KUN) Chengkou, Chongqing, China OM836062 OM884451 OM884454 OM884457 OM884460
Paraphlomis nana Y.P. Chen, C. Xiong & C.L. Xiang 2 C. Xiong & H.L. Zhou XC21126 (KUN) Wushan, Chongqing, China OM836063 OM884452 OM884455 OM884458 OM884461
Paraphlomis pagantha Dunn L.X. Yuan et al. s.n. (KUN) Qionghai, Hainan, China OP605345 OP609840 OP609847 OP609854 OP609861
Paraphlomis paucisetosa C.Y. Wu 1 X.X. Zhu s.n. (KUN) Malipo, Yunnan, China MW602125 MW602092 MW601997 MW602029 MW602061
Paraphlomis paucisetosa C.Y. Wu 2 X. Li LX200704 (GXF) Napo, Guangxi, China MW602133 MW602103 MW602008 MW602040 MW602071
Paraphlomis reflexa C.Y. Wu & H.W. Li Z.Z. Yang et al. s.n. (HIB) Tongshan, Hubei, China MW602122 MW602089 MW601994 MW602026 MW602058
Phlomis fruticosa L. Y. Tong s.n. (KUN) Shanghai, China (cultivated) MW602119 MW602086 MW601991 MW602023 MW602055
Phlomoides dentosa var. glabrescens (Danguy) C.L. Xiang & H. Peng Y.P. Chen EM360 (KUN) Beijing, China (cultivated) MW602120 MW602087 MW601992 MW602024 MW602056