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Taxonomic and nomenclatural notes on Pedicularis (Orobanchaceae): I. One new species from northwest Yunnan, China
expand article infoXin Li§, Hong Wang|, De-Zhu Li|, Wen-Bin Yu§
‡ University of Chinese Academy of Sciences, Beijing, China
§ Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Mengla, China
| Kunming Institute of Botany, Chinese Academy of Sciences, Kunming, China
¶ Southeast Asia Biodiversity Research Institute, Chinese Academy of Science, Yezin, China
Open Access

Abstract

Pedicularis multicaulis W.B.Yu, H.Wang & D.Z.Li (series Oliganthae Prain) is a new species described and illustrated herein. This new species is endemic to northwest Yunnan and only two populations were found in Weixi county. Phylogenetic analyses support P. multicaulis as a new species, sister to P. taihaiensis Bonati and P. macilenta Franch. Morphological comparisons between P. multicaulis and P. macilenta and P. taihaiensis also support P. multicaulis as a new species to science.

Keywords

Orobanchaceae, Pedicularis multicaulis, Mountains of Southwest China, phylogenetic analysis

Introduction

Pedicularis Linn., with around 600 species, is the largest genus of Orobanchaceae and widely distributed throughout the North temperate region (Fischer 2004, Stevens 2001, Yu et al. 2015). More than 350 species have been recognised in China (Yang et al. 1998). Of them, about two-thirds of the species are restricted in the Hengduan Mountains, which belongs to the Mountains of Southwest China hotspot (Wang 2006, Wang and Wu 1994). Due to the previously limited accessibility of the Mountains of Southwest China before the 21st century, several new species of Pedicularis have subsequently been discovered and described in the 2000s, owing to the construction of a road system under China’s Great Western Development Strategy (Liu and Yu 2015, Yang et al. 2003, Yu et al. 2010, Yu et al. 2018).

According to the phylogeny of the Pedicularis species with well-represented samples from the Hengduan Mountains region, 18 taxa were not categorised as any recognised species, based on both molecular and morphological data (Yu et al. 2015), which could be potential new species or new records to China. Of them, two taxa had been described as new species, P. wanghongiae M.L.Liu & W.B.Yu (Liu and Yu 2015) and P. millina W.B.Yu, D.Z.Li & H.Wang (Yu et al. 2018). In this study, we described and illustrated another new species, P. multicaulis W.B.Yu, H.Wang & D.Z.Li, from the remaining 16 taxa after carefully examining morphological characters and in comparisons with herbarium specimens of the close relatives, P. taihaiensis Bonati and P. macilenta Franch. (Yu et al. 2015). Pedicularis multicaulis is strongly supported as a new species, based on the revised phylogenetic analyses. Meanwhile, the pollen morphology of P. multicaulis was investigated using a scanning electron microscope (SEM).

Material and methods

The fresh specimens of the new species were collected from Pantiange and Lidiping in Weixi county, northwest Yunnan, China. Pollen samples were collected from the type specimens, then observed under SEM (ZEISS EVO LS10, Germany). For the morphological comparisons, we examined specimens or specimen images of the closest relatives from the herbaria E, K, KUN, LA, P and PH. Selected type specimens of P. macilenta and P. taihaiensis are presented in Suppl. material 1: Figures S1 and S2.

According to the published phylogeny of Pedicularis (Yu et al. 2015), P. multicaulis, P. macilenta and P. taihaiensis were chosen as ingroups and P. cephalantha Franch. ex Maxim. and other species from series Oliganthae Prain, Strobilaceae Tsoong and Amplitubae Li were also included (Table 1). Pedicularis axillaris Franch. ex Maxim. was specified as the outgroup. In this study, we had two samples of the new species from Pantiange (W.-B. Yu et al. 2014102) and Lidiping (W.-B. Yu et al. 2014096), respectively, two samples of P. tahaiensis from Luquan (C.-L. Xiang et al. HP9544) and Huize (W.-B. Yu et al. HW10369), respectively and one sample of P. macilenta. Four DNA regions (nrITS, matK, rbcL and trnL-F) were used and the new sequences generated following Yu et al. (2011). Bayesian Inference (BI), Maximum Likelihood (ML) and Maximum Parsimony (MP) methods were used to reconstruct the phylogenies. The BI analysis was performed using MrBayes 3.26 (Ronquist and Huelsenbeck 2003). The total dataset was partitioned (see Suppl. material 2: Dataset 1) and the DNA substitution model of Bayesian Information Criterion (BIC) for four DNA regions was estimated using jModeltest 2 (Darriba et al. 2012). The ML analysis was conducted with RAxML 8.2.10 (Stamatakis et al. 2008). The MP analysis was carried out using PAUP* 4.a165 (Swofford 2003). Parameters for the three analyses followed the previous studies (Yu et al. 2013, Yu et al. 2015).

Voucher information and GenBank accessions of samples used in phylogenetic analyses.

Taxon Source Voucher information ITS matK rbcL trnL-F
P. amplituba H.L. Li Yunnan: Luquan Yu et al., LIDZ1519A (KUN) JF977469 JF955063 JF942952 KF277605
P. axillaris Franch. ex Maxim. (3) Yunnan: Dali Yu et al., YWB2014097 (KUN) KT022428 KT022531 KT022705 KT022883
P. cephalantha Franch. ex Maxim. Yunnan: Lijiang W. Jiang, 08727 (KUN) JF977493 JF955087 JF942976 KF277613
P. cephalantha affinis Yunnan: Eryuan Yu et al., YWB2014063 KT022501 KT022661 KT022841 KT022967
P. dissectifolia H.L. Li Yunnan: Shangeri-La Yu et al., HW10133 (KUN) KF277539 KR707763 KF277641 KF277641
P. fengii H.L. Li (1) Yunnan: Shangeri-La Yu et al., HW10102 (KUN) JF977553 JF955146 JF943036 KT022910
P. fengii H.L. Li (2) Yunnan: Shangeri-La Yu et al., Yu606 (KUN) JF977564 JF955157 JF943047 KF277646
P. gracilicaulis H.L. Li Xizang: Chayu Jin et al., STET0522 (PE) KF277547 no data no data KF277654
P. macilenta Franch. ex Forbes ex Hemsl. Yunnan: Zhaotong Li et al., 8484 (KUN) KF277558 KT022606 KT022780 KF277680
P. multicaulis W.B.Yu, H.Wang & D.Z.Li Yunnan: Weixi Yu et al., YWB2014096 KT022502 KT022662 KT022842 KT022968
P. multicaulis W.B.Yu, H.Wang & D.Z.Li Yunnan: Weixi Yu et al., YWB2014102 MK983380 MK983381 MK983382 MK983383
P. strobilacea Franch. Yunnan: Shangeri-La Cai et al., 11CS3261 (KUN) KT022508 KT022673 KT022852 KT022977
P. pseudocephalantha Franch. Xizang: Linzhi Gao et al., GLM123906 (KUN) KR707794 KR707760 KR707780 KR707807
P. tachanensis Bonati Sichuan: Mianning Yu et al., LIDZ1062 (KUN) JF977743 JF955333 JF943226 KF277740
P. tahaiensis Bonati Yunnan: Luquan Xiang et al., HP9544 (KUN) JF977552 JF955145 JF943035 KF277741
P. tahaiensis Bonati Yunnan: Huize Yu et al., HW10369 (KUN) JF977563 JF955156 JF943046 no data

The conservation status of P. multicaulis was assessed in accordance with IUCN Red List Criteria (IUCN 2012).

Taxonomy

Pedicularis multicaulis W.B.Yu, H.Wang & D.Z.Li, sp. nov.

Figures 1, 2A–F and 3

Vernacular name

Duo Jing Ma Xian Hao (多茎马先蒿) (Chinese).

Type

CHINA. Yunnan: Weixi, Lidiping, wet meadow, alt. 3180 m, 27°9'16.06"N, 99°24'48.70"E, 30 Aug 2014, W.-B. Yu, X.-L. Yang & H. Tang 2014096 (holotype: HITBC! (accession no. 169315); isotypes: HITBC!, KUN!).

Diagnosis

Pedicularis multicaulis W.B.Yu, H.Wang & D.Z.Li is distinguished from P. macilenta and P. taihaiensis in having taller and more ascending stems, partially crawling stems with fibrous roots, shorter petiole and leaf blade of cauline leaves in middle and upper parts and smaller corollas with a shorter beak.

Description

Herbs perennial, 20–50 cm tall, glabrescent, drying slightly black; taproots slender, fusiform; stems caespitose, mostly (3) 5 to 9 (12) from a caudex, ascending or partially crawling (with fibrous roots) and branchlets (0) 1–3 (10), glabrescent or sparely pubescent along the lines. Basal leaves absent. Cauline leaves alternate; petiole up to 10 mm long or distal ones sessile or subsessile, glabrescent; leaf blade ovate-elliptic or oblong, 5–30 mm × 7–15 mm, glabrous on both surfaces, pinnatisect; segments 2 to 5 pairs, ovate to lanceolate-oblong, incised-pinnatifid or double dentate. Inflorescences racemose, up to 30 cm long; bracts leaflike, distal ones shorter than flowers. Pedicel 1.0–2.5 mm long. Calyx tube ca. 5 mm long, glabrescent, 1/3 cleft anteriorly; lobes 3, unequal, posterior one acicular, lateral pair larger, leaf-like and toothed. Corolla rose, 10–14 mm long; tube erect, ca. 8–10 mm long ; galea ±falcate, not crested, not twisted, with 1 distinct reflexed marginal tooth on one side; beak straight, ca. 3 mm, slightly 2-cleft at apex, not ciliate; lower lip 5–6 mm × 6–8 mm, sparely ciliate, lobes 3 unequal; middle lobes apex slightly cucullate. Filaments 4 glabrous, equal length, ca.13 mm long, inserted in the middle of corolla. Ovary long ovoid, ca. 3 mm long; Capsule lanceolate-oblong, 10–15 mm × 4–5 mm. Seeds narrowly ovoid, ca. 1.0–1.2 mm.

Figure 1. 

Line drawing of Pedicularis multicaulis W.B.Yu, H.Wang & D.Z.Li A habit B leaf C flower D open flower showing the anthers and style E fruit. Drawn by Zhen-Long Liang from the holotype (A–D) and an isotype (E), W.-B. Yu, X.-L. Yang & H. Tang 2014096 (KUN).

Etymology

The specific epithet “multicaulis” refers to the new species having many ascending stems that are branched in the middle and upper parts.

Phenology

This new species was found in flowering from middle June (in a field trip in 2006) to August and in fruiting from July to September.

Pollen morphology

Pollen grains are radially symmetrical, isopolar, spheroidal and medium in size (polar length: 23.71–25.47 μm × equatorial diameter: 18.86–20.29 μm). Pollen apertures are bisyncolpate (Figures 2G and H) and the colpi are usually wide and sunken (Figure 2G); exine ornamentation is perforated tectum with microfoveolate ornamentation (Figure 2I).

Figure 2. 

Field photos and pollen of Pedicularis multicaulis W.B.Yu, H.Wang & D.Z.Li A–C overview of habitat and plants D inflorescence E flowers F infructescence G equatorial view of pollen H polar view of pollen I exine ornamentation.

Figure 3. 

The holotype of Pedicularis multicaulis W.B.Yu, H.Wang & D.Z.Li (W.-B. Yu, X.-L. Yang & H. Tang 2014096, HITBC, accession no. 169315).

Phylogenetic analyses

All analyses strongly supported P. taihaiensis as sister to P. multicaulis (ML/MP/BI = 88/76/1.00, Figure 4) and the two samples of P. multicaulis (ML/MP/BI = 100/100/1.00) and of P. taihaiensis (ML/MP/BI = 100/99/1.00) are monophyletic, respectively. Then, P. macilenta is sister to P. taihaiensis + P. multicaulis (ML/MP/BI = 100/100/1.00).

Figure 4. 

The major-rule consensus tree of Bayesian Inference analysis using the total data by concatenating four DNA regions (nrITS, matK, rbcL and trnL-F). Bootstrap values of Maximum Likelihood/Parsimony and posterior probability values of Bayesian Inference are presented above branches. The bottom scale bar represents the number of substitutions per site.

Distribution

Pedicularis multicaulis was only found in two populations in Weixi county, northwest Yunnan (Figure 5). It occurs in wet meadow or the margin of wetland between 2900 m and 3200 m a.s.l.

Figure 5. 

Distribution map of Pedicularis multicaulis W.B.Yu, H.Wang & D.Z.Li and related taxa.

Conservation assessment

To date, we only collected this new species from two populations in Weixi county, northwest Yunnan. There are around 100 and 300 individuals in Pantiange and in Lidiping, respectively. It is restricted to wet meadow, which is likely to be threatened by grazing in these areas. According to IUCN Red List Criteria (IUCN 2012), P. multicaulis can be classified as Vulnerable (VU).

Additional examined specimens

Pedicularis multicaulis W.B.Yu, H.Wang & D.Z.Li. CHINA. Yunnan: Weixi, Pantiange, wet grassland, alt. 2930 m, 27°20'39.48"N, 99°16'59.30"E, 27 Aug 2014, W.-B.Yu, X.-L.Yang & H.Tang 2014102 (KUN!). Pedicularis macilenta Franch. CHINA. Yunnan: Eryuan (Mountain Yentzehay), in humid localities on the slopes, 8 Aug 1888, Delavay 3698 (types, P!, PH!, LA!); Yunnan: Zhaotong, Dashanbao, Dahaizi reservoir, alt. 3044 m, 27°44'89.2"N, 103°31'94"E, 7 Aug 2008, H.Li et al. 8078 (KUN!). Pedicularis taihaiensis Bonati. CHINA. Yunnan: Huize, Dahai, Jul 1913, E.E. Maire 678 (holotype: E [E00284020]!); ibid. 30 Jul 2010, W.-B.Yu et al. HW10369 (KUN!); Yunnan: Luquan, Wumeng Mountains, alt. 3700 m, 2 Jul 1990, R.Z.Fan & Z.W.Lyv 061 (KUN!); Yunnan: Luquan, Jiaozi Mountain. 8 Jul 2008, C.L.Xiang et al. HP9544 (KUN!).

Discussion

The galea of P. multicaulis bears one pair of distinct reflexed marginal teeth on both sides, which is the key character of series Oliganthae Prain. Phylogenetic analyses did not support series Oliganthae as monophyletic (Yu et al. 2015). The previous study indicated that P. macilenta and P. taihaiensis formed a weakly supported clade, then sister to P. multicaulis (= Pedicularis sp. (9)) by using one sample of each species. In this study, both P. multicaulis and P. taihaiensis had two samples from different populations and our results showed that P. multicaulis and P. taihaiensis formed a strongly supported clade, then sister to P. macilenta. The relationship amongst the three species was well resolved. Therefore, population level sampling is very important for species delimitation and phylogeny of recently derived lineage.

Morphological characters differentiate P. multicaulis from the two most closely related species (Table 2). The key diagnostic characters of P. multicaulis are having taller and more branched stems, partially crawling stems with fibrous roots, shorter petiole of cauline leaves and smaller corollas with a short beak. The three species are also isolated geographically (Figure 5). According to herbarium records, P. taihaiensis occurs in Luquan and Huize, north Yunnan and P. multicaulis is only found in Weixi, northwest Yunnan. The distribution of P. macilenta is disjunct, with one population in Eryuan, northwest Yunnan and another in Zhaotong, northeast Yunnan. As all three species were mainly confined to the habitat of wet meadow, we assume that geographical isolation may play an important role in species divergence in this lineage.

Morphological comparison amongst Pedicularis multicaulis, P. macilenta and P. taihaiensis.

Characters P. multicaulis P. macilenta P. taihaiensis
Plant height (cm) 20–50 20–30 15–30
Rooting stems Yes No No
Stems (3) 5-9 (12) 1–5 2–4
Branchlets per stem (0) 1-3 (10) 1–3 1–3
Leaf bade size (mm) 5–20 × 7–15 30–50 × 10–15 15–30 × 8–11
Petiole length (mm) 3–11 5–20 8–25
Leaf lobes (pairs) 2–5 5–7 5–7
Leaf lobe size (mm) 3–8 × 2–4 3–7 × 2–5 3–6 × 1–4
Calyx length (mm) 4–5 6–7 5–7
Corolla colour Rose White with purple beak Rose
Corolla length (mm) 10–14 11–13 17–20
Corolla tube length (mm) 8–10 6–7 11–15
Beak length (mm) 3 3–4 4–5
Galea Not crested Slightly crested Not crested

Acknowledgements

We are grateful to Hui Tang and Xiu-Long Yang for their kind help in the field; to Zhen-Long Liang for the line drawing; to Ting Tang and Yin Zhao for preparing SEM investigation; and to the curators of the herbaria K, KUN, LA, P and PH for making specimens available for access in the herbarium or online database. This study was supported by grants from the Large-scale Scientific Facilities of the Chinese Academy of Sciences (2017-LSFGBOWS-02), Chinese Academy of Sciences Strategic Priority Research Program of the Chinese Academy of Sciences (XDB31000000) and the National Natural Science Foundation China (31470323, 31870196).

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