CC BY
39
Turczaninowia 24, 3: 175-193 (2021) DOI: 10.14258/turczaninowia.24.3.14 http://turczaninowia.asu.ru
I ISSN 1560-7259 (print edition)
TURCZANINOWIA
I ISSN 1560-7267 (online edition)
УДК 582.382:581.45+57.065 A taxonomic revision of Selaginella monospora Spring (Selaginellaceae)
A. P. Shalimov1, 2, 3, X.-C. Zhang1, 4*
1 State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, Beijing, 100093, China
2 University of Chinese Academy of Sciences, Beijing, 100049, China
3South Siberian Botanical Garden, Altai State University, Leninapr., 61, Barnaul, 656049, Russian Federation.
ORCID iD: https://orcid.org/0000-0002-8806-5217
4E-mail: zhangxc@ibcas.ac.cn; ORCID iD: https://orcid.org/0000-0003-3425-1011
*Corresponding author
Keywords: China, India, lectotypification, new synonyms, phylogenetic analysis, Selaginella, taxonomy.
Summary. During a taxonomic revision of Selaginella from the Himalaya which is a part of the ongoing "Flora of Pan-Himalaya" project, we studied the taxonomy of Selaginella monospora and its closely related species. We found that S. microclada once treated as a synonym of S. monospora should be accepted as a distinct species. In contrast, S. trichophylla, S. monospora var. ciliolata, S. medogensis, and S. effusa var. dulongjiangensis are reduced to synonymy of S. monospora based on molecular phylogenetic analyses and morphological characters. Selaginella monospora is characterized by denticulate to ciliolate leaves and strongly dimorphic sporophylls vs. denticulate leaves and slightly dimorphic sporophylls in S. microclada. A lectotypification of S. medogensis was also made in the present study.
Таксономическая ревизия Selaginella monospora Spring (Selaginellaceae)
А. П. Шалимов1, 2, 3, С.-Ч. Жан1
1 Лаборатория систематики и эволюционной ботаники, Институт ботаники Китайской Академии Наук (КАНН), г. Пекин, 100093, Китай
2 Университет Академии наук Китая, г. Пекин, 100049, Китай
3Южно-Сибирский ботанический сад, Алтайский государственный университет, просп. Ленина, 61, г. Барнаул, 656049, Россия
Ключевые слова: Китай, Индия, лектотипификация, новые синонимы, таксономия, филогенетический анализ, Selaginella.
Аннотация. В процессе таксономической ревизии рода Selaginella Гималаев, которая является частью продолжающегося проекта «Флора Пан-Гималаев», мы обнаружили, что после недавней типификации традиционное понимание таксона S. monospora изменилось, что потребовало пересмотра номенклатуры данного вида и его родства. Selaginella microclada, традиционно принимавшаяся как синоним S. monospora, должна быть признана отдельным видом, а S. trichophylla, S. monospora var. ciliolata, S. medogensis и S. effusa var. dulongjiangensis сводятся в синонимы к S. monospora на основе изучения гербарных коллекций, в том числе типовых, и молекулярно-филогенетического анализа рассматриваемой группы. Selaginella monospora ха-
Поступило в редакцию 18.08.2021 Принято к публикации 10.09.2021
Submitted 18.08.2021 Accepted 10.09.2021
рактеризуется зубчатыми или реснитчатыми листьями и сильно диморфными спорофиллами, в отличие от X ткгос^а, которой свойственны зубчатые листья и слегка диморфные спорофиллы. В настоящем исследовании также обнародуется лектотипификация X medogensis.
INTRODUCTION
Selaginella P. Beauv. is the only genus of family Selaginellaceae comprising approximately 700-800 species (Jermy, 1990; Zhang, 2004; Zhang et al., 2013; Weststrand, Korall, 2016). The taxonomy of some species groups in this genus have remained unresolved although the infrageneric classification has been the subject of several recent studies (Zhou et al. 2015a; Weststrand, Korall, 2016; Zhang et al. 2020). Selaginella monospora Spring is widely distributed from Indo-Himalaya to China, Myanmar, Laos, and Vietnam (Alston, 1934, 1945; Chun et al., 1964; Panigrahi, Dixit, 1966; Iwatsuki, 1975, 1988; Tagawa, Iwatsuki, 1979; Ching, Wu, 1983; Dixit, 1992; Thapa, 2002; Zhang, 2004; Ghosh et al., 2004; Singh, Panigrahi, 2005; Chu, 2006; Wu, 2006; Jiang, 2013; Zhang et al., 2013; Fraser-Jenkins et al., 2015; Fraser-Jenkins et al., 2017; Shalimov et al., 2019a). It is rather a common species in Southwest China, and similar plants were described from Xizang and Yunnan as S. medogensis Ching et S. K. Wu (1983), S. trichophylla K. H. Shing (1993), S. monospora var. ciliolata W. M. Chu (2006), and S. effusa var. dulongjiangensis W. M. Chu (2006). However, the taxonomic status of these taxa was uncertain (Zhang et al., 2013).
Herein, we use molecular and morphological data to draw further taxonomic conclusions and to stimulate and direct future taxonomic work in S. monospora group.
Material and Methods Molecular study
We sampled 57 individuals representing 35 species of Selaginella including the type specimens of the S. medogensis, S. trichophylla, S. monospora var. ciliolata, and S. effusa var. dulongjiangensis from GenBank and our data. Outgroups were chosen from different evolution lineages in Selaginella (S. nipponica, S. helvetica, S. laxistrobila, S. braunii, S. uncinata, S. delicatula, S. krausiana, S. remotifolia, S. deflexa, S. selaginoides) (Appendix 1).
Phylogenetic reconstruction was based on the chloroplast genes rbcL, atpI, psbA. Total genomic
DNA was extracted from silica-gel-dried leaf materials following the protocols and the primers and procedures described in Shalimov et al. (2019b). All sampling information is provided in Appendix 1. The raw sequences of Selaginella were assembled with ContigExpress. The alignments produced by Clustal X v.1.83 (Thompson et al., 1997) followed by manual adjustment in BioEdit v.7.1.11 (Hall, 1999), and gaps were treated as missing data.
Phylogenetic trees were constructed using maximum likelihood (ML) and Bayesian inference (BI). Models of nucleotide substitution were selected by jModelTest 0.1.1 (Posada, 2008). The ML analysis was run in RAxML 7.2.6 (Stamatakis, 2006) with 1000 bootstrap replicates under the GTRGAMMA model. Bayesian inference using the GTR+I+G model (four gamma categories) was executed in MrBayes version 3.1.2 (Huelsenbeck, Ronquist, 2001). Bayesian trees were started from random trees, and four Markov chain Monte Carlo (MCMC) simulations were run simultaneously and sampled every 1000 generations for 10 million generations. The average standard deviation of split frequencies (< 0.01) was used to assess the convergence of the two runs. Bayesian posterior probabilities (PP) were calculated as the majority consensus of all sampled trees with the first 25 % discarded as burn-in.
Morphological study
This revision is based on numerous specimens deposited at CDBI, CSH, E, GXMG, KUN, KYO, L, NY, PE, PYU, TI, U, and US. Type specimens and general collections from other herbaria (B, K, KYO, L, P, and US) were examined through digital images. We also used the JSTOR Global Plants project database (https://plants.jstor.org) and GBIF Global Biodiversity Information Facility (https:// www.gbif.org). Herbarium abbreviations are according to Thiers (2021).
Morphological characters, such as ventral (lateral), dorsal (median) and axillary leaves, strobili and arrangement of branches were carefully observed. The morpho-photographs of the plants were taken with a Nikon DXM 1200F camera connected to a stereomicroscope (Nikon SMZ 1000) and computer, and measurement were done by
D 3.10. Morphological features for included species were illustrated with axillary, dorsal, ventral leaves and strobili.
For the study of spore morphology, scanning electron microscopy (SEM) was used. Morphology of spores was examined from several collections for S. microclada, S. monospora, S. trichophylla, and S. monospora var. ciliolata. The spores were taken from mature sporangia and fixed on double line tape, and then covered with gold-palladium mixture. Spores were photographed and measured under different magnifications using a Hitachi S-4800 at 10-20 kV. The terminology used to describe the species is based on Tryon and Lugardon (1991) and Zhou et al. (2015b). For the black background,
contrast and brightness of the images were optimized in Adobe Photoshop.
Results
Interpretation of the lectotype of S. monospora designated by Fraser-Jenkins et al. (2017)
S. monospora was described by Spring (1850) based on several collections from India, Nepal, and Bhutan. We carefully checked the syntypes of S. monospora and specimens from the whole distribution area and realized that the syntypes represented two distinct species (Table 1-2).
Table 1
Taxonomic identity of the syntypes of Selaginella monospora
Herbarium barcode Collection information Taxa
B 20 0141434 India: Assam, Griffith s. n. species A
B 20 0141435 India: Assam, Griffith s. n. species A, with misidentification of upper left plants as S. semicordata
B 20 0141541 India: Assam, Griffith s. n. species A
K001067477 India: Assam, Griffith s. n. species A
K001067478 Bhutan ('Bootan'), Griffith W. 391 species B Lectotype of S. monospora chosen by Fraser-Jenkins et al. (2017)
K001067480 Nepal, Buchanan-Hamilton F. s. n. 1810 species A
K001067481 India, J. Mack 2 species A
K001067482 [Myrung] wood, Khasya, Griffith W. s. n. mixed gathering species A species B
K001067483 India, J. Mack 2 species A
K001067484 India: Khasiya, Griffith W. s. n. species B
P00523054 India, Assam, Griffith s. n. species B
US 00134380 Bootan, Griffith W. 391 mixed gathering species B species A
US 01392970 India: Khasiya, Griffith s. n. species B
K000009111 India: Khasiya, Griffith s. n. species B
Table 2
Morphological comparison between the taxa
Characteristics Species A Species B
Axillary leaves ovate, narrowly ovate, or narrowly elliptic, 2-3 x 0.8-1.6 mm, margin denticulate. ovate, 1.2-2.5 x 1-2 mm, margin ciliolate
Dorsal leaves ovate-lanceolate or elliptic, 1-1.6 x 0.3-0.7 mm, base obtuse, margin denticulate, acuminate or shortly aristate at apex ovate, 1.2-1.9 x 0.5-1 mm, base subcordate or obtuse, margin ciliolate, aristate to cuspidate at apex
Ventral leaves ovate-triangular or oblong-falcate, 2.6-5.5 x 0.92.3 mm, margin subentire, entire or denticulate ovate-triangular, 2-3.2 x 0.8-1.6 mm, margin denticulate to ciliolate
Strobili 3-20 x 1.9-3.2 mm, margin minutely denticulate 6.0-10 x 1.2-2.3 mm, margin denticulate
Sporophylls slightly dimorphic strongly dimorphic
The Species A is characterized by dorsal leaves ovate-lanceolate or elliptic, base obtuse, margin denticulate, apex acuminate or shortly aristate; strobili dorsiventrally complanate, sporophylls slightly dimorphic; lateral branches densely arranged, ventral leaves slightly ascending or spreading, ovate-triangular or oblong-falcate, margin denticulate, apex subacute. The Species B is characterized by dorsal leaves ovate, base obtuse or subcordate, margin ciliolate to denticulate, apex aristate to cuspidate; strobili dorsiventrally complanate, sporophylls strongly dimorphic; lateral branches flabellate arranged, ventral leaves ovate-triangular, oblong or oblong-ovate, margin ciliolate to denticulate, apex acute.
Fraser-Jenkins et al. (2017) lectotypified S. monospora Spring based on one of the original specimens cited in Spring's protologue: "Bhutan, W. Griffith, Herbarium Hookerianum", i. e. W. Griffith 391 in K (K001067478). This specimen belongs to the above Species B, which is characterized by strongly dimorphic strobili, flabellate lateral branches; dorsal leaves with denticulate to ciliolate at margin, aristate to cuspidate at apex; ventral leaves denticulate to ciliolate at acroscopic base.
Spring (1850) described S. gorvalensis at the same time with S. monospora. The type was cited as "Hindustania superiori, Gorval, Griffith" (Spring, 1850). Alston (1945) noticed that this collection was in fact from Khasia, and reduced it to S. monospora.
We found that the morphology of S. gorvalensis matched the above Species B in the syntypes of
S. monospora, and S. trichophylla (Shing, 1993) as well as S. monospora var. ciliolata (Chu, 2006) also belong to the Species B. The remaining part of the original syntypes of S. monospora have the same morphology characters (Species A) and are identical to the morphology of S. microclada Baker which was regarded as a synonym of S. monospora until now. Therefore, the resurrection of S. microclada is a consequence of the lectotypification of S. monospora by Fraser-Jenkins et al. (2017).
Morphology
The traditional concept of S. monospora s. l. is in fact a mixture of two different species, i. e., S. microclada and S. monospora. Selaginella microclada (species A) is characterized by slightly dimorphic sporophyll and thus the strobili appear more or less uniform; ventral and dorsal leaves denticulate; dorsal leaves oval-lanceolate or elliptic, base obtuse, apex acuminate or shortly aristate (Fig. 1, 1A-D). While, Selaginella monospora (species B) is characterized by strongly dimorphic sporophylls; ventral and dorsal leaves ciliolate to denticulate; dorsal leaves ovate, base subcordate or obtuse, apex cuspidate-aristate (Fig. 1, 2A-D).
Selaginella trichophylla, and S. monospora var. ciliolata are two similar eco-types of S. monospora with short spines on the dorsal and ventral leaves (Table 3).
Species/ Characteristics S. microclada S. monospora (incl. S. gorvalensis, S. trichophylla, S. monospora var. ciliolata) S. effusa var. dulongjiangensis S. medogensis
Main stems long creeping, 3585 cm or more creeping, 20-50 cm creeping, ca. 20-25 cm creeping, ca. 25 cm
Axillary leaves ovate, narrowly ovate, or narrowly elliptic, 2-3 x 0.81.6 mm, margin denticulate ovate, 1.2-2.5 x 1-2 mm, margin ciliolate broadly elliptic or ovate-elliptic, 1.12.0 x 0.7-1.7 mm, margin denticulate ovate-triangular or narrowly elliptic, 1.2-2.3 x 0.61.1 mm, margin denticulate
Base of axillary leaves base exauriculate base exauriculate base obtuse base exauriculate
Dorsal leaves ovate-lanceolate or elliptic, 1-1.6 x 0.30.7 mm, carinate or strongly carinate, margin denticulate ovate, 1.2-1.9 x 0.5-1 mm, not carinate, margin ciliolate (more densely ciliolate at base) obliquely ovate-elliptic 0.5-1.4 x 0.30.8 mm, margin denticulate ovate, 0.9-1.6 x 0.4-0.7 mm, carinate or strongly carinate, margin denticulate
Table 3
Morphological comparisons among Selaginella microclada, S. monospora (incl. S. gorvalensis, S. trichophylla, S. monospora var. ciliolata), S. effusa var. dulongjiangensis, and S. medogensis
Table 3 (end)
Species/ Characteristics S. microclada S. monospora (incl. S. gorvalensis, S. trichophylla, S. monospora var. ciliolata) S. effusa var. dulongjiangensis S. medogensis
Base of dorsal leaves obtuse, not peltate, obtuse or subcordate, not peltate obtuse oblique, not peltate
Apex of dorsal leaf acuminate or shortly aristate aristate to cuspidate shortly aristate caudate
Ventral leaves ovate-triangular or oblong-falcate, 2.65.5 x 0.9-2.3 mm, margin subentire or entire or denticulate; apex subacute ovate-triangular, 2-3.2 x 0.8-1.6 mm, margin denticulate to ciliolate, apex acute oblong or oblong-ovate 1.7-3.0 x 0.9-1.7 mm, margin denticulate, apex obtuse oblong or oblong-ovate, 1.6-3.5 x 1-2.0 mm, margin denticulate, apex subacute
Acroscopic base of ventral leaf enlarged, broader, strongly overlapping stem and branches, enlarged, broader, overlapping stem and branches, enlarged, broader, strongly overlapping stem and branches, margin denticulate at base in middle to apex subentire acroscopic base enlarged, broader, strongly overlapping stem and branches, margin denticulate at base (denticulate or subentire to apex)
Basiscopic base of ventral leaf margin subentire or entire margin entire margin entire margin denticulate at base (elsewhere denticulate or subentire)
Strobili 3-20 x 1.9-3.2 mm, margin minutely denticulate 6.5-10 x 1.2-2.3 mm, margin denticulate 4.0-7.0 x 1.11.7 mm, margin denticulate 3.4-4.8 x 1.82.2 mm, margin denticulate
Sporophylls slightly dimorphic dimorphic dimorphic dimorphic
Selaginella medogensis from Xizang is characterized by lateral leaves base broader, apex acute, margin denticulate; dorsal leaves narrow ovate or ovate, not carinate, short acuminate at apex, margin denticulate; dorsal sporophylls linear-lanceolate, ventral sporophylls triangle-lanceolate (Fig. 1, 3A-D).
Selaginella effusa var. dulongjiangensis is similar to S. effusa and S. medogensis, however, the main stems of S. effusa var. dulongjiangensis are creeping with spaced short flabellate branches (Fig. 1, 4A-D).
Micromorphology of spores
Our analysis of the morphology of spores also confirmed the micromorphological differences of mega- and microspores between S. microclada (in previous study considered as S. monospora) and S. monospora. The proximal and distal surfaces of
megaspores of S. microclada covered with verrucae or papillate ornamentation (Fig. 2A, C), sometimes with vermiculate structures, whereas S. monospora are covered by verrucae or papillate ornamentation on proximal and distal sides (Fig. 2E, G). The surface of megaspores varied from smooth with rare vermiculation (in present study) to more vermiculate without spinulose micro-sculptures (see in Zhou et al., 2015b for S. trichophylla), or surface with vermiculate microstructure and covered by rarely spinulose micro-sculptures (see in Zhou et al., 2015b for S. monospora var. ciliolata). Our observation of S. monospora showed verrucate ornamentation with vermiculate surface, which covered by spinulose micro-sculptures (Fig. 2E, G).
The morphology of spores of S. medogensis was reported by Liu et al. (2005) for S. mutensis (illegal name for S. medogensis) and Zhou et al. (2015b). As report by Liu et al. (2005), its megaspores have irregular verrucate ornamentation and
spinulose micro-sculptures on both surfaces. Zhou et al. (2015b) mentioned verrucate or pappilate ornamentation, but did not report vermiculate
structures of the surface (see Zhou et al., 2015b) and spinulose micro-sculptures.
2A
I - .vi-, -fc k jjjy; JätJfSp1 |r
- r.. 1 -
1.5 mm
1C
ygbk
s 3 3
3A
jffifki.
ApHk
JSt?- '.'•'■ t'-^v..- •
r ' .1»
't
3
3
3B
V
•
V j
s 1Jr
i
fei-
Fig. 1. Differences in vegetative leaves and strobilus: 1A-D - S. microclada (Feng 11473, PE); 2A-D - S. monospora (from paratype of S. monospora var. ciliolata (Lu et Zhang 27625-B, PE)); 3A-D - S. medogensis (Qinghai-Xizang Complex Exped. 74-4375, PE); 4A-D - S. effusa var. dulongjiangensis (W. M. Chu et Z. R. He 31299, PE). A - Axillary leaves; B - Dorsal leaves; C - Ventral leaves; D - Strobila.
Megaspores of S. effusa var. dulongjiangensis were reported by Zhou et al. (2015b), the surface had verrucae and was sometimes with vermiculate ornamentation, micro-sculptures were usually covered with dense spinules.
Microspores of S. microclada on proximal and distal sides had verrucae or spherulate ornamentation, surface covered with spinulose micro-sculptures (Fig. 2B, D). Our results considered previous report of Zhou et al. (2015b) for "S. monospora".
Microspores of S. monospora was previously reported for S. trichophylla and S. monospora var. ciliolata by Zhou et al. (2015b). They have verrucate or spherulate ornamentation, that is the same with our result, with fine reticulate micro-sculptures (Fig. 2F, H).
Microspores of S. medogensis were reported by Liu et al. (2005) and Zhou et al. (2015b), surface had verrucate or spherulate ornamentation, but Liu et al. (2005) additionally reported fine reticulate micro-sculptures.
As reported by Zhou et al. (2015b), microspores of S. effusa var. dulongjiangensis are covered with irregularly sized and spaced verrucae, the surface with dense spinules micro-sculptures. Summary on the difference between mega- and microspores of Selaginella microclada and S. monospora (incl. S. gorvalensis, S. trichophylla, and S. monospora var. ciliolata) with additional data on S. medogensis and S. effusa var. dulongjiangensis are presented in Table 4.
Table 4
Morphological comparison of mega- and microspores Selaginella microclada, S. monospora (incl. S. trichophylla, S. monospora var. ciliolata), S. medogensis, and S. effusa var. dulongjiangensis
S. microclada* S. monospora (incl. S. trichophylla*, S. monospora var. ciliolata*) S. effusa var. dulongjiangensis* S. medogensis* **
Megaspores the proximal and distal surfaces
verrucate and/or papillate verrucate and/or papillate verrucate verrucate or irregular sizes verrucae
Megaspores micro-sculptures
irregular rugae, spinulose irregular rugae, spinulose spinulose microsculptures spinulose microsculptures
Microspores
Microspores the proximal surfaces
verrucate or/and spherulate, with reticulate microsculptures verrucate or/and spherulate irregularly sized and spaced verrucae verrucate or/and spherulate
Microspores the distal surface
spherulate ornamentation, with reticulate microsculptures spherulate ornamentation, fine reticulate micro-sculptures dense spinules spherulate ornamentation, fine reticulate microsculptures
Note: * More detail and photo see in Zhou et al. (2015b); ** More detail and photo see in Liu et al. (2005).
Phylogenetic Analysis
The combined plastid dataset (rbcL + atpI+psbA) data matrix contains 57 accessions representing 35 Selaginella species. The alignment has 2040 characters, 382 (382/2041 = 18.71 %) of which are potentially parsimony informative. The ML tree with PP and ML bootstrap values (BS) is shown in Fig. 3. Phylogenetic analysis result showed that S. microclada (included 6 samples, clade A) is sister to S. monospora (included 16 samples, clade B) (PP = 1. 0, BS = 98; Fig. 3). Clade B includes S. trichophylla, S. monospora var. ciliolata, S. medogensis, and S. effusa var. dulongjiangensis
with strong support (PP = 1.0, BS = 99). This is confirmed by their gross morphology and micromorphology of spores described above.
Discussion
The molecular and morphological evidence suggests that S. monospora s. l. represents two distinct species. We resurrected the older name S. microclada from S. monospora for specimens with slightly dimorphic strobili. We treated S. medogensis, S. trichophylla, S. monospora var. ciliolata, and S. effusa var. dulongjiangensis as synonyms of S. monospora.
Fig. 2. Mega- and microspores of Selaginella microclada (A-D) and Selaginella monospora (E-H): A, C, E, G -Megaspores; B, D, F, H - Microspores. A, B from X. P. Qi Q070 (PE); C-D from D. D. Tao 1011 (PE); E-F from Lu et Zhang 27625-B (PE); G-H from Lu et Zhang 27625-B (PE).
The type specimens of S. trichophylla and S. monospora var. ciliolata are morphologically similar to S. monospora but leaf surface with spinules which are found from shaded and wet habitat, and the types of S. medogensis and S. effusa var. dulongjiangensis are morphological abnormalities (Fig. 1), there are only two nucleotide substitutions in the rbcL sequences with S. monospora.
We have observed enough specimens from the distribution ranges of S. monospora, made critical studies of the type specimens and conducted molecular phylogenetic analysis with related species. The traditional recognized S. monospora is a mixture of two distinct taxa, i. e., S. microclada and S. monospora.
Fig. 3. The 50 % majority rule consensus tree derived from maximum likelihood showing the position of Selaginella microclada and S. monospora. Support values (BSML/PP ) are shown above the main branches; the dash (-) indicates BS < 50 %. The two distinct species are shown in bold. BI
TAXONOMIC TREATMENT
Key to Selaginella monospora and its related species
1. Sporophylls slightly dimorphic, dorsal leaves ovate-lanceolate or elliptic, base obtuse, margin
denticulate, apex acuminate or shortly aristate .......
...........................................................S. microclada
+ Sporophylls strongly dimorphic, dorsal leaves obliquely ovate-elliptic and ciliolate at margin or ovate-lanceolate or elliptic and denticulate at margin ............................................................................... 2
2. Ventral leaves strongly overlapping stems and branches, axillary leaves on main stems ovate, ovate-triangular, or ovate-elliptic .............S. effusa
+ Ventral leaves not overlapping stems and branches, axillary leaves on main stems ovate to broadly ovate....................................S. monospora
Selaginella microclada Baker, 1884, J. Bot. 22: 246.
Type: "India, Sikkim, Chong-tong, alt. 4000 ft. 22 VII 1862. Dr. Anderson 1404" (K - [K001067485] image online!).
- Selaginella monospora sensu Alston, 1945, Proc. Nat. Inst. Sci. India 11: 228, p. p.; K. Iwats., 1975, Fl. E. Himalaya 8: 168, p. p; K. Iwats., 1988, Univ. Mus. Univ. Tokyo Bull. 31: 243; Dixit, 1992, Selaginell. India: 92, p. p; Thapa, 2002, Pterid. Nepal: 27; X. C. Zhang, 2004, Fl. Reipubl. Popularis Sin. 6(3): 186; Singh, Panigrahi, 2005, Ferns and Fern-Allies ofArunachal Pradesh 1: 62; X. C. Zhang et al., 2013, Fl. China 2-3: 61; Fraser-Jenkins et al., 2015, Ferns Fern-Allies Nepal 1: 73, p. p; Fraser-Jenkins et al., 2017, Annot. Checkl. Indian Pterido-phytes: 39, p. p.; Shalimov et al., 2019a, PhytoKeys 133: 34, p. p.
Plants 35-85 cm, main stems long creeping. Rhizophores at intervals throughout length of main stem, borne on ventral side in axils of branches. Main stems branched throughout, pinnately branched, stramineous, 1.5-2 mm in diam. in lower part, stem oval or terete, not sulcate, branches many, some primary lateral branches developed into long branch systems; primary leafy branches 8-12 pairs, once or twice pinnately branched or 2 or 3 times forked. Axillary leaves ovate, narrowly ovate, or narrowly elliptic, 2-3 x 0.8-1.6 mm, base exauriculate, margin denticulate. Dorsal leaves ovate-lanceolate or elliptic, 1-1.6 x 0.3-0.7 mm, carinate or strongly carinate, base obtuse, not peltate, margin denticulate, apex acuminate or shortly aristate. Ventral leaves ovate-triangular or oblong-falcate, 2.6-5.5 x 0.9-2.3
mm, basiscopic base decurrent, margin subentire or entire; acroscopic base enlarged, broader, strongly overlapping stem and branches, margin denticulate; apex subacute. Strobili solitary, dorsiventrally complanate (sometimes appearing uniform), 3-20 x 1.93.2 mm; sporophylls slightly dimorphic, resupinate, not white-margined; dorsal sporophylls lanceolate, sharply carinate, margin minutely denticulate, apex acuminate, with sporophyll-pteryx complete and denticulate; ventral sporophylls ovate-lanceolate, carinate, base dilated, margin denticulate; microspores yellowish orange or pale yellow, megaspores whitish.
Ecology: On damp banks in forests. (400)1300-1800(2600)m.
Distribution: BHUTAN, CHINA (Hainan, Guangdong, Guangxi, Guizhou, Xizang, Yunnan), NEPAL (Kathmandu, Kabhrepalanchok, Udayapur, Solukhumbhu, Sankhuwasabha, Tehrathum, Jha-pa, Ilam, Taplejung), INDIA (Uttarakhand, Assam State, Sikkim, Daijeeling, Manipur, Meghalaya, West Bengal), LAOS, CAMBODIA, THAILAND, VIETNAM.
Selected specimens examined:
INDIA: Assam: "Tura Peak, Garo Hills, 4000 ft. IV 1888. G. Mann 65" (US01392971); "l. c. G. Mann s. n." (L.3498106). Meghalaya: "Barapani, Khasi Hills, deep forest, on boulder, 3000 ft. 22 V 1949. W. N. Koelz 22751" (MICH1507940); ibid. "W. N. Koelz 22951" (MICH1507942). Uttarakhand: "Mussoorie, 1956. P. S. Sabharwa s. n." (U). Darjiling: "7000 ft. IX 1958. S. S. Bir s. n." (PE01622251; US01392974; U.1603187); ibid., "2185 m, 3 XII 1932. T. N. Liou L 6336" (PE); ibid., "3000 ft. 23 VIII 1869. F. Henderson s. n." (E00754770); ibid., "2500 ft. 31 X 1870. F. Henderson 13644" (E00754773; E00800129); ibid., "19 VIII 1841. Dr. Samerson 430" (E00800136); ibid., "7000 ft. 22 XII 1936, W. Koelz 10487" (US01392972); ibid., "Lebong forest, 5500 ft. 8 VIII 1956. S. S. Bir 197" (US01392975). Sikkim: "Gangtok, common on moist banks, 5000 ft. 7 IV 1939, F. Ludlow, G. Sherriff, G. Taylor 4005" (PE 01622250; MICH1578741); ibid., "Dikchu, 2010 m, 1972, Pradhan et al. 61" (E).
NEPAL: E Nepal: Sankhuwasabha: "en route from Harelo to Chichila, Castanopsis hystrix forest, on the moist rock. Alt. 1935 m, 2 VI 1978. H. Tabata, K.R. Rajbhandari, Y. Shimizu 11051" (PE 01622247); "Above Shinbun-Hatia Gola, alt. 1600-2100 m, 3 VIII 1977. H. Ohashi et al. 771954" (TI); "Seduwa, in shade under rocks, 5000 ft. 7 VI 1965. Banerjee, Upadhyay, Baskola 3322"
(US01392965); "Seduwa (Kasuwa Khola) prostrate, alt. 3000 ft. 6 V 1965. Banerjee, Upadhyay, Baskola 3374" (US01392966). Solukhumbhu: "Near Namche, in shady of rocks, rare, alt. 8000 ft. 9 V 1965. Banerjee, Upadhyay, Baskola 3418" (US01392968); "Near Namche, in shade, alt. 8000 ft. 9 V 1965, Banerjee, Upadhyay, Baskola 3420" (US01392967). Taplejung: "Khebang below Siling Tzokupa, 20 XI 1963. H. Hara et al." (TI). Ilam: "Densely forested, rocky stream-gully of Sudhung Khola, shortly below Sudhung khola, shortly below Sudhung, leading south below road, below Sundergaon, W. of Pashupatinagar on Ilam Road, 9 X 2001. C. R. Fraser-Jenkins 29549 (FN 5494)" (US01392964; L.3726784); "in Oak-forest on slopes above streams, between Chitregaon and Manebhan-jyang, ca. 4-5 km N. E. of Pashupatinagar, on footpath to Manebhanjyang near indian border, N. E. of Ilam, 23 X 2001. C. R. Fraser-Jenkins 29069 (FN 5584)" (US01392963; L.3726782). Jhapa: "ca. / km below and S. of Kuttedara on road to Bhudaba-re, N. of Charali on road to Phikal and Ilam, NE of Birtamod and NW of Kakkarbhitta, forested Khola (stream-gully), planted with ginger, belowroad, alt. 5000 ft. 16 VIII 1998, C. R. Fraser-Jenkins 26586 (FN 2564)" (US01392962; L.3726781).
BHUTAN: "Yongpu La, 6000 ft, on shady bank in forest, 10 XI 1938. F. Ludlow, G. Sherriff, G. Taylor 7210" (PE 01622248); "Shamgong, 11.6 km South of Shamgong on the road to Gaylegh-phug, 1500 m, growing on the forest floor, 9 V 1984.
B. Bartalomew 1620A" (PE 01622249); "Gayleg-phug, slope below Shan Khara, 27°01' N, 90°34' E, warm broad-leaved forest on steep hillside. Se-laginella forming carpets on steep bank, 1250 m. 29 III 1982. A. J. C. Grierson, D. G. Long 4092" (E00754762); "Chukka district: Tabji Khola, Rai-dak valley, 26°51'N, 89°34'E, warm broad-leaved forest on shady mossy bank, 1700 m. 23 II 1982. A. J. C. Grierson, D. G. Long 3148" (E00800130); "1 km S of Tongsa, 27° 30' N, 90° 30' E, damp rock face in Quercus forest, ca. 2250 m. 17 V 1979, A. J.
C. Grierson, D. G. Long 1115" (E00800133); "2 km below Putlibhir, on damp shady rock at roadside, ca. 2000 m. 7 VI 1975. A. J. C. Grierson, D. G. Long 18" (E00800140); "Tronsa/Trashiling, 7500 ft, wet shady banks, 5 VIII 1949, F. Ludlow, G. Sherriff, J.K. Hicks 17028" (L.3498102); "Yuto La, between Bumthang and Trongsa, shady banks in deciduous forest, 8500 ft. 4 VIII 1949. F. Ludlow, G. Sherriff, J. K. Hicks 17023" (L.4328981).
CHINA: Hainan: "Diaoluoshan, 8 I 1975. W. M. Chu 5895" (PYU); "Diaoluoshan, 6 I 1975. W.
M. Chu 5896" (PYU); "Wuzhishan, 28 XII 1974. W. M. Chu 1768" (PYU); "Jianfengling, 13 XII 1974. W. M. Chu 1512" (PYU). Guangxi: "Napo, Xiahua, Guinongshan, 1300 m. 15 IV 1998. H. N. Qin et al. 958" (KUN1288810). Xizang: "Nyingchi, Linzhi, 936 m. 9 V 2011. Q. Tian et al. TQ00517" (CSH0071243); "Megog, Beibeng, 810 m. 11 VII 1974. Qingzang team 74-1911" (PE00405934; PE00405933); "Medog, 12 IX 2016. Y. H. Yan et al. ZXL07275" (CSH0137167). Yunnan: "Zhen-kang, 2800 m. III 1936. C. W. Wang 72509" (PE00405931); "Congyuan, Nangunhe nature reserve, 1 III 2001. J. C. Zhao 2000-3" (PE00405930); "Lushui, 2380 m. 18 VI 1981. Y. F. Han et al. 81768" (PE01593691); "Lushui, II 1935. C. W. Wang 71971" (PE00244926, PE00244931); "Longling Xian, Longjiang Xiang. Xiaoheishan Provincial Preserve. Xiaotai Shan, Xiaoshui He, on W side of Gaoligong Shan, S of the new road from Baoshan to Tenchong via Nankang Yakou. 2011 m. 24°49'44"N, 98°45'36"E. Subtropical evergreen broadleaf forest. Disturbed by felling. Secondary forest with grassy cleared areas along streams. Growing under shrubs at edge of forest, 21 VIII 2003. Gaoligong Shan Biodiversity Survey 17312" (KUN1407313); "Gongshan, 1700 m. 1935. C. W. Wang 67309" (KUN); ibid., "Chiu-Kiang, W of Champutung, 1700 m. X 1935. C. W. Wang 67309" (KUN0801785); ibid., "Hills around Jengyueh, II 1913. G. Forrest 9694" (E00823623); "Hills around Jengyueh, 6-8000 ft, II 1913. G. Forrest 9697" (E00823624); ibid., "Hills around Jengyueh, 5000 ft. V 1912. G. Forrest 7705" (E00823625); "Western Yunnan, without exact locality, 1952. R. C. Ching 50510" (PE00244911); "l. c., R. C. Ching 50513" (PE00244913); "Tenchong Xian, Jietou Xiang, Datang Cun. Vicinity of Erda-gou Wailinggan, NNE of Datang. W side of Gaoligong Shan. 25°44'26.1"N, 98°42'9.8"E. 2080-2150 m. Growing under forest along trail. Herb. ca. 20 cm tall. 14 V 2006. Gaoligong Shan Biodiversity Survey 30168" (KUN1407319); ibid., "Wuhe, Xieng, km 24.7 along Higway S, 317.1 km north along road to Tenghang village and ridge to the W of the river along trail to the Zhengding forest reserve station. 2146 m. 24°51'22"N, 98°45'23"E. Growing in a partially shaded dell on forest floor. 28 V 2005. Gaoligong Shan Biodiversity Survey 25074" (KUN1407318); ibid., "Jietou Xiang, Zhoujiapo on the road between Jiefou and Datang, 25°33'11"N, 98°40'6"E, disturbed secondary broadleaved evergreen forest and scrub, 1680 m, 28 X 1998. L. Heng, B. Bartholomew, D. Zhilia 11139" (KUN1407312, E00246016); "Honghe, Pingbian, Wantang, 24 XI
2017. Y. F. Li B170094" (CSH0150753); "Luc-hun, Laobian, 1500 m. 21 X 1973. D. D. Tao 1011" (KUN0801833); "Luchun, Laobian, 1500 m., 28 IV 1974. Luchun Team 178" (KUN0801880); "Yuanyang, Fengchunling, Jiejipo, alt. 1300 m. 7 VI 1974. Luchun Team 1619" (KUN0801884, KUN0801866); "Luchun, alt. 1800 m. 11 V 1974. Luchun Team 675" (KUN0801879, KUN0801878); "Wenshan, Laojunshan, alt. 1500 m. 6 V 1962.
G. M. Feng 22524" (KUN0801709, KUN0801708); "Xichou, Fadou, 14 III 1986. W. M. Chu et al. 21694" (PYU); "Jinping, Fenshuiling, Hetouzhai,17 VII 1980. W. M. Chu 3835" (PYU); "Jinping, 16 V 1976. W. M. Chu et al. 6356" (PYU); "Luchun, 10 VI 1976. W. M. Chu et al. 6605" (PYU); "Gongshan, Dulongjiang, 2 X 2004. W. M. Chu, R. Zhao 31295" (PYU); "Gongshan, Dulongjiang, 13 IX 1999. W. M. Chu et al. 29310" (PYU); "Simao, Caiyanghe Nature Reserve, forest on clay slope along road in forest, 1500 m. 8 VIII 2007. K. Iwatsuki, S. K. Wu 0701" (KUN1226629); "Lu-chun, Niu-bo-shui-ku, along trail at edge of forest, 1850-1950 m. 2 VIII 1994. S. K. Wu, X. Cheng, Y. P. Yang, K. Iwatsuki, J. Murata, H. Nagamasu 1466" (KUN0801724); "Lu-chun, Ba-dong-dao-ban, along trail in sparse secondary forest, 1650 m. 1 VIII 1994. S. K. Wu, X. Cheng, Y. P. Yang, K. Iwatsuki, J. Murata,
H. Nagamasu 1435" (KUN0801722); "Lu-chun, Fen-shui-ling, along trail at edge of dense secondary forest, 1700 m. 1 VIII 1994. S. K. Wu, X. Cheng, Y. P. Yang, K. Iwatsuki, J. Murata, H. Nagamasu 1420" (KUN0801721); "Shweli river drainage basin to summit of Shwell-Salween watershed east of Tengyueh, XI 1922, J. Rock 7689" (US01392969).
MYANMAR: Kachin: "Putao District, Putao Township, Maliraing area, buffer zone of Hkakab-orazi National Park, around camp 3. 27°41'25.8"N, 97°23'49.4"E. Growing along the riverside, 638 m. 11 XI 2016. T. Y. New, K. Armstrong, D. Little, P. P. Hnin, B. Shaung, T. Naing, S. N. Syn, J. Dan, D. Ram 543" (NY02654079); "Kachin State. Putao District. Naungmung Township, buffer zone of Hkakab-orazi National Park, around Hton Wan rest house. 27°30'10.3"N, 98°0'12.6"E. 729 m. 16 IX 2015. D. Little, K. Armstrong, S. Lwin, M.M. San, Z. N. Tun, P. S.Aung, A. Syn,A. Bai 1444" (NY02649226); "Kachin State. Putao District. Naungmung Township, buffer zone of Hkakaborazi National Park, around Tup Kwan rest house. 27°36'15.6"N, 98°9'31.3"E, 729 m. 16 X 2015. D. Little, K. Armstrong, S. Lwin, M. M. San, Z. N. Tun, P. S. Aung, A. Sun, A. Bai 1444" (NY02649226); "Kachin State. Putao District. Naungmung Township, buf-
fer zone of Hkakaborazi National Park, around Tup Kwan rest house. 27°36'15.6"N, 98°9'31.3"E, 1929 m. 25 IX 2015. K. Armstrong, D. Little, S. Lwin, M.M. San, Z.N. Tun, P.S. Aung, A. Syn, A. Bai 987" (NY02648619).
Sagaing: "Stilwell Road, Burma Assam Borden, 1942, H. Young 36" (MICH1173545); "l. c. H. Young 24" (MICH1173544).
THAILAND: Loei: "Northeastern: Phu Kra-dung, S. of Loi, 16°53rN, 101°53'E, 1100 m. 7-9 XI 1970. Ch. Charoenphol, K. Larsen, E. Warncke 4709 (3rd Expedition 1970)" (L.3484642). Udawn: "Loey: Phu Kradung, on the plain at its summit, alt. ca. 1100-1200 m, on rather dry slope in forest. 28 XI 1965. M. Tagawa, K. Iwatsuki, N. Fukuoka 587" (L.3484655).
LAOS: "Sam-Nena, 9 X 1920. E. Poilane 2031" (L.3726783; P01229420); "env. de Nape, 13 X 1928. M. Delacour s. n." (P01229412; P01254282).
VIETNAM: Annam: "Massif du Honba, Aout, 1919, F. Vincens s. n." (P01229422). Tuyên Quang: "Tam Dao National Park: North of Vietnam, 9001100 m., 21°27'47" N, 105°38'68"E, 31 X 2004. L. Q. Li, N. T. Hiep, Z. Y. Zhang, X. C. Zhang, T. G. Gao, Z. T. Wang, N. S. Khang, N. X. Tam 0638" (PE01849460).
Selaginella monospora Spring, 1850, Mém. Acad. Roy. Sci. Belgique. 24: 135, no. 80; Alston, 1945, Proc. Nat. Inst. Sci. India 11: 228; Panigra-hi, Dixit, 1968, Proc. Nat. Inst. Sci. India, 34B (4): 201; Dixit, 1992, Selaginell. India: 92, p. p.; Fraser-Jenkins et al., 2015, Ferns Fern-Allies Nepal 1: 73, p. p.; Fraser-Jenkins et al., 2017, Annot. Checkl. Indian Pteridophytes: 39, p. p.
= Lycopodium monosporum (Spring) Hook, 1857, Bot. Misc. 9: 362.
= Selaginella plumosa var. monospora (Spring) Baker, 1883, J. Bot. 21: 145.
Type: (lecto - designated by Fraser-Jenkins et al., 2017) [BHUTAN] "Bootan [Khegumpa, N. of Dewangiri (Deotang)], W. Griffith 391 Journal [24 I 1838], Herbarium Hookerianum" (K -[K001067478] image online!).
= S. gorvalensis Spring, 1850, Mém. Acad. Roy. Sci. Belgique. 24: 256, no. 195.
= Lycopodioides gorvalensis (Spring) Kuntze, 1891, Revis. Gen. Pl. 2: 826.
Type: [INDIA] "Gorval, Griffith s. n., in error for Khasia" (K - [K000009117] image online!).
= S. medogensis Ching et S. K. Wu, 1983, Fl. Xizang. 1: 23, pl. 6, syn. nov.
= S. effusa var. medogensis (Ching et S. K. Wu) W. M. Chu, 2006, Fl. Yunnan. 20: 719.
Lectotype (designated here): [CHINA] "Xizang austro-orient., Medog, in sylvis alt. 800 m, 19 VIII 1974. Qinghai-Xizang Complex Exped. 74-4375" (PE! - [PE00405946]; iso - PE! [PE00405947], KUN! - [KUN1217049], PYU! fragm.). - [INDIA] "Pachak-shiri Distr., Laung, plastered on the moss covered banks in dense mixed forest, 2000-2500 m. 9 V 1938. F. Ludlow, G. Sherriff, G. Taylor 3723" (paralecto - PE! - [PE00405945], BM?).
= S. trichophylla K. H. Shing, 1993, Acta Phy-totax. Sin. 31(6): 569, pl. 2, syn. nov.; W. M. Chu, 2006, Fl. Yunnan. 20: 83; X. C. Zhang et al., 2013, Fl. China 2-3: 61; Shalimov et al., 2019, PhytoKeys 133: 35.
= S. monospora subsp. trichophylla (K. H. Shing) X. C. Zhang, 2004, Fl. Reipubl. Popularis Sin. 6(3): 189.
Type: "China, Yunnan prov., Gongshan, Dulongjiang River, alt. 1450-1500 m, in sylvis frondosis, 22 VIII 1982, Qinghai-Xizang Exped. 9451" (PE! - [PE00452190]).
= S. monospora var. ciliolata W. M. Chu, 2006, Fl. Yunnan. 20: 719, syn. nov.
Type: "China, Yunnan prov., Jingdong, Ailao Shan, Xujiaba, alt. 2450 m, under evergreen broad-leaved forest, 22 IV 1982. J. J. He 13352" (PYU!; iso - PE!); "China, Yunnan prov., Wuliang Shan, 20 IV 1964. R. S. Wang et al. 14" (para - PYU!); "China, Yunnan prov., VII 1964, X. W. Ren 3824" (para - PYU!); "China, Yunnan prov., Wuliang Shan, Mengling, alt. 1800 m, on moist cliff in evergreen broad-leaved forest, 9 IX 1979. W. M. Chu, Z. Z. Jin 9073" (para - PYU!); "China, Yunnan prov., Wuliang Shan, Wenpu, Erdaoya, Qingtou, alt. 2100 m, under evergreen broad-leaved forest. 9 IX 1979. W. M. Chu, Z. Z. Jin 9299" (para - PYU!); "China, Yunnan prov., Malipo, Laojuan Shan, alt. 1600 m, under evergreen broad-leaved forest, 30 VII 1993. S. G. Lu, G. F. Zhang 27625-B)" (para -PYU!, PE!); "China, Guandong prov., Xinyi, Datian Ding, alt. 1600 m, under evergreen broad-leaved forest, 23 VIII 1984. W. M. Chu et al. 18296" (para -PYU!, PE!); "NEPAL, East Nepal, Mai Majuwa-Dhara Pani, alt. 1500-1600 m, 4 XII 1963. H. Hara, H. Kanai, S. Kurasawa, G. Murata, M. Togashi, T. Tuyama s. n." (para - KUN! - [KUN0801546], L - [L.3498103] image online!).
= S. effusa var. dulongjiangensis W. M. Chu, 2006, Fl. Yunnan. 20: 718, syn. nov.
Type: "China, Yunnan prov., Gongshan, Dulongjiang valley, between Kongdan and Xianjiudan,
on cliff at edge of evergreen broad-leaved forest, alt. 1500 m, 3 X 2004. W. M. Chu, Z. R. He 31299" (PYU!; iso - PE! - [PE n 2526065]), "China, Yunnan prov., Gongshan, Dulongjiang valley, Bukawa-ng, on cliff at edge of evergreen broad-leaved forest, alt. 1440 m, 2 X 2004. W. M. Chu & Z. R. He 31291" (para - PYU! - [PYU01065937]).
Plants creeping, 30-50 cm. Rhizophores at intervals throughout length of creeping stem and branches, borne on ventral side in axils of branches. Main stems pinnately branched throughout, stramineous, terete, not sulcate; primary leafy branches 10-15 pairs, once or twice pinnately branched, secondary branches forked or once pinnately branched, tertiary branches forked, branchlets sparse, adjacent primary branches on main stem 2-4 cm apart; leafy portion of main stem including leaves 5-6 mm wide at middle, ultimate branches 2.8-3.2 mm wide including leaves. Axillary leaves ovate, 1.2-2.5 x 1-2 mm, base exauriculate, margin ciliolate. Dorsal leaves ovate, 1.2-1.9 x 0.5-1 mm, not carinate, base obtuse or subcordate, not peltate, margin cili-olate, apex aristate to cuspidate. Ventral leaves on branches distant, slightly ascending or spreading, ovate-triangular, oblong or oblong-ovate 2-3.2 x 0.8-1.6 mm, margin denticulate, apex acute; ac-roscopic base enlarged, broader, overlapping stem and branches, margin denticulate, ciliolate. Sometimes upper surface ventral leaves with spines. Stro-bili solitary, dorsiventrally complanate, 6.0-10 x 1.2-2.3 mm; sporophylls dimorphic, resupinate, not white-margined; dorsal sporophylls ovate-lanceolate, carinate, margin denticulate, apex acuminate, with sporophyll-pteryx complete and denticulate; ventral sporophylls ovate-lanceolate, carinate, margin denticulate; megasporophylls in basal portion on lower side of strobilus; microsporangia transversely elliptic, rather thin, upper margin cells differentiated; microspores pale yellow, megaspores white-yellow.
Ecology: on moist cliff in evergreen broad-leaved forest. Alt. 1100-3033 m.
Distribution in Pan-Himalaya region: NEPAL (C Nepal, E Nepal), INDIA (Sikkim), BHUTAN, CHINA (Yunnan, Sichuan).
General distribution: INDIA (Assam, Sikkim, Meghalaya), NEPAL (Dolakha, Sankhuwasabha), BHUTAN, CHINA (Hainan, Guangxi, Guizhou, Guandong, Yunnan, Sichuan), MYANMAR (Kachin), VIETNAM (Cao Bang, Quang Ninh).
Notes: In the protologue, Ching and S. K. Wu (in S. K. Wu, 1983) cited single specimens "Qinghai-Xizang Complex Exped. 4375" with the label as
"type" kept at HP (= PE). Both of these sheets had the field number "4375" attached, but the herbarium label already had the entry "74-4375"; to correct this omission we add the citation numbers from the herbarium label. Besides, these herbarium sheets contain a signature on the herbarium label with the name "Selaginella mutensis Ching and S. K. Wu sp. nov." nom. in shed. herb., this name was later used in the work of Liu et al., 2005.
Selected specimens examined:
BHUTAN: "Rukubi (2600) - Chendebi (2300) -Charikhachor (2250) - Neylong (2200), 14 IV 1967. H. Hara, H. Kanai, G. Murata, H. Ohashi, O. Tana-ka, T. Yamazaki 4105" (KYO); "Yuto La, between Bumthang and Trongsa, 8500 ft. Shady banks in deciduous forest, 4 VIII 1949. F. Ludlow, G. Sherriff, J.H. Hicks 17023" (KYO; L.4328981).
CHINA: Hainan: "Wuzhi Mt. 18°53'N, 109°41'E, 1800 m, in forest, 18 XII 2004. Wuzhishan Fern Survey 134" (PE); "Wuzhishan, 1700-1800 m. 16 XI 2000. X. C. Zhang, G. M. Zhang, D. Li 1698" (PE); "Wuzhishan, 1700-1800 m. 16 XI 2000. X. C. Zhang, G. M. Zhang, D. Li 1704" (PE); "Wuzhishan, 1700-1800 m. 16 XI 2000. X. C. Zhang, G. M. Zhang, D. Li 1699" (PE); "Qiongzhong li and Miao autonomous county, Wuzhi Mt. one peak to two peaks, 1800 m. 28 XII 1974. 236-6 n 01798" (PE 00405937); "Wuzhi Mt. 18°89813'N, 109°70399'E. 28 XI 2018. Y. R. Wang 9375" (PE); "Wuzhi Mt. 18°89810'N, 109°20349'E, 2 XI 2018. Y. R. Wang 9370" (PE); "Wuzhi Mt. 18°89716'N, 109°70121'E, 2 XI 2018. Y. R. Wang 9371" (PE); "Qiongzhong, Wuzhishan, 1800 m. 28 XII 1974. 236-6 Team 01798" (PE00405937). Guizhou: "Pana, Chengfeng, on cliff, 25 IX 1930. Y. Tsiang 4418" (PE00405630); "T. N. Liou 6718" (PE00405632); "Du Yun City, Cape Mountain Nature Reserve, under forest, alt. 998 m. 7 X 2007. Z. X. Lin, Y. L. Lan 07313" (KUN); "l. c., Z. X. Lin, Y. L. Lan 07315" (KUN); "Duyun, Doupengshan Nature Reserve, 7 X 2007. Z. X. Lin, Y. L. Lan 07313" (HUST); ibid., "7 X 2007. Z. X. Lin, Y. L. Lan 07315" (HUST); "Zhenfeng, 25 X 1930. Y. Tsiang 4418" (PE); "T. N. Liou 6718" (PE00405632). Guandong: "Anonymous 615" (PE00405928); "Sun Yat-sen University 615" (PE 00244908); "ZXL 10058" (CSH0044647); "Xin yi county, Fog ridge, 13 VIII 2004. Y. H. Yan, N. L. Jin, Y. Wang, X. F. Zhu WYD 404" (CSH0068766); "Xinyi, Dawuling, 3 VIII 2003. Y. H. Yan et al. 1403" (PE). Guangxi: "Gui Ping City, Da Ping Shan, Lv Chong, evergreen broad-leaved forest, sandstone, alt. 600 m.
24 IX 1987. H. G. Zhou, Y. Li 939" (PYU); "Gui Ping City, Lv Chong, bamboo forest, sandstone, alt. 200 m. 27 IX 1987, Y. Li, H. G. Zhou 1012" (PYU); "Gui Lin City, Yong Fu county, Baoli township Qingping village siding, broad-leaved forest, forest, yellow brown soil, alt. 326 m. 24°50'52.8"N, 110°08'24.0"E, 4 III 2013. Yongfu county census team, 450326130304011LY" (GXMG 0152819); "Jinxiu mountain, 2 XI 2018. Y. R. Wang 9540" (PE); "Guiping, Taipingshan, 24 IX 1987. H. G. Zhou, Y. Li 939" (PYU); "Guiping, Luchong, 27 IX 1987. Y. Li, H. G. Zhou 1012" (PYU). Yunnan: "Xichou county, Fadou village, dank humus under the forest, alt. 1600 m. 15 III 1987. W. M. Chu et al. 21738" (PYU); "Ping Bian county, Da Wei Shan, Y. Jiao 2000-03" (PYU); "Jing Dong county, on a dry hillside, alt. 2100 m. 1 X 1956. Y. Qiu 52468" (KUN0801134); "Jing Dong county, under a forest on a hillside, alt. 1800 m. 5 VI 1986. Wu Liang Shan team 0023" (KUN); "Houqiao, Tengchong county, forest, 1750-1800 m, 31 III 2003. X. C. Zhang 2992" (PE); "Gongshan, alt. 1300-1400 m, 24 I 2017. X. C. Zhang et al. 8121" (PE); Tengchong, Dahetou, 14 V 2016. Z. D. Fang et al. DJDC-653" (SABG); "Tengchong, Houqiao, 31 III 2003. X. C. Zhang 1403" (PE); "Tengchong, Houqiao, 12 IX 2003. X. C. Zhang 3007" (PE); "Tengchong, Houqiao, 12 IX 2003. X. C. Zhang 3009" (PE); 'Yongde, Daxueshan, 8 XI 1982. W. M. Chu et al. 14863 (A)" (PYU); "Xichou, Fadou, 15 III 1987. W. M. Chu et al. 21738" (PYU); "Pingbian, Daweishan, Y. Jiao 2000-03" (PYU); "Jinping, 1400 m. 10 IV 1959. Class 1960 Plant Resources Investigation Team 1014" (PYU); "Fugong Xian, Lumadeng Xian, ca. 2.8 direct km SSW of the Yaping bridge, on the W side of the Nujiang, ca. 22.5 direct km N of Fugong city. 27°6'29''N, 98°52'11''E, disturbed secondary forest. Growing in thicket, moist, near stream in forest shade area, creeping perennial herb, 1255 m. 28 IV 2004. Gaoligong Shan Biotic Survey Expedition 19963" (E00268722); "Fugong Xian, Yaping Xian, Vicinity of Shibali logging station on the road from the Nujiang to Yaping pass, E side of Gaoli-gong Shan, 2485 m. 27°9'48"N, 98°46'58"E, growing on rock in forest, creeping herb, 28 IV 2004. Gaoligong Shan Biotic Survey Expedition 20013" (E00258354). Sichuan: "Jun Lian county, liberation commune Shui Tou creek, canyons, under broad-leaved forests, on rock walls, 14 VI 1978. H. S. Kung 5255" (PYU); "Jun Lian county, liberation commune shui tou creek, canyons, under broad-leaved forests, on wet rock walls, alt. 1100 m, 14
VI 1978, H. S. Kung 5229" (PYU); ibid., "14 VI 1978. H. S. Kung 5225" (CDBI); "Shimian, Caoke, Tangjiashan, 15 IX 1991, s. l. 1434" (CDBI). Xi-zang: "Medog, Beibeng, 800-1400 m, 2 V 2021. Zhang X. C. et al. 11429" (PE).
INDIA: Assam: "Indochina, Assam, Griffith s. n." (P00523054). Sikkim: "E Sikkim District. Above and S of Penlang Bazaar, below and on way up to Namphung Peak of the Tinjure ridge, W. of Tashi viewpoint, across valley to the north of Gangtok. Just below crest on N side of densely mixed-forest ridge, 29 IX 1998. C. R. Fraser-Jen-kins 27054 (FN 3031)" (L.4328985); "Pachak-Shiri Distr., Laung, «plastered on the moss covered banks in dense mixed forest», 28°45'N, 94°15'E, 7-8000 ft. 9 V 1938. F. Ludlow et al. 3723" (PE00405945, ?BM). Meghalaya: "Arwah Cave, under forest, 4 IV 2019. CPG Expedition CPG40561 (Lai Y. J., Zhang Q.)" (PE).
NEPAL: C NEPAL: Dolakha: "Jiri, Dolakha, alt. ca. 2200 m, 4 X 1988. T. Nakaike 3076" (PE). E NEPAL: Sankhuwasabha: "Seduwa, in shade under rocks, 5000 ft. 7 VI 1965. Banerjee, Upad-hyay, Baskola 3322" (US01392965). ILAM: "Mai Majuwa-Dhara Pani, alt. 1500-1600 m, 4 XII 1963. H. Hara, H. Kanai, S. Kurasawa, G. Murata, M. To-gashi, T. Tuyama s. n." (KUN; L.3498103).
MYANMAR: Kachin: "Kachin state, 27°31'56.4"N, 96°08'51.1"E, alt. 2393 m. 18 X 2016. T. Y. Nwe et al. Tyn380" (PE); "Kachin sta-
te, 27°30'55.2"N, 96°78'03.2"E, alt. 3033 m. 20 X 2016. T. Y. Nwe et al. Tyn410" (PE).
VIETNAM: Quang Ninh: "Tonkin (Dam-Ha: Sai Wong Mo Shan (Sai Vong Mo Leng) Lomg Ngong Village, 18 VII - 9 IX 1940. W. T. Tsang 30240" (E00829326).
Acknowledgements
We are grateful to Dr. Dmitry A. German of Altai State University (Barnaul, Russia), Dr. Xiang-Yun Zhu of Institute of Botany (CAS, Beijing, China) and Dr. Alexander N. Sennikov of Finnish Museum of Natural History, University of Helsinki (Helsinki, Finland) for their valuable and helpful comments on a draft of this manuscript and nomenclatural discussion relating to this paper. We are also grateful to the curators and the staffs of several herbaria B, CDBI, CSH, E, K, KUN, KYO, L, NY, P, PE, PYU, TI, U, and US for permitting the study of the collections including their digitized herbarium images. We are also thankful to the staff of the Global Plants project database (JSTOR, https://plants.jstor.org) and Global Biodiversity Information Facility (GBIF, https:// www.gbif.org) for open access digitized herbarium images. The authors thank Hongrui Zhang, Siqi Liang, Meng-Hua Zhang, and Ya-Rong Wang for discussion, manuscript language checking, and for help with experiments. The joined author was sponsored by CAS-TWAS President's Fellowship for International PhD Students.
References
Alston A. H. G. 1934. An enumeration of the Chinese species of Selaginella. Bull. Fan Mem. Inst. Biol., Bot. 5(6): 261-304.
Alston A. H. G. 1945. An enumeration of the Indian species of Selaginella. Proc. Natl. Inst. Sci. India 11: 211-235. ChingR. C., Wu S. K. 1983. Selaginella. In: FloraXizangica. Vol. 1. Beijing: Science Press. Pp. 19-21. Chu W. M. 2006. Selaginellaceae. In: Flora Yunnanica. Vol. 20. Beijing: Science Press. Pp. 35-93, 719. Chun W. Y., Chang C. C., Chen F. H. 1964. Selaginella. In: Flora Hainanica. Vol. 1. Beijing: Science Press. Pp. 9-13.
Dixit R. D. 1992. Selaginellaceae of India. Dehra Dun, Bishen Singh Mahendra Pal Singh. 196 pp. Fraser-Jenkins C. R. 2008. Selaginellaceae. In: Taxonomic revision of three hundred Indian subcontinental Pte-ridophytes with a revised census-list (A new picture of fern-taxonomy and nomenclature in the Indian subcontinent). Dehra Dun: Bishen Singh Mahendra Pal Singh. Pp. 11-18.
Fraser-Jenkins C. R., Kandel D. R., Pariyar S. 2015. Selaginellaceae. In: Ferns and fern-allies of Nepal. Vol. 1. Kathmandu, Nepal: National Herbarium and Plant Laboratories, Department of Plant Resources, Ministry of Forests and Soil Conservation. Pp. 58-86.
Fraser-Jenkins C. R., Gandhi K. N., Kholia B. S., Benniamin A. 2017. Selaginellaceae. In: An annotated checklist of Indian Pteridophytes, Pt. 1 (Lycopodiaceae to Thelypteridace). Dehra Dun: Bishen Singh Mahendra Pal Singh. Pp. 27-53.
Ghosh S. R., Ghosh B., Biswas A., Ghosh R. K. 2004. Selaginellaceae. In: The Pteridophytic Flora of Eastern India. Vol. 1. Kolkata: Botanical Survey of India. Pp. 88-131.
Hall T. A. 1999. BioEdit: A user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series 41: 95-98.
Huelsenbeck J. P., Ronquist F. 2001. MRBAYES: Bayesian inference of phylogenetic trees. Bioinformatics 17: 754-755.
Iwatsuki K. 1975. Pteridophyta. In: Flora of Eastern Himalaya. Vol. 8. Tokyo: The University Museum, the University of Tokyo. Pp. 166-205.
Iwatsuki K. 1988. An enumeration of the pteridophytes of Nepal. In: The Himalayan Plants. Vol. 1. H. Ohba, S. B. Malla (eds). Tokyo: University of Tokyo Press. Pp. 231-339.
JermyA. C. 1990. Selaginellaceae. In: Thefamilies andgenera of vascular plants, pteridophytes and gymnosperms. 1st eds. Vol. 1. Berlin: Springer. Pp. 39-45.
Jiang R. H. 2013. Selaginella. In: Flora of Guangxi. Vol. 6. Pteridophyta. Nanning: Guangxi Science and Technology Publishing House. Pp. 15-37.
Liu J.-X, Yan X.-F. 2005. Observations on spore morphology of four species of Selaginellaceae from Xizang (Tibet). Chinese Bulletin of Botany 22, 1: 44-49.
Panigrahi G., Dixit R. D. 1966. Studies in the systematics of Indian Selaginella. I. Proc. natn. Inst. Sci. India 34, B 4: 201-202.
Posada D. 2008. jModelTest: Phylogenetic model averaging. Molecular Biology and Evolution 25(7): 1253-1256. DOI: 10.1093/molbev/msn083
Shalimov A. P., Wu Y. D., Zhang X. C. 2019a. A taxonomic revision of the genus Selaginella (Selaginellaceae) from Nepal. PhytoKeys 133: 1-76. DOI: 10.3897/phytokeys.133.37773
Shalimov A. P., Zhu Y. M., Zhang M. H., Zhang X. C. 2019b. Selaginella dianzhongensis (Selaginellaceae), a new spikemoss from China. PhytoKeys 118: 75-87. DOI: 10.3897/phytokeys.118.30375
Shing K. H. 1993. Some new species of Pteridophytes from Hengduan Mountains. Acta Phytotaxonomica Sinica 31(6): 569-574.
Singh S., Panigrahi G. 2005. Selaginellaceae. In: Ferns andfern-allies of Arunachal Pradesh. Vol. 1. Fern-Allies and Ferns: Adiantaceae to Nephrolepidaceae. Dehra Dun: Bishen Singh Mahendra Pal Singh. Pp. 55-76.
Spring A. F. 1850. Monographie de la Famille des Lycopodiacées, Seconde partie. Mém. Acad. Roy. Sci. Belgique 24: 1-358.
StamatakisA. 2014. RAxML Version 8: A tool for phylogenetic analysis and post-analysis of large phylogenies.
Bioinformatics (Oxford, England) 30(9): 1312-1313. DOI: 10.1093/bioinformatics/btu033
Tagawa M., Iwatsuki K. 1979. Selaginellaceae. In: Flora of Thailand. Vol. 3(1) Pteridophytes. Bangkok: The Tistr Press. Pp. 14-32.
Thapa N. 2002. Pteridophytes of Nepal. Bull. Department Plant Resources 19: 1-175.
Thiers B. [2021]. Index Herbariorum: A global directory of public herbaria and associated staff. In: New York Botanical Garden's Virtual Herbarium. URL: http://sweetgum.nybg.org/science/ih/ (Accessed 26 July 2021).
Thompson J. D., Gibson T. J., Plewniak F., Jeanmougin F., Higgins D. G. 1997. The CLUSTAL_X windows interface: Flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research 25(24): 4876-4882. DOI: 10.1093/nar/25.24.4876
Tryon A. F., Lugardon B. 1991. Spores of the Pteridophyta: surface, wall structure, and diversity based on electron microscope studies. Berlin: Springer-Verlag. 670 pp.
Zhang H. R., Wei R., Xiang Q. P., Zhang X. C. 2020. Plastome-based phylogenomics resolves the placement of the sanguinolenta group in the spikemoss of lycophyte (Selaginellaceae). Molecular Phylogenetics and Evolution 147: 106788. DOI: 10.1016/j.ympev.2020.106788
Zhang X. C. 2004. Selaginellaceae. In: Flora Republica Popularis Sinicae. Vol. 6(3). Beijing: Science Press. Pp. 86-219.
Zhang X. C., Nooteboom H. P., Kato M. 2013. Selaginellaceae. In: Flora of China. Vol. 2-3. Beijing: Science Press and St. Louis: Missouri Botanical Garden Press. Pp. 37-66.
Zhou X. M., Zhang L. B. 2015a. A classification of Selaginella (Selaginellaceae) based on molecular (chloroplast and nuclear), macromorphological, and spore features. Taxon 64: 1117-1140. DOI:10.12705/646.2
Zhou X. M., Jiang L. J., Zhang L., Gao X. F., He Z. R., Zhang L. B. 2015b. Spore morphology of Selaginella (Selaginellaceae) from China and its systematic significance. Phytotaxa 237: 001-067. DOI: 10.11646/ phytotaxa.237.1.1
Weststrand S., Korall P. 2016. A subgeneric classification of Selaginella (Selaginellaceae). American Journal of Botany 103(12): 2160-2169. DOI: 10.3732/ajb.1600288
Wu T. 2006. Selaginellales. In: Flora of Guandong. Vol. 7. Guangzhou: Guandong Science and Technology Press. Pp. 15-22.
Appendix 1
Specimen information and GenBank accession numbers (* indicates new accession)
Taxon Voucher Country rbcL atpI psbA
Selaginella albociliata P. S. Wang Zhang X.-C. 7242 (PE) Guizhou, China MH814882 MH814826 MH814854
Selaginella amblyphylla Alston Zhang X.-C. 7951 (PE) Yunnan, China MH814884 MH814828 MH814856
Selaginella amblyphylla Alston Zhang X.-C. 2924 (PE) Yunnan, China MH814883 MH814827 MH814855
Selaginella bodinieri Hieron. Zhang X.-C. 5 (PE) Chongqing, China, MH814885 MH814829 MH814857
Selaginella bodinieri Hieron. Zhang X.-C. 526 (PE) Sichuan, China MH814886 MH814830 MH814858
Selaginella bodinieri Hieron. Zhang X.-C. 7069 (PE) Guizhou, China MH814887 MH814831 MH814859
Selaginella braunii Baker Zhang X.-C. 1332 (PYU, CDBI) Hainan, China KT161420.1 - -
Selaginella chaetoloma Alston Zhang X.-C. 7347 (PE) Guizhou, China MH814889 MH814833 MH814861
Selaginella chaetoloma Alston Guo Z.-Y. 2016014 (PE) Guizhou, China MH814888 MH814832 MH814860
Selaginella chingii Alston Zhang X.-C. 7904 (PE) Guangxi, China MH814890 MH814834 MH814862
Selaginella chrysocaulos (Hook. et Grev.) Spring Zhang X.-C. 86 (PE) Sichuan, China MH814891 MH814835 MH814863
Selaginella ciliaris (Retz.) Spring Zhang X.-C. 7780 (PE) Yunnan, China MH814892 MH814836 MH814864
Selaginella decipiens Warb. Zhang X.-C. 1130 (PE) Guangxi, China MH814893 MH814837 MH814865
Selaginella deflexa Brack. - - AF093253.1 - -
Selaginella delicatula (Desv. ex Poir.) Alston Gao et al. HGX10734 (CDBI) - KT161441.1 - -
Selaginella dianzhongensis X.-C. Zhang Zhu Y.-M. 8158 (PE) Yunnan, China MH814909 MH814853 MH814881
Selaginella effusa var. dulongjiangensis W. M. Chu Chu et al. 31299 (PYU) Yunnan, China KT161449.1 - -
Selaginella helvetica (L.) Link Zhou 093 (CDBI) - KT161472.1 - -
Selaginella heterostachys Baker Zhang X.-C. 7088 (PE) Guizhou, China MH814896 MH814840 MH814868
Selaginella heterostachys Baker Zhang X.-C. 7268 (PE) Guizhou, China MH814897 MH814841 MH814869
Selaginella kraussiana (Kunze) A. Braun Zhou 062 (CDBI) Yunnan, China, Cult. KT161498.1 - -
Selaginella kurzii Baker Zhang X.-C. 1934 (PE) Yunnan, China MH814898 MH814842 MH814870
Selaginella labordei Hieron. ex Christ Zhang X.-C. 3356 (PE) Hubei, China MH814899 MH814843 MH814871
Selaginella laxistrobila K. H. Shing Chu et al. 24449 (PYU) Yunnan, China, KT161509.1 - -
Taxon Voucher Country rbcL atpI psbA
Selaginella medogensis Ching et S. K. Wu Qinghai-Xizang Complex Exped. 744375 (PE) Tibet, China OK247696* - -
Selaginella megaphylla Baker Jin X.-H. 19301 (PE) Tibet, China MH814901 MH814845 MH814873
Selaginella microclada Spring 100000 mountain investigation team 109 (PE) Guangxi, China OK247697* OK247681* OK247713*
Selaginella microclada Baker Wei Ran 342 (PE) Yunnan, China OK247699* OK247682* OK247714*
Selaginella microclada Baker Zhang X.-C. et al. 7296 (PE) Guizhou, China 0K247700* OK247683* OK247715*
Selaginella microclada Baker Zhang X.-C. 7889 (PE) Guangxi, China MH814902 MH814846 MH814874
Selaginella microclada Baker Zhang X.-C. et al. 8107 (PE) Yunnan, China 0K247701* OK247684* OK247716*
Selaginella microclada Baker Zhang X.-C. et al. 8307 (PE) Yunnan, China 0K247702* OK247685* OK247717*
Selaginella microclada Baker Zhang X.C. et al. 11429 (PE) Xizang, China OK247698* - -
Selaginella monospora Spring CPG Expedition CPG40561 (PE) Meghalaya, India 0K247709* OK247692* OK247722*
Selaginella monospora Spring Zhang X.-C. et al. 8121 (PE) Yunnan, China 0K247705* OK247688* 0K247720*
Selaginella monospora Spring Zhang X.-C. et al. 8122 (PE) Yunnan, China 0K247706* OK247689* OK247721*
Selaginella monospora Spring Zhang X.-C. 2992 (PE) Yunnan, China 0K247703* OK247686* OK247718*
Selaginella monospora Spring Yan Y.-H. et al. WYD404 (PE) Guangdong, China OK247711* OK247694* OK247724*
Selaginella monospora Spring Zhang X.-C. 6016 (PE) Guangxi, China 0K247704* OK247687* OK247719*
Selaginella monospora Spring Yan Y.-H. et al. WYD427 (PE) Guangdong, China OK247712* OK247695* OK247725*
Selaginella monospora Spring Wang Y.-R. 9370 (PE) Hainan, China 0K247707* 0K247690* -
Selaginella monospora Spring Wang Y.-R. 9371 (PE) Hainan, China 0K247708* OK247691* -
Selaginella monospora var. ciliolata W. M. Chu Lu S. G., Zhang G. F. 27625-B (PE) Yunnan, China 0K247710* OK247693* OK247723*
Selaginella nipponica Franch. et Sav. Zhou et al. DJY07479 (CDBI) Sichuan, China KT161542.1 - -
Selaginella ornata (Hook. et Grev.) Spring Zhang X.-C. 8520 (PE) Yunnan, China MH814903 MH814847 MH814875
Selaginella remotifolia Spring Zhou 005 (PYU, CDBI) Yunnan, China KT161580.1 - -
Selaginella repanda (Desv. ex Poir.) Spring Li B.-G. sn_20 (PE) Yunnan, China MH814906 MH814850 MH814878
Selaginella repanda (Desv. ex Poir.) Spring Zhang X.- C. 9273 (PE) Yunnan, China MH814905 MH814849 MH814877
Taxon Voucher Country rbcL atpI psbA
Selaginella repanda (Desv. ex Poir.) Spring Zhang X.-C. 5655 (PE) Yunnan, China MH814904 MH814848 MH814876
Selaginella selaginoides (L.) P. Beauv. ex Schrank et Mart. S. Weststrand 104 (UPS) Sweden KY023148.1 - -
Selaginella trichophylla K. H. Shing Zhang et al. 6784 (CDBI, MO, VNMN, PYU) Cao Bang, Vietnam KT161624.1 - -
Selaginella trichophylla K. H. Shing Chu et al. 31925 (PYU) Yunnan, China KT161621.1 - -
Selaginella trichophylla K. H. Shing Chu et al. 29310 (PYU) Yunnan, China KT161622.1 - -
Selaginella trichophylla K. H. Shing Jiang 318 (PYU, CDBI) Hainan, China KT161623.1 - -
Selaginella uncinata (Desv. ex Poir.) Spring Zhang et Zhou DJY04101 (CDBI) Sichuan, China KT161626.1 - -
Selaginella vaginata Spring Zhang Z.-S. 161 (PE) Shaanxi, China MH814907 MH814851 MH814879
Selaginella xipholepis Baker Zhang X.-C. 7422 (PE) Guizhou, China MH814908 MH814852 MH814880
