Li-Bing Zhang

Missouri Botanical Garden · Science & Conservation

A taxonomic revision of Ctenitis (C. Chr.) C. Chr. (Dryopteridaceae) from Africa and the Western Indian Ocean was carried out. Forty species are recognized, of which 12 are described as new, eight are transferred to Ctenitis from other genera, and one is elevated from variety to species and transferred from Aspidium Sw. to Ctenitis. These 21 nomenc...

As the second most genera-rich fern family, Pteridaceae contain more than 1000 species contributing to ca. 10% of extant leptosporangiate fern diversity. The subfamily Pteridoideae is one of the five subfamilies often recognized. The circumscription of Pteridoideae has not been clear. A large number of species have not yet been included in any mole...

The largest genus of seed-free vascular plants Selaginella alone constitutes the family Selaginellaceae, the largest of the lycophyte families. The genus is estimated to contain ca. 800 species distributed on all continents except Antarctica, with the highest species diversity in tropical and subtropical regions. The monophyly of Selaginella has ra...

The circumscription and the phylogeny of the fern family Tectariaceae have been controversial. Previous molecular studies have supported the monophyly of this family, with 4-5 genera. However, these studies were exclusively based on plastid markers and relatively small sampling, especially of the non-Tectaria genera. In the present study, DNA seque...

The pantropical monospecific fern genus Didymochlaena had long been placed in the family Dryopteridaceae before the advent of molecular phylogenetics. Recent molecular studies confirmed that it is a member of eupolypods I, but its familial placement has been controversial. It has been resolved using molecular data either as a member of Hypodematiac...

A cytotaxonomic study is conducted for Lepisorus from eastern, southern, and southwestern China, involving 120 samples of about 26 species in seven sections. The chromosome numbers of 10 species are reported for the first time. The basic chromosome numbers, x = 19 and x = 22 discovered in L. cespitosus and L. lineariformis, respectively, are found...

A new fern species, Whittieria hengduanensis (Ophioglossaceae), from Sichuan, Xizang, and Yunnan, Southwest China (eastern Himalaya), is described and illustrated. This species is similar to W. engelmannii in the Americas in having a cylindrical rhizome and complex-reticulate venation. In addition, both species grow in open habitat on basic soil. H...

The unresolved phylogenetic framework within the Selaginellaceae subfamily Gymnogynoideae (ca. 130 species) has hindered our comprehension of the diversification and evolution of Selaginellaceae, one of the most important lineages in land plant evolution. Here, based on plastid and nuclear data extracted from genomic sequencing of more than 90% spe...

The species diversity of certain groups of ferns in Indo-Burma Biodiversity Hotspot remains largely unexplored using combined molecular and morphological approaches. Here, based on morphology and an updated phylogeny of Leptochilus with 121 (115% increase of the latest sampling) new samples across Asia added, specifically those of three systematica...

The proposal by Wan & al. intended to maintain maximal nomenclatural stability by proposing to conserve Selaginella (Selaginellaceae) with a conserved type, regardless of how and when Selaginella is to be split into several or more genera, as has already been done in some recent studies. Valdespino & al. do not seem to have understood the nomenclat...

Five new combinations in the lycophyte family Selaginellaceae are made and transferred from Selaginella sensu lato. These five names include Boreoselaginella baodongii, B. sajanensis, Chuselaginella neei, C. negrosensis, and Hypopterygiopsis odishana. We also designated the lectotypes of S. neei and S. negrosensis.

The genus Diplazium in Thailand is revised. Thirty-one species are recognized, of which D. thailandicum is new to science, and two species, D. bellum and D. pallidum are new to Thailand. These species are described and illustrated. The lectotypes of D. dilatatum, D. leptophyllum, D. petolotii and D. petrii are designated here. A key to the species,...

Three new species of the shield fern genus, Polystichum sunhangii, P. wusugongii, and P. xinfeniae, were described based on morphological and molecular evidence. The new species were discovered in Medog county, Xizang, at elevations ranging from 1200 to 2300 meters. The differences between each new species and their morphologically similar species...

Three new species, Polystichum pseudodangii, P. xuansonense, and P. unicum, and a new record, P. omeiense, are described and reported here, respectively. These four species were found in limestone habitat in northern Vietnam, with the first species collected from Phu Tho Province and the rest from Ha Giang Province. Detailed descriptions, images, t...

Grammitidoideae are the largest subfamily in Polypodiaceae and contain about 911 species. Progress has been made in understanding the overall phylogeny and generic boundaries in the light of recent molecular works. However, the majority of species, especially Asian species, and some critical type species of genera remain unsampled . In this study,...

Selaginella is the largest and most taxonomically complex genus in lycophytes. The fact that over 750 species are currently treated in a single genus makes Selaginellales/Selaginellaceae unique in pteridophytes. Here we assembled a dataset of six existing and newly sampled plastid and nuclear loci with a total of 684 accessions (74% increase of the...

Intercontinental disjunct distributions can arise either from vicariance, from long-distance dispersal, or through extinction of an ancestral population with a broader distribution. Tectariaceae s.l., a clade of ferns in Polypodiales with ca. 300 species mainly distributed in the tropics and subtropics, provide an excellent opportunity to investiga...

Following results of recent studies, Huperzia myriophyllifolia (Huperziaceae), Lycopodium subinundatum (Lycopodiaceae), and Polystichum pseudodeltodon, P. pseudostenophyllum, P. subapiciflorum, and P. subobliquum (Dryopteridaceae) are recognized/reinstated. A new combination, Pseudolycopodiella subinundata, is made based on L. subinundatum. Spinulu...

Following a recent phylogenetic study, we here review the circumscription of the grammitid fern genus Oreogrammitis (Polypodiaceae: Grammitidoideae). We propose three new genera Calligrammitis, Devolia, and Glabrigrammitis, to accommodate the three clades resolved outside of the core Oreogrammitis. The taxonomic treatment is presented, and the morp...

A new species of Hymenasplenium (Aspleniaceae), H. hovenkampii, from the Lesser Sunda Islands of Indonesia is described. This species was preliminarily identified as H. unilaterale, but can be easily distinguished by its bipinnatisect laminae (vs. 1-pinnate lamina) and reddish brown (vs. dark brown to black) stipes and rachises. In fact, all specie...

The fern genus Didymochlaena is characterized by having scaly rhizomes and petioles, tufted and bipinnate leaves, dimidiate pin-nules, and fewer than 11 sori per pinnule. For a long time only one species was widely recognized in the genus. Most recently, six and seven species from Madagascar and Asia-Pacific regions, respectively, have been recogni...

The pantropical fern genus Didymochlaena (Didymochlaenaceae) has long been considered to contain one species only. Recent studies have resolved this genus/family as either sister to the rest of eupolypods I or as the second branching lineage of eupolypods I, and have shown that this genus is not monospecific, but the exact species diversity is unkn...

We describe a new species of Cyrtomium, C. calcis, from northern Vietnam and southern China, and report new records of six species in two fern genera, Arachniodes and Polystichum, of Dryopteridaceae, i.e., Arachniodes amabilis (from Lamdong Province), A. amoena (Quangtri Province), A. fengii (Nghean Province), A. globisora (Thanhhoa, Quangtri, and...

The papilionoid legume genus Sophora (Fabaceae) exhibits a worldwide distribution, but a phylogenetic framework to understand the evolution of this group is lacking to date. Previous studies have demonstrated that Sophora is not monophyletic and might include Ammodendron, Ammothamnus, and Echinosophora, but the relationships among these four genera...

We reviewed the hybrid genera of ferns and propose three new nothogeneric names, ×Apseudathyrium, ×Glaphyrocyclosorus, and ×Osmunimunda, and four new combinations to accommodate the nothospecies related to these three new nothogenera.

Polystichum suiyangense (Dryopteridaceae), a new species from Suiyang County, Guizhou, China, is described and illustrated. It is a member of P. sect. Sphaenopolystichum and similar to P. auriculum, P. bifidum, and P. martinii, but differs from them in its frond texture, leaf color, scales on petiole and rachis, pinna shape, and pinna apex. The new...

Bolbitis is a pantropical fern genus of Dryopteridaceae with ca. 80 species mainly in tropical Asia. Earlier studies confirmed the monophyly of Bolbitis when Mickelia is excluded and identified three major clades in Bolbitis. However, earlier studies are based on relatively small sampling and the majority of Asian species are not sampled. In this s...

Polystichum gonggashanense (Dryopteridaceae: sect. Hypopeltis) is described and illustrated from Mt. Gongga (Minyar Gonkar), Sichuan, China. It can be distinguished from its most similar species, P. sinense, by the blackish brown petiole scales, the oblong-lanceolate lamina, and narrowly lanceolate rachis scales. The new species is found in four po...

Adder's tongue ferns or Ophioglossaceae are best known among evolutionary biologists and botanists for their highest chromosome count of any known organisms, the presence of sporophores, and simple morphology. Previous studies recovered and strongly supported the monophyly of the family and the two multi-generic subfamilies, Botrychioideae and Ophi...

A new fern species, Polystichum jinpingense, a member of P. subg. Haplopolystichum sect. Haplopolystichum, is described and illustrated from southeastern Yunnan, China. Polystichum jinpingense is most similar to P. subacutidens in plant size and having numerous pinnae, but the former has thinner (0.7–0.9 mm in diam.) rachis, oblong pinnae, pinna ma...

A new species of Ophioglossum, O. isanense, from Northeast Thailand is described and illustrated. It may be diagnosed from other congeners by a combination of morphological characters including: its smaller size, cylindrical rhizome, stoloniferous roots, brownish green common stalk and trophophylls, the presence of persistent leaf sheath at the top...

A new fern species, Athyrium aberrans (Athyriaceae), currently only known from southeastern Xizang, China is described. Morphologically, A. aberrans is superficially similar to A. falcatum in the habit and pinna shape, but the new species can easily be distinguished from the latter by having purplish petioles and rachises, a bulbil on the rachis be...

A cytotaxonomic study was conducted of 38 fern species from China, based on collections from Chongqing, Guangxi, Guizhou, Jiangxi, Sichuan, Xizang, and Yunnan provinces, and their reproductive modes are estimated by counting the spore numbers per sporangium. The chromosome numbers of 11 species are reported here for the first time, and cytotaxonomy...

Lycopodiales are one of the oldest lineages of any living vascular plants and contain about 400 species distributed on all continents except Antarctica but with its highest diversity in tropical regions. Following a recent molecular study, here we classify Lycopodiales into two families: Huperziaceae and Lycopodiaceae, and Lycopodiaceae are further...

As one of the earliest land plant lineages, Selaginella is important for studying land plant evolution. It is the largest genus of lycophytes containing 700–800 species. Some unique characters of Selaginella plastomes have been reported, but based only on 20 species. There have been no plastome phylogenies of Selaginella based on a relatively large...

A new fern species, Hypodematium eglandulosum, is described and illustrated from southern China and Southeast Asia, and a new combination is made, H. chrysolepis, from Africa. Hypodematium eglandulosum is distinguished from its similar species by plants eglandular, leaves 3-pinnatifid, and stipes upward sparsely covered with acicular hairs or nearl...

The Old World fern genera Hypodematium and Leucostegia had long been placed in the families Dryopteridaceae and Davalliaceae, respectively, before the advent of molecular phylogenetics. Recent molecular studies confirmed the recognition of the family Hypodematiaceae composed of these two genera, but the relationships within each of these two genera...

Lycopodiaceae are one of the oldest lineages of any living vascular plants and contain about 400 species distributed on all continents except Antarctica, with its highest diversity in tropical regions. Previous studies on the morphology, anatomy, and molecular systematics of Lycopodiaceae have made substantial progress in understanding the diversit...

Based on morphological and phylogenetic studies, Selaginella qingchengshanensis, a new species in S. sect. Heterostachys from Sichuan, China is described and illustrated. Morphologically, the new species is similar to S. amblyphylla, but has sterile leaves with denticulate to slightly ciliate margins, dorsal leaves with long arista (as long as leav...

We describe a new species of Asplenium (Aspleniaceae) from Vietnam as A. quangbinhense. Morphologically, it is most similar to A. oldhamii. The new species is closely related to A. pseudopraemorsum, and a morphological comparison is presented between the two species. Three species, A. cyrtosorum, A. maguanense, and A. pseudopraemorsum, are reported...

Comprising about 82% of the extant fern species diversity, Polypodiales are generally believed to have diversified in the Late Cretaceous. We estimated the divergence times of Polypodiales using both penalized likelihood and Bayesian methods, based on a dataset consisting of 208 plastomes representing all 28 families and 14 fossil constraints refle...

A taxonomic study of the Hypodematium glabrum complex (Hypodematiaceae) is presented based on morphological evidence. Five species are recognized in this complex, four of which are described as new. The four new species are all from Yunnan, China and include H. brevipilosum sp. nov., H. parvisorum sp. nov, H. subglabrum sp. nov, and H. truncatum sp...

The Asplenium wrightii complex is morphologically variable and difficult in species delimitation. Owing to lack of comprehensive sampling in phylogenetic studies, the taxonomy of this complex remains unresolved. Based on extensive field observations, specimen examination and our recent molecular data, the present study aims to clarify the identitie...

Hypodematium shingii (Hypodematiaceae), a new fern species, is described and illustrated from limestone regions in southern China and Southeast Asia. Hypodematium shingii is different from true H. crenatum in morphology and distribution.

A new fern species, Selliguea wusugongii (Polypodiaceae), is described from southeastern Xizang, China. Morphologically, S. wusugongii is distinguished in the genus by having long need-like fronds with width of the lamina only 0.8–1.1 mm, and is the species with the narrowest laminae in selligueoid ferns. The new species is most similar to some Mal...

Two new fern species, Polystichum dongchuanense and P. menglaense (Dryopteridaceae), are described and illustrated from Yunnan, China. Both are members of P. subg. Haplopolystichum sect. Haplopolystichum. Polystichum dongchuanense is most similar to P. acutidens in plant size and pinna size, but the two can be distinguished from each other by pinna...

A taxonomic revision of Didymochlaena (Didymochlaenaceae) from Asia and the Pacific region is conducted based on morphological and molecular evidence. Seven species are recognized, of which four are described as new and a new status is raised to a species from a variety. These four new species include D. fijiensis from Fiji, D. philippensis from th...

A new fern species, Leptochilus gracilis, is described and illustrated from southeastern Yunnan, China. Leptochilus gracilis is most similar to L. ellipticus. Compared with L. ellipticus, L. gracilis has a smaller habit with fronds less than 20 cm long, laminae with only one or two pairs of lobes, and midrib of lobes at angles of 45–60 degree with...

The fern family Didymochlaenaceae contains the genus Didymochlaena only. This genus is generally believed to comprise one species only, while morphological polymorphism in this species has long been observed. Here we conducted a taxonomic revision of Didymochlaena from Madagascar and recognized six species including two new species, three new statu...

New Guinea is the world’s largest tropical island and has fascinated naturalists for centuries. Home to some of the best-preserved ecosystems on the planet and to intact ecological gradients—from mangroves to tropical alpine grasslands—that are unmatched in the Asia-Pacific region, it is a globally recognized centre of biological and cultural diver...

The biological species concept is not exclusively applicable in many groups of organisms including ferns. Interspecific fern hybrids are not rare: there are 16 intergeneric hybrid genera in ferns confirmed with molecular data. Here we add one more hybrid genus in the tribe Lepisoreae of Polypodiaceae, ×Lepinema, formed via hybridization between par...

A taxonomic study of Hypodematium (Hypodematiaceae) from Japan is presented. Three species are recognized including H. faurei, H. glanduloso-pilosum, and H. ryukyuense (sp. nov.).

A new species, Selaginella coriaceifolia, is described and illustrated from central Vietnam. The new species can be distinguished from other species in Asia by its habit epilithic, main stem nearly isotomously branched, leaves strongly leathery, ventral leaves imbricate on stem and branch, dorsal leaves with reflexed arista at apex, and sporophylls...

A new fern species, Polystichum neoacutidens (Dryopteridaceae), is described and illustrated from a limestone area in northeastern Thailand. It is distinguished from its closest relative, P. acutidens, by the color of petioles and rachises, density of petiole scales, and position of sori.

A new fern of Polystichum (Dryopteridaceae), P. asia-minoris, is described and illustrated from Horma Canyon in the borders of Küre Mountains National Park in Kastamonu Province located in the north of Turkey. Polystichum asia-minoris is distinguishable from the morphologically similar P. obliquum by having pinnae abaxially densely covered with bro...

Lepisoroid ferns (tribe Lepisoreae, Polypodiaceae) are arguably one of the most confusing fern groups in Polypodiaceae in terms of delimitation of genera largely because of their simple morphology. Previous molecular studies either had very small taxon sampling of the non-Lepisorus genera and did not well resolve the relationships among these gener...

The taxonomy of the cliff fern family Woodsiaceae has been controversial with 22–59 species in 1–7 genera recognized in various classifications. This is mainly due to limited taxon sampling (up to 33 accessions representing up to 21 species) and poor resolution in molecular studies so far. In the present study, DNA sequences of five plastid markers...

As an ancient lineage of ferns, Ophioglossaceae are evolutionarily among the most fascinating because they have the highest chromosome count of any known organism as well as the presence of sporophores, subterranean gametophytes, eusporangiate sporangia without annuli, and endophytic fungi. Previous studies have produced conflicting results, identi...

The first chloroplast genome of the fern genus Polystichum Roth (Dryopteridaceae) is reported here. Polystichum deltodon (Baker) Diels belongs to subgenus Haplopolystichum (Polystichum; Dryopteridaceae), many species of which are endangered or critically endangered species. The complete chloroplast genome of P. deltodon was determined for the first...

Ophioderma redactophylla is described and illustrated as a new species from the Malay Peninsula. It is characterized by vermiform underground parts without tuberous rhizomes, the flat stalks, the linear trophophore, and the sporophore surpassing the trophophore. A taxonomic description and illustrations of the new species are presented. A key to sp...

A taxonomic study of the Hymenasplenium unilaterale subclade (Aspleniaceae) is presented based on morphological and molecular evidence. Twelve species are recognized, nine of which are described as new. The nine new species include H. kinabaluense, H. solomonense, H. madagascariense, H. nigricostatum, H. neocaledonicum, H. oligosorum, H. queensland...

Four new species of Hymenasplenium (Aspleniaceae) from Africa and Asia are here described based on both morphological and molecular evidence. These four new species include H. guineense and H. kenyense from mainland Africa and H. sabahense and H. wusugongii from Asia. These new species have been erroneously treated as H. unilaterale by earlier pter...

A new asplenioid fern species, Asplenium simaoense (Aspleniaceae), found growing on an exposed slope of a limestone mountain in southwestern Yunnan, China, is described and illustrated. A preliminary phylogenetic analysis showed that A. simaoense is strongly supported as sister to A. tenuifolium, whereas A. simaoense is morphologically most similar...

Petrocosmea qiruniae M.Q.Han, Li Bing Zhang & Yan Liu (Gesneriaceae), a new species from Guizhou, China, is described and illustrated. It is most similar to P. leiandra (W.T.Wang) Z.J.Qiu in leaf blade shape and corolla type, especially in the colour of corolla throat and spots, indumentum of the stamens and the anther and ovary morphology. It was...

Eight new fern species of Polystichum subg. Haplopolystichum sect. Haplopolystichum (Dryopteridaceae) are described and illustrated from Guizhou Province, southwestern China. These eight species include P. anshunense, P. asperrimum, P. confusum, P. kropfii, P. libingii, P. macrodon, P. perditum, and P. yifanii. Each of these eight species was disco...

The newly defined fern genus Leptochilus contains about 50 species occurring in subtropical to tropical Asia and adjacent Pacific islands. The circumscription and phylogeny of the genus have been ambiguous and its species had been included in various genera such as Colysis, Dendroglossa, Kontumia, Microsorum, and Paraleptochilus. Previous molecular...

The extinct fern genus Coniopteris was a typical component in the Mesozoic flora with a wide distribution in both Northern and Southern Hemispheres from Early Jurassic to Early Cretaceous, and it played a very important role in stratigraphic correlation, while its character evolution and systematic position have both been unsolved mysteries. In thi...

We describe a new species, Woodsia kungiana (Woodsiaceae), based on morphological characters. The new species is described and illustrated from high mountains of western Sichuan, China. The new species is distinguishable from the morphologically similar, closely related Woodsia cycloloba in indumentum and morphology of indusia.

Chlamydogramme Holttum (1986: 157) was described mainly based on the linear sori of its type C. hollrungii (Kuhn in Schumann & Hollrung 1889: 8) Holltum (1986: 157). Later, Holttum (1991) added one more species to this genus, C. elata Holttum (1991: 556). Kramer et al. (1990) pointed out that Chlamydogramme should probably be included in Tectaria C...

A new fern of the genus Arachniodes (Dryopteridaceae), A. hehaii, is described and illustrated from a cave entrance in southwestern China’s Guizhou Province. The new species is similar to A. chinensis, but differs in the color of stipe scales, lamina width and shape, the morphology of lamina apex, and indusium thickness.

Arachniodes (Dryopteridaceae) is one of the most confusing and controversial fern genera in terms of its circumscription, nomenclature, and taxonomy. Estimates of species number range from 40 to 200. Previous molecular works included only 2–17 accessions representing 2–12 species of Arachniodes and allied genera, leaving most of the Asian species r...

Recent molecular work has found that the fern genera Colysis C. Presl (1849: 146), Kontumia S.K. Wu & K.L. Phan (2005: 245), and Paraleptochilus Copeland (1947: 198) should be subsumed under Leptochilus Kaulfuss (1824: 147) (Dong et al. 2008, Kreier et al. 2008, Kim et al. 2012, Zhang et al. in press) so that Leptochilus can be maintained as monoph...

Three new species of the fern genus Arachniodes (Dryopteridaceae) are described from Vietnam: A. daklakensis, A. longicaudata, and A. quangnamensis. The first two occur in Central Highlands and Central Vietnam, respectively, while the last is found in the north (Bavi, Langson and Tamdao) as well as in the central Vietnam (Nghean, Quangbinh, Quangtr...

Following the results of a molecular phylogeny we made nine new combinations in Leptochilus based on morphological, ecological, and distributional evidence. The new combinations include: Leptochilus chilangensis, L. chingii, L. dissimilialatus, L. evrardii, L. flexilobus, L. fluviatilis, L. pentaphyllus, L. poilanei, and L. saxicola.

A new species of Polystichum (subg. Haplopolystichum; Dryopteridaceae), P. recavum, is described from a limestone cave from southern Guangxi, China. Polystichum recavum is most similar to P. cavernicola in having oblanceolate lamina and oblong pinnae, but distinguishable from the latter in number of sterile pinna pairs, shape of pinna apex, and sha...

Eight new fern species of Polystichum subg. Haplopolystichum (Dryopteridaceae) are described and illustrated from Guangdong and Yunnan provinces, southern and southwestern China, respectively. These eight species include P. deltatum, P. gejiuense, P. malipoense, P. oblongipinnarum, P. pingbianense, P. rectum, and P. superum from Yunnan, and P. hanm...

The pantropical genus Tectaria Cav. (Tectariaceae) is one of the largest fern genera. It has been estimated to contain ca. 210 mostly tropical species in Africa, the Americas, Asia, and the Indian Ocean and Pacific Ocean islands. Tectaria had perhaps been the most confusing fern genus in terms of its circumscription and phylogeny. Recent studies ha...

Nine new species of Hymenasplenium (Aspleniaceae) from Asia are here described based on both morphology and our recent molecular phylogenetics of the genus. Of the nine new species, H. chingii, H. denticulatum, H. sinense, H. speluncicola, and H. wangpeishanii are from southern China, whereas H. distans, H. ngheanense, H. phamhoanghoi, and H. quang...

Asplenium serratifolium (Aspleniaceae), a new fern species from central Vietnam, is described and illustrated. The new species is characterized by plants 10-18 cm tall, laminae pinnatipartite, lobe margins entire or with shallow teeth, and veins simple or forked. Molecular phylogenetic analysis based on five plastid markers (atpB, rbcL, rps4, rps4-...

A new fern species, Tectaria moranii (sect. Tectaria; Tectariaceae), is described from central Costa Rica. Tectaria moranii is most similar to T. vivipara in having rachis bulbils, but the two are distinguishable by the habit of rhizome, shape of lamina, petiole and rachis color, proliferability of rachis bulbils, number of pairs of pinnae per lami...