Phylogeny,character evolution,and classification of Selaginellaceae (lycophytes)

Xin-Mo Zhou ,Li-Bing Zhng

a School of Ecology and Environmental Science,Yunnan University,Kunming 650504,Yunnan,China

b Missouri Botanical Garden,4344 Shaw Blvd,St.Louis,Missouri 63110,USA;Chengdu Institute of Biology,Chinese Academy of Sciences,Chengdu 610041,Sichuan,China

Keywords: Generic classification Homoplasy Lycophyte phylogeny Megaspore types Microspores Nuclear 18S and 26S genes

ABSTRACT 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 earlier largest sampling)representing ca.300 species to infer a new phylogeny.The evolution of 10 morphological characters is studied in the new phylogenetic context.Our major results include:(1)the nuclear and plastid phylogenies are congruent with each other and combined analysis well resolved and strongly supported the relationships of all but two major clades;(2)the Sinensis group is resolved as sister to S.subg.Pulviniella with strong support in two of the three analyses;(3)most morphological characters are highly homoplasious but some characters alone or combinations of characters well define the major clades in the family;and (4) an infrafamilial classification of Selaginellaceae is proposed and the currently defined Selaginella s.l. is split into seven subfamilies (corresponding to the current six subgenera+the Sinensis group) and 19 genera (the major diagnosable clades)with nine new species-poor genera.We support the conservation of Selaginella with a new type, S. flabellata,to minimize nomenclatural instability.We provide a key to subfamilies and genera,images illustrating their morphology,their morphological and geographical synopses,a list of constituent species,and necessary new combinations.This new classification will hopefully facilitate communication,promote further studies,and help conservation.

1.Introduction

Selaginella(Selaginellaceae) is the largest genus in lycophytes and known for its special phylogenetic position in vascular plants,its heterospores,its resurrection ability,and its pharmacological value(Banks et al.,2009).Selaginellais among the earliest diverging lineages of vascular plants and diverged from its closest living relative about 383 million years ago (Ma) in the Devonian Period(Klaus et al.,2017).

Following PPG I (2016),Selaginellaalone constitutes the family Selaginellaceae and the order Selaginellales.However,it is unique for an order/family in pteridophytes with 700 or more species(Jermy,1990;Tryon and Lugardon,1991;Zhou and Zhang,2015)to contain only one genus,considering the most recent classifications of pteridophytes,for example,ca.380 species in 17 genera in Lycopodiales (Chen et al.,2022;Zhang and Zhou,2022),the sister order of Selaginellales+Iso¨etales,ca.265 species in 25 genera in Blechnaceae (PPG I,2016;de Gasper et al.,2016;González et al.,2020),ca.1200 species in 37 genera in Thelypteridaceae (Fawcett and Smith,2021),andca.900 species in 42 genera in Polypodiaceae subfam.Grammitidoideae(Yang et al.,2023;Zhou et al.,2023).

In fact,morphological heterogeneity within the broadly definedSelaginellaor Selaginellaceae has been observed for a long time and a number of studies to recognize several genera in Selaginellaceae have been published.SplittingSelaginellainto several genera dates back to Lamark and Mirbel (1803) and Palisot de Beauvois (1804)when they described and recognizedDidiclisP.Beauv.ex Mirb.,DiplostachyumP.Beauv.,GymnogynumP.Beauv.,andStachygynandrumP.Beauv.ex Mirb.Kuntze (1891) transferred a number of species ofSelaginellatoLycopodioidesbut he actually did not split the genus into two,because he recognizedLycopodioidesonly.Börner (1912) separatedHeterophylliumHieron.ex Börner fromSelaginellabased onSelaginella helvetica,although this generic name is a later homonym ofHeterophyllium(Schimp.)Müll.Hal.ex Kindb.and a nomenclatural synonym ofDiplostachyum.Rothmaler(1944) recognized three genera in Selaginellaceae:Didicliswith a single macrosporangium at the base of strobili surrounded by sterile sporophylls,Lycopodioideswith multiple macrosporangia at the base of strobili without any sporophylls,andSelaginellawith uniform and spirally arranged leaves.Kung (1988) dividedSelaginella s.l.intoLycopodioidesandSelaginella s.s.and restricted the latter toSelaginella selaginoides(L.)P.Beauv.ex Schrank&Mart.and its allies that have no rhizophores and have loosely spirally arranged sterile leaves and cylindric strobili(S.subg.Selaginellasensu Zhou and Zhang,2015).Selaginellasensu Kung(1988)has the same circumscription as Rothmaler's (1944).Satou (1997) and Tzvelev(2004) adopted Rothmaler's (1944) and Kung's (1988) classifications.Soják(1992)acceptedSelaginellasensu Rothmaler(1944)and Kung (1988) and splitLycopodioidessensu Kung (1988) into the narrowly definedLycopodioidesandBryodesmaSoják;the latter was based onSelaginella rupestris(L.)Spring and included those species with spirally arranged sterile leaves and decussate sporophylls.Most recently,Weakley (2012) has recognized three genera in the family,Bryodesma,Lycopodioides,andSelaginella s.s.,for the Flora of the Southern and Mid-Atlantic States.Weakley(2022)and Weakley and Southeastern Flora Team (2022) included one more genus in the family,Gymnogynum,in the Flora of the Southeastern United States.

One of the major reasons that the broadly definedSelaginellawas often adopted(e.g.,Jermy,1990;Valdespino,1993;Zhang et al.,2013;PPG I,2016) is that the current type of the genus,S.selaginoides,is a member of a two-species subunit with no rhizophores at all,dramatically different from all other species in the genus.Recognizing multiple genera would result in name changes of all but the two species without rhizophores,although the nameSelaginellamay be proposed for conservation with a conserved type(Turland et al.,2018).

The second major reason that the broadly definedSelaginellawas often adopted is that the phylogenetic relationships within the genus had been largely unclear in spite of earlier phylogenetic studies (Korall et al.,1999;Korall and Kenrick,2002,2004;Arrigo et al.,2013) and the morphological homoplasy had not been adequately assessed in an appropriate phylogenetic context.These have been improved greatly in the past few years.Weststrand and Korall (2016a) and Zhou et al.(2016 available online in 2015)published large-scale phylogenies based on plastid and nuclear markers.Based on plastidrbcLand nuclear ITS data of 394 accessions representingca.200 species (ca.25% of the extant diversity of the genus)ofSelaginella,Zhou et al.(2016)identified six deep-level clades and 20 major clades with strong resolution of the relationships among most of these clades.Zhou and Zhang(2015) thus recognized these six deep-level clades as six subgenera ofSelaginella(S.subg.Selaginella,S.subg.BoreoselaginellaWarb.,S.subg.PulviniellaLi Bing Zhang &X.M.Zhou,“S.subg.EricetorumJermy” (=S.subg.Gymnogynum(P.Beauv.) Weststrand&Korall),S.subg.HeterostachysBaker,andS.subg.Stachygynandrum(P.Beauv.ex Mirb.) Baker) and three of subgenera (S.subg.Ericetorum,S.subg.Heterostachys,andS.subg.Stachygynandrum)were further classified into six,five,and seven sections,respectively.However,some relationships,e.g.,those of theSelaginella sanguinolentaclade,were not strongly supported in Zhou et al.(2016) and the enigmaticSinensisgroup,which had caused poor phylogenetic resolution in the previous study (Korall and Kenrick,2002),was not sampled in their study.Based on plastidrbcLand nuclearpgiCandSQD1data of 340 accessions representing 223 species ofSelaginellaincluding 10 species of theSinensisgroup,Weststrand and Korall(2016a)resolved theSinensisgroup as sister to those taxa (S.subg.Pulviniella,S.subg.Ericetorum,S.subg.Heterostachys,andS.subg.Stachygynandrumsensu Zhou and Zhang (2015)) with rhizophores clearly originated on the ventral side of the stem with moderate support (PP=0.97) in Bayesian inference (BI).The relationships of five out of the seven major clades identified by Weststrand and Korall (2016a) were not resolved.Depending on different DNA markers,the relationships ofS.subg.Boreoselaginella[includingSelaginella nummularifoliaChing,Selaginella rossii(Baker) Warb.,S.sanguinolenta(L.) Spring,etc.)] were resolved differently (Weststrand and Korall,2016a;Zhou et al.,2016).Based on plastid genomes of 26 species ofSelaginella,Zhang et al.(2019) resolvedS.subg.Boreoselaginellaas sister to the superclade with ventral rhizophores with strong support,same as the position β found by Weststrand and Korall(2016a).Zhang et al.(2020,2022) then speculated that high GC content,extensive RNA editing sites,and elevated substitution rate of plastid genomes maybe strongly shook the stability of phylogenetic topology ofSelaginella.A recent phylogenomic analysis based on 59 plastomes failed to well resolve the phylogenetic relationships inSelaginellawith confidence,highlighting the difficulty in resolving the phylogeny and evolution of this particularly important land plant lineage (Zhou et al.,2022).

Morphologically,state of rhizophores,arrangement of vegetative leaves and sporophylls,and spore morphology are traditionally used for the classification ofSelaginella(e.g.,Jermy,1986;Zhou and Zhang,2015;Weststrand and Korall,2016b).However,because of the homoplastic nature of some morphological characteristics,some taxa are very difficult to define and identify clearly (Zhou et al.,2015;Weststrand and Korall,2016b).For example,rosette-forming habit is present in three subgenera:S.subg.Stachygynandrum[e.g.,S.pallescens(C.Presl) Spring),S.subg.Gymnogynum[e.g.,Selaginella lepidophylla(Hook.&Grev.)Spring,S.novoleonensisHieron.),andS.subg.Pulviniella[e.g.,S.pulvinata(Hook.&Grev.) Maxim.,S.tamariscina(P.Beauv.)Spring),and dimorphic sporophylls appear inS.subg.Stachygynandrum[e.g.,S.flagellataSpring,S.moritzianaSpring,S.radiataBaker,S.simplexBaker] andS.subg.Heterostachys[e.g.,S.leptophyllaBaker,S.nipponicaFranch.&Sav.,S.repanda(Desv.ex Poir.)Spring].Similarly,some morphological character states,e.g.,rhizophores borne on dorsal sides of the stem,reticulate surfaces of megaspores,stele with more than two vascular bundles,and diverse sporangial arrangement on the strobili,are also present in more than one section and/or subgenus.

Selaginellais heterosporous.A number of studies on the morphology of megaspores and microspores (especially megaspores) ofSelaginellahave been published (e.g.,Zhou et al.,2015;Valdespino et al.,2015;Valdespino et al.,2018a;Valdespino et al.,2018b;Valdespino,2015,2017a,2020;Bauer et al.,2016;Wang et al.,2018).These studies showed thatSelaginellaholds extremely high diversity in spore morphology.Spore data have provided very important and useful taxonomic information which has been incorporated in the recent classifications of the genus(Zhou and Zhang,2015;Weststrand and Korall,2016a,2016b).In addition to spore ornamentation,the size of both megaspores and microspores also shows great variation inSelaginella.Megaspore sizes range from 150 to 1500 μm and microspore sizes from 15 to 50 μm (Mickel and Hellwig,1969;Zhou et al.,2015;Bauer et al.,2016).Taxonomically,species ofS.subg.Gymnogynumoften have larger(500-1000 μm)megaspores,whereas those ofS.subg.HeterostachysandS.subg.Pulviniellausually have smaller(200-400 μm) megaspores.

Plastome studies (Tsuji et al.,2007;Smith,2009;Wicke et al.,2011;Jansen and Ruhlman 2012;Ruhlman and Jansen 2018;Mower et al.,2019;Xu et al.,2018;Zhang et al.,2019a,Zhang et al.,2019b;Kang et al.,2020;Xiang et al.,2022;Zhou et al.,2022)showed that plastomes of Selaginellaceae and their infrafamilial lineages known so far have a number of unique and diverse features: (a) plastomes of most plant lineages are highly conserved with quadripartite structure composed by a large single copy(LSC),a small single copy (SSC),and two inverted repeats (IRa and IRb),whereas plastomes of Selaginellaceae exhibit DR structure(also can be R,DR,IR,or DR-IR coexisting types) with small to medium repeats existed in SC,and plastome conformations ranged from one to 24(Zhou et al.,2022);(b)plastome sizes in most land plants are 120-160 kb but those in Selaginellaceae are 78-190 kb;(c) a plastome in other vascular plants usually contains approximately 120-130 genes but those of Selaginellaceae contain 36-128 genes;(d)accD/cemA/infA/psaM/rpl20/rpl21/rpl32/rpl33/rps12/rps15/rps16/ycf66/ycf94and most of the rRNA,tRNA,and introns are generally lost or pseudogenetized in Selaginellaceae;(e) GC content in land plant plastomes ranges from 34% to 40%,but plastomes of Selaginellaceae often are extremely GC-rich (>50%);and (f) overall distinctions of plastomes among subgenera even among sections inSelaginellaare much greater than those among orders/families/subfamilies/genera in other vascular plants.

In this study our goals included: (1) better resolving the relationships withinSelaginellawith expanded taxon sampling by including the elusiveSinensisgroup and with expanded character sampling by including two highly conserved nrDNA markers,18S and 26S,to be sampled for the first time forSelaginella;(2) for the first time evaluating the evolution of gross morphology and micromorphology of megaspores and microspores and identifying morphological synapomorphies of various clades in the context of the new phylogeny;and(3) proposing an infrafamilial classification of Selaginellaceae based on new molecular and morphological results.

2.Materials and methods

2.1.Taxon sampling

Silica gel-dried materials and herbarium samples either were collected in the fields or herbaria(CDBI,KUN,MO,and PYU).Taxon sampling for this study consisted of 686 accessions representingca.300 species inSelaginellaincluding all subgenera/sections recognized by Zhou and Zhang (2015) and Weststrand and Korall(2016b).For morphological analysis,291 accessions to representca.275 species ofSelaginellawere included.

2.2.DNA extraction,amplification and sequencing

Total genomic DNA was extracted from silica-dried material or sometimes from herbarium specimens using a TIANGEN plant genomic DNA extraction kit (TIANGEN Biotech.,Beijing,China)following the manufacturers’ protocols.In total six genes,one plastid marker (rbcL),two single-copy nuclear genes (pgiCandSQD1),and three nuclear rDNA markers (18S,5.8S,and 26S),were used in the phylogenetic study.The primer sequences and PCR conditions are listed in Table S1.PCR products were purified and sequenced by TSINGKE Biological Technology (Chengdu,China).Sequencher 4.14(Gene Codes Corp.,Ann Arbor,MI,USA)was used to assemble and edit contiguous sequences.

2.3.Sequence alignment and phylogenetic analysis

Sequences for each marker were initially aligned with MAFFT v.7(Katoh and Standley,2013)and manually adjusted in BioEdit(Hall,1999).The ambiguously aligned areas were deleted.jModeltest 2.1.7 (Darriba et al.,2012) was used to select the best-fitting likelihood model for Bayesian analyses.The Akaike information criterion(Akaike,1974)was used to select among models instead of the hierarchical likelihood ratio test,following Pol (2004) and Posada and Buckley (2004).The best-fitting models and parameter values are provided in Table 2.The alignment has been deposited in FigShare with study number#22730093.

Equally weighted maximum parsimony (MP) jackknife (JK) analyses(Farris et al.,1996)for each locus and the combined sequence data were conducted using PAUP* v.4.0b10 (Swofford,2002),with insertions and deletions coded as missing data,the removal probability set to approximately 37%,and “jac” resampling emulated analyses.One thousand replicates were performed with 10 TBR searches per replicate and a maximum of 100 trees held per TBR search.

For the concatenated analysis of all nucleotide characters,maximum likelihood(ML;Felsenstein,1973)tree searches and ML bootstrapping (BS) were conducted using the web server RAxMLHPC2 on TG v.7.2.8 on CIPRES web server (Miller et al.,2010),with 1000 rapid bootstrap analyses and -m GTRCAT as the default model followed by a search for the best-scoring tree in a single run(Stamatakis et al.,2008).

Bayesian inference (BI) was conducted using MrBayes 3.2.7(Ronquist and Huelsenbeck,2003) on CIPRES (Miller et al.,2010),transition/transversion rate ratio set as a free parameter,and GTR+I+Γ model(lset nst=6 rates=invgamma)for all partitions,all model parameters were unlinked across four partitions (unlink statefreq=[all],revmat=[all],shape=[all],pinvar=[all]),and all partitions were allowed to have different rates(prset applyto=[all]ratepr=variable).Two independent runs,each with four chains(one cold,three heated),were conducted,each beginning with a random tree and sampling one tree every 1000 generations of 10,000,000 generations.Convergence among generations and stationarity were assessed using Tracer v.1.4 (Rambaut and Drummond,2007) and the first 25% of the trees was discarded as burn-in to ensure that stationarity in log-likelihood had been reached.The remaining trees were used to calculate a 50%majorityrule consensus topology and posterior probabilities (PP).

2.4.Morphological studies

Morphological data (habit,sterile leaves,states of rhizophores,strobilus morphology,sporophylls,etc.) were obtained from field observations,herbarium investigations (specimens at CDBI,MO,NY,PYU,SING,and UC),and literature studies.Megaspores and microspores for some representative species were observed and studied using scanning electron microscope (SEM) (QUANTA 200 Scanning Electron Microscope,FEI Co.,USA) at Yunnan University,Kunming,China.

2.5.Character evolution analysis

Based on previous morphological works and our studies of the morphology ofSelaginellaspecies worldwide,10 morphological characters were analyzed: presence of rhizophores (absent,present),positions of rhizophores (dorsal side of stems and/or branches,ventral side of stems or/and branches,uncertain),dimorphism of overall sterile leaves (dimorphic,monomorphic),dimorphism of sterile leaves at the base of main stems(dimorphic,monomorphic),arrangement of sterile leaves [(spiral,four rows,decussate (at least on stem)],number of megasporophylls on strobili (only one to few at base,several on dorsal or/and ventral side),dimorphism of sporophylls on strobili (dimorphic,monomorphic),resupination of strobili with dimorphic sporophylls(resupinate,non-resupinate,N/A),surfaces of megaspores (reticulate,non-reticulate),reticulate megaspores [theBryodesmatype(with prominent and dense muri: Fig.5Q),theGymnogynumtype(with often high and strong or wing-like muri and slightly regular meshes: Fig.5N-Q),theStachygynandrumtype(with more or less open and irregular meshes: Fig.10H,K-N),theTetragonostachyaetype-1(with very fine muri and open meshes:Fig.9M and N),and theTetragonostachyaetype-2 (with fine muri formed by verrucate ornamentation: Fig.9L),N/A].Similar to megaspores,microspores ofSelaginellaalso showed high variation (e.g.,Valdespino et al.,2015;Zhou et al.,2015;Wang et al.,2018),but microspore data were often unavailable and thus were not included in our analysis.Mesquite v.2.75 (Maddison and Maddison,2011) was used to optimize morphological features on the most likely ML tree.The ancestral states were estimated based on parsimony criterion.For most species with multiple samples,only one or two samples of each species were selected on the tree for analysis.

3.Results

3.1.Molecular phylogeny

A total of 1551 sequences including 665 sequences ofrbcL,128 ofpgiC,143 ofSQD1,187 of 18S,284 of 5.8S,and 144 of 26S were used in the study,of which 420 (ca.30% of total sequences) sequences were newly sequenced for this study.An additional 1131 sequences were downloaded from GenBank.Voucher information and Gen-Bank accession numbers for all sequences are listed in Table S1.Characteristics of alignments are summarized in Table 1.The tree topologies using maximum likelihood,Bayesian inference,and maximum parsimony analyses were generally concordant when using the concatenated dataset.

Table 1 Data matrices and tree statistics for each of the dataset.PI chars.include the outgroups.

Table 2 Best-fitting models and parameter values for separate (rbcL,18S,26S,5.8S, pgiC,and SQD1),simultaneous nuclear,and simultaneous nuclear+plastid datasets and in this study.“G”=gamma distribution shape parameter(Yang,1994).“GTR”=general-time-reversible model(Tavaré,1986).“I”=proportion of invariable sites.“pInv”=proportion of invariable sites.“Simultaneous-1”=18S+26S+5.8S + pgiC + SQD1. “Simultaneous-2”=18S+26S+5.8S + pgiC + SQD1 + rbcL.

A comparison of the trees resulting from MPJK analyses of the plastid marker,individual nuclear markers,combined nuclear markers,and combined plastid and nuclear dataset did not identify any well-supported conflicts in MP analyses (MPJK >70%;Mason-Gamer and Kellogg,1996;Zhang and Simmons,2006).The ML tree based on the plastid marker is shown in Fig.S1,that based on the combined nuclear markers is in Fig.S2,that based on combine plastid and nuclear markers is in Fig.S3,and the simplified Fig.S3 is in Fig.1.The results described below were based on combined plastid and nuclear markers.

The monophyly ofSelaginellawas strongly supported (MLBS:100%;MPJK: 100;BIPP: 1.00).WithinSelaginella,seven deep-level clades corresponding toS.subg.Selaginella,S.subg.Boreoselaginella,S.subg.Gymnogynum(“S.subg.Ericetorum” sensu Zhou and Zhang,2015),S.subg.Pulviniella,S.subg.Heterostachys,S.subg.Stachygynandrumsensu Zhou and Zhang (2015),and theSinensisgroup were strongly supported (MLBS: 100%;MPJK ≥99;BIPP: 1.00).

Selaginellasubg.Selaginella,S.subg.Boreoselaginella,andS.subg.Gymnogynumwere resolved as the first,second,and third diverging lineages,respectively,all with strong ML and BI support.Selaginellasubg.Pulviniellawas resolved as sister to theSinensisgroup with strong ML and BI support.They together were sister toS.subg.Heterostachys+S.subg.Stachygynandrum.The MPJK values were generally low for the inter-subgeneric relationships.Within theSinensisgroup,three deep geographically differentiated clades were identified: one with Asian and northern African species,one withSelaginella australiensis,and one with species from Indian Ocean islands.WithinS.subg.Gymnogynum,six major clades corresponding six sections sensu Zhou and Zhang(2015)were strongly supported as monophyletic: sect.Megalosporarumwas strongly supported as sister toS.sect.Myosurus,andS.sect.Lepidophyllaewas strongly supported as sister toS.sect.Homoeophyllae,but other inter-sectional relationships were not strongly supported.WithinS.subg.Heterostachys,five sections defined by Zhou and Zhang(2015)were all strongly supported as monophyletic (MLBS ≥99%;MPJK≥88;BIPP: 0.98) and their relationships were well resolved and strongly supported (MLBS ≥95%;MPJK ≥80;BIPP: 1.00),but same as those in Zhou et al.(2016).WithinS.subg.Stachygynandrum,S.sect.Ascendenteswas resolved as sister to the rest,followed byS.sect.CircinataeandS.sect.Plagiophyllaewhich formed a clade sister to the remaining four sections sensu Zhou and Zhang (2015).

3.2.Morphological evolution

Ten common morphological characters and their states are shown in Fig.2A-J.It took one step for the parsimony reconstruction of rhizophores(Fig.2A);three steps for that of positions of rhizophores (Fig.2B),four steps for that of dimorphism of overall sterile leaves (Fig.2C);20 steps for that of dimorphism of sterile leaves at the base of main stems (Fig.2D);three steps for that of arrangement of sterile leaves(Fig.2E);four steps for that of number of megasporophylls on strobili (Fig.2F);15 steps for that of dimorphism of sporophylls on strobili(Fig.2G);15 steps for that of resupination of strobili with dimorphic sporophylls(Fig.2H);seven steps for that of surfaces of megaspores(Fig.2I);and eight steps for that of reticulate megaspores (Fig.2J).

Fig.2.Morphological characters of Selaginellaceae optimized onto the maximum likelihood tree.A.Presence of rhizophores.B.Position of rhizophores.C.Dimorphism of overall sterile leaves.D.Dimorphism of sterile leaves at the base of main stems.E.Arrangement of sterile leaves.F.Number of megasporophylls on strobili.G.Dimorphism of sporophylls on strobili.H.Resupination of strobili with dimorphic sporophylls.I.Surfaces of megaspores.J.Reticulate ornamentation of megaspores.

4.Discussion

4.1.Major new findings in the phylogeny

With the largest phylogeny so far in terms of the number of species sampled,our study made a number of new findings in comparison with earlier such works(Weststrand and Korall,2016a;Zhou et al.,2016).These new findings might provide foundation for future studies of interspecific relationships and discoveries of hidden diversity.Below we focus on and update the major relationships in the genus.In order to compare our current results with previous ones,we use subgeneric and sectional names in the Zhou and Zhang (2015) classification except theSinensisgroup,which was not treated in our earlier classification,andS.subg.Ericetorumwhich is changed toS.subg.Gymnogynumhere(Weststrand and Korall,2016a).

Overall,the major new findings include: (1) TheSinensisgroup is resolved as sister toS.subg.Pulviniellawith strong support in two of the three analyses;(2)S.sect.CircinataeandS.sect.Plagiophyllaeformed a clade with strong support in two of the three analyses;(3)S.sect.MegalosporarumandS.sect.Myosurusare resolved as sister to each other;(4)S.sect.Ascendentesis resolved as sister to the rest ofS.subg.Stachygynandrumwith strong support;and (5)S.sect.Ascendentesis found to contain African species.We will discuss them in detail below.

4.2.Deep relationships in Selaginella

4.2.1.Selaginella subg.Selaginella (Fig.3)

Fig.3.Morphology of subfamily Selaginoidoideae 圆穗卷柏亚科(Selaginella subg. Selaginella;Zhou and Zhang,2015).A-C,G and L-O.Selaginoides spinulosa (A.Habit;B.Showing plant without rhizophore;C and G.Showing densely and spirally arranged sporophylls.L and M.Spiny or verrucate ornamentation on megaspore surface.N and O.Spiny ornamentation on microspore surface (N.Distal surface in tetrad and O.Proximal surface).D,E,F,and H-K.S.deflexa (A.Habit;D and E.Densely and spirally arranged sporophylls;F.Spirally arranged sterile leaves on stem;H and I.Spiny ornamentation on megaspore surface;J and K.Spiny ornamentation on microspore (J.proximal surface and K.distal surface).

Selaginellasubg.Selaginellais the first diverging lineage inSelaginella.It contains only two species,S.selaginoidesandS.deflexaBrack.Its relationships have been quite consistent (Korall et al.,1999;Korall and Kenrick,2002,2004;Arrigo et al.,2013;Weststrand and Korall,2016a;Zhou et al.,2016).This clade is well characterized by spiral and monomorphic sporophylls and absence of rhizophores (Figs.2A,C,E and 3).

4.2.2.Selaginella subg.Boreoselaginella (Fig.4)

It corresponds to theSanguinolentagroup (Weststrand and Korall,2016a).The resolutions ofSelaginellasubg.Boreoselaginellahave been controversial.Depending on datasets and analysis methods,there have been three resolutions: (1) sister to the remaining taxa with rhizophores [e.g.,BI based onrbcL,SQD1,orrbcL+SQD1: Weststrand and Korall (2016a);BI and ML based onrbcL+ITS: Zhou et al.(2016)];(2) sister to a clade containingS.subg.Pulviniella+S.subg.Heterostachys+S.subg.Stachygynandrumsensu Zhou and Zhang(2016)based onpgiCdata(Weststrand and Korall,2016a)and based on plastome data(Zhou et al.,2022);and (3) sister to the ventral-rhizophore clade with strong support based on plastid gnomes of 26 species ofSelaginellabut only a few species ofSelaginellawere sampled and did not include theSinensisgroup(Zhang et al.,2020).

Morphologically,Selaginellasubg.Boreoselaginellahas large number of inexplicable features inSelaginella,e.g.,nearly monomorphic sterile leaves,rhizophores borne on dorsal side of stems and branches,xerophytic habit,monomorphic sporophylls,larger megaspores,and rugate surface of microspore.It is similar toS.sect.Articulatae,S.sect.Megalosporarum,andS.sect.Myosurusin having large spore size,dimorphic sterile leaves,and rhizophores borne on dorsal side of stems and branches.It is similar toS.sect.Homoeophyllaein having xerophytic habit,rhizophores borne on dorsal side,and rugate surface of microspores (Fig.4L-N).Some species(S.sanguinolentaandS.nummularifolia)with nearly monomorphic sterile leaves are similar toS.sect.Lepidophyllae.However,this subgenus also has some characters (e.g.,strobili with more than one megasporophylls,sterile leaves dimorphic and in four rows)similar toS.subg.HeterostachysandS.subg.Stachygynandrumsensu Zhou and Zhang (2015).In particular,except rhizophores borne on dorsal side and strobili with more than one megasporophyll,this subgenus is strongly similar to some species (e.g.,S.albocinata) of theSinensisgroup in Asia in having xerophytic habit and monomorphic sporophylls,and they usually share same distribution area in Asia.

In our current studySelaginellasubg.Boreoselaginellais well supported as the second diverging lineage in our ML and BIanalyses(second resolution above),although the support value is <50% in our MP analysis(Fig.1).Being the second diverging lineage has so far most frequently supported.

4.2.3.Selaginella subg.Gymnogynum (Fig.5)

We usedSelaginellasubg.Ericetorumin our earlier papers(Zhou and Zhang,2015;Zhou et al.,2016) which turned out to be a synonym ofS.subg.Gymnogynum(Weststrand and Korall,2016a).This subgenus is strongly supported as the third diverging lineage in our current study,consistent with our earlier results(Zhou et al.,2016).It shares a synapomorphy of rhizophores borne on dorsal side of stems;this character state is undetermined in some species inS.sect.Lyallia,one of the six sections in the subgenus (Zhou and Zhang,2015).Reticulate megaspore surfaces are a synapomorphy of this subgenus (see below).Within this subgenus,six wellsupported clades/sections,S.sect.Lyallia,S.sect.Megalosporarum,S.sect.Myosurus,S.sect.Articulatae,S.sect.Homoeophyllae,andS.sect.Lepidophyllaeare well identified.Selaginellasect.HomoeophyllaeandS.sect.Lepidophyllaeare strongly supported as sister to each other (Weststrand and Korall,2016a;Zhou et al.,2016;our Figs.1 and S3).Selaginellasect.MegalosporarumandS.sect.Myosurusare resolved as sister to each other,which was not found in our earlier study(Zhou et al.,2016),but consistent with the results of Weststrand and Korall (2016a).Other internal relationships of this subgenus are poorly resolved as previous studies (Weststrand and Korall,2016a;Zhou et al.,2016) and more studies are needed.

Selaginellasubg.Exaltataesensu Weststrand and Korall (2016b)includes ourS.sect.MegalosporarumandS.sect.Myosurus(Zhou and Zhang,2015).These two sections,not recognized by Weststrand and Korall (2016b),have different geographical distributions (the former in America and the latter in Africa) and specialized gross morphology and microspores (Fig.5).Selaginellasect.Megalosporarumhas microspores with rod-like ornamentation on surfaces,butS.sect.Myosurushas microspores with equatorial ring and verrucous ornamentation on surfaces(Fig.5S vs.5T).In addition,the divergence between them has been dated asca.200 million years ago(Klaus et al.,2017).

4.2.4.The Sinensis group (Fig.6)

Weststrand and Korall (2016a) resolved theSinensisgroup as sister to the rest of theirSelaginellasubg.Stachygynandrum(=Selaginellasubg.Pulviniella+S.subg.Heterostachys+S.subg.Stachygynandrumsensu Zhou and Zhang (2015)).In our current study,theSinensisgroup is for the first time resolved as sister toS.subg.Pulviniellawith relatively strong support in ML and BI analyses,although only 50%support value in MP analysis(Figs.1 and S3).The monophyly of theSinensisgroup is strongly supported.Morphologically,theSinensisgroup can be easily distinguished from other taxa inSelaginellain having rhizophores borne on the ventral side of stems,only one (to a few) megasporophyll at the base of strobili,large megaspores (usually >600 μm in diam.)(Figs.2B,F and 6).Within theSinensisgroup,three well-supported subclades (theSelaginella sinensissubclade,theS.australiensissubclade,and theS.fissidentoidessubclade) are identified,consistent with results by Weststrand and Korall(2016a).These subclades can be easily distinguished from one another by gross morphology,microspores,megaspores,and geographical distributions (see below).Although strobili with only one megasporophyll (megasporangium) at the base are also present inS.subg.Gymnogynum,the latter has rhizophores borne on dorsal side of stems,reticulate ornamentation on megaspore surfaces,and often spiny ornamentation on microspore surfaces (Fig.5).

Recent plastome study has shown that 48 species ofSelaginellasampled had high GC content (>50%) but all three species in theSinensisgroup had～ 30% GC content in plastomes,similar to most vascular plants;theSinensisgroup has the smallest plastomes in land plants except some parasitic plants,and their plastomes do not contain any tRNAs (Zhou et al.,2022).

4.2.5.Selaginella subg.Pulviniella (Fig.7)

Selaginellasubg.Pulviniellawas resolved as sister toS.subg.Heterostachys+S.subg.Stachygynandrumsensu Zhou and Zhang(2015) with moderate support in our earlier study (Zhou et al.,2016) but theSinensisgroup was not sampled.Weststrand and Korall (2016a) sampled theSinensisgroup but only BI analysis was presented.Our current study resolvedS.subg.Pulviniellaas sister to theSinensisgroup with relatively strong support in ML and BI analysis,although only 50% support value in MP analysis(Figs.1 and S3).This resolution has not been found before.Morphologically and ecologically,this subgenus comprises species mostly with rosette-forming habit and extremely drought habitat.This subgenus and theSinensisgroup share drought habitat,but morphological synapomorphies to support their sister relationship are unclear.

4.2.6.Selaginella subg.Heterostachys (Figs.8 and 9)

Consistent with our previous results (Zhou et al.,2016),our current study resolvedSelaginellasubg.Heterostachysas sister toS.subg.Stachygynandrumwith strong support and identified five strongly supported clades within the subgenus corresponding to the five sections proposed by Zhou and Zhang(2015):S.sect.Auriculate,S.sect.Homostachys,S.sect.Heterostachys,Didiclis(“S.sect.Oligomacrosporangiatae”),andS.sect.Tetragonostachyae.Although Weststrand and Korall (2016a,b) did not recognize this subgenus and our sections,their results were consistent with ours with five sections sensu Zhou and Zhang (2015) strongly supported.This subgenus,strongly supported as monophyletic (Weststrand and Korall,2016a;Zhou et al.,2016;our Figs.1 and S3) and characterized by non-reticulate megaspores(Fig.2I),mainly comprises species with dimorphic sporophylls of strobili in Asia and those species with verrucate ornamentation on their microspore surfaces but this reversed to finely reticulate ornamentation (Tetragonostachyaetype-1 and type-2) in some pacific island species ofS.sect.Tetragonostachyae(Fig.2I and J).

4.2.7.Selaginella subg.Stachygynandrum (Fig.10)

This subgenus is also always strongly supported as monophyletic(Korall and Weststrand,2016a;Zhou et al.,2016;our Figs.1 and S3).Reticulate megaspore surfaces are a synapomorphy ofS.subg.Stachygynandrum(see below).In our earlier study,we recognized seven sections/clades in this subgenus:S.sect.Ascendentes,S.sect.Austroamericanae,S.sect.Circinatae,S.sect.Heterophyllae,S.sect.Pallescentes,S.sect.Plagiophyllae,andS.sect.Poceres(Zhou and Zhang,2015) and resolved two paraphyletic clades,S.sect.CircinataeandS.sect.Plagiophyllae,as the first diverging lineages with weak support (Zhou et al.,2016).In our current study,these two clades formed a clade strongly supported as monophyletic in two of the three analyses and as the second diverging lineage in the subgenus with strong support.Selaginellasect.Ascendentesis resolved as sister to the rest of the subgenus.These results are consistent with those in Korall and Weststrand(2016a).Selaginellasect.Ascendenteswas only known to be distributed in Asia and Pacific islands (Zhou et al.,2016),but now an African species (Selaginella versicolor) is also resolved in this clade (Korall and Weststrand,2016a;our Figs.1 and S3).Korall and Weststrand(2016b) recognized this subgenus with a very broad circumscription of 600+species including three of our subgenera:S.subg.Heterostachys,S.subg.Pulviniella,andS.subg.Stachygynandrum,plus theSinensisgroup.They recognized none of our sections in this subgenus either,although five out of our seven sections were strongly supported and the remaining two formed a strongly supported monophyletic clade in their study (our Table 3).

Table 3 A comparison of this classification to two recent classifications[Zhou and Zhang's(2015)and Weststrand and Korall's(2016b)]and Weststrand and Korall's(2016a)phylogeny.

4.3.Morphological evolution in Selaginella

4.3.1.Character 1-Presence of rhizophores (Fig.2A)

Two species ofSelaginella,S.selaginoidesandS.deflexa,have no rhizophores(Fig.3C),and all other species have.These two species are the only member ofS.subg.Selaginella(Zhou and Zhang,2015).The ancestral state of rhizophores inSelaginellais ambiguous because the character state inIso¨etesis unknown or not applicable.Absence of rhizophores can be a synapomorphy or a symplesiomorphy ofS.subg.Selaginella.Presence of rhizophores can also be a synapomorphy or a symplesiomorphy of the rest ofSelaginellaexcludingS.subg.Selaginella(Fig.2A).

4.3.2.Character 2-Position of rhizophores (Fig.2B)

The rhizophores inSelaginellacan be borne on dorsal or ventral side of stems or/and branches.The rhizophores borne on dorsal side are the ancestral state of rhizophores inSelaginella(Fig.2B).The rhizophores ofS.sect.Lyalliaare strictly restricted to the base of erect stems and the dorsal/ventral position of rhizophores is unknown (Fig.5C).The ventrally borne rhizophores evolved once inSelaginellaand are a synapomorphy ofS.subg.Pulviniella+S.subg.Heterostachys(Fig.8D,G)+S.subg.Stachygynandrum+theSinensisgroup(Fig.6B).

Fig.8.Growth forms of subfamily Lycopodioidoideae 异穗卷柏亚科(Selaginella subg.Heterostachys;Zhou and Zhang,2015).A.Hypopterygiopsis bodinieri;B.H.heterostachys;C.H.monospora;D.H.ciliaris;E.H.repanda;F.H.vaginata;G.Lycopodioides nipponica;H.Didiclis bisulcata;I.D.delicatula;J.D.mairei;K.D.siamensis;L.D.willdenowii.

4.3.3.Character 3-Dimorphism of overall sterile leaves (Fig.2C)

The sterile leaves ofSelaginellacan be monomorphic or dimorphic.Although only about 8% of the species have monomorphic sterile leaves inSelaginella,monomorphic sterile leaves have been frequently used to identify and define some taxa,e.g.,S.subg.Selaginella(Fig.3A-D),S.sect.HomoeophyllaeSpring sensu Zhou and Zhang (2015) (Fig.5A,H),andS.subg.Ericetorumsensu Jermy(1986)(Fig.5C,K).Our morphological analysis suggests that the ancestral state of sterile leaves inSelaginellais ambiguous.The widespread dimorphic sterile leaves evolved only once from monomorphic state inSelaginella.Dimorphic sterile leaves evolved to monomorphic leaves once independently each inS.sect.HomoeophyllaeandS.sect.Lyallia.

4.3.4.Character 4-Dimorphism of sterile leaves at the base of main stems (Fig.2D)

Except all species ofSelaginellasubg.SelaginellaandS.sect.Homoeophyllae,and some ofS.sect.Lyalliawith monomorphic sterile leaves throughout plants in Character 3,those species with erect stems and dimorphic sterile leaves on branches can have monomorphic sterile leaves at the base of the main stem.Most species have dimorphic sterile leaves at the base of the main stem(Figs.3-10) inSelaginellaand this dimorphism is a morphological synapomorphy or ancestral state ofSelaginellaexcludingS.subg.Selaginella.Monomorphic sterile leaves at the base of the main stem have evolved at least 18 times from dimorphic sterile leaves independently in the genus.Monomorphic sterile leaves at the base of the main stem(Fig.5C)appear to be a synapomorphy ofS.sect.Lyalliaand it is reversed to dimorphic sterile leaves on lower parts of main stem independently.

4.3.5.Character 5-Arrangement of sterile leaves (Fig.2E)

Arrangement of sterile leaves inSelaginellacan be spiral(Figs.3A,B,D,F and 5A,H),in four rows (Figs.4A-H,5B,D,E,I,6A-H,7A-D,8A-L and 10A-C,F),or decussate (at least on stem;Fig.5C,K).The ancestral state of arrangement of sterile leaves inSelaginellais ambiguous and can be either spiral or in four rows.Sterile leaves in four rows can be a synapomorphy or a symplesiomorphy ofSelaginellaexcludingS.subg.Selaginella.Sterile leaves in four rows evolved to spiral sterile leaves once inS.sect.Homoeophyllae.Decussate sterile leaves evolved from those in four rows and are a synapomorphy ofS.sect.Lyallia.Decussate sterile leaves on branches are restricted toS.sect.Lyallia,while some other species with decussate sterile leaves on main stems have sterile leaves in four rows on branches.We hypothesize that decussate sterile leaves inS.sect.Lyalliaare homologous,but more developmental and anatomical studies are necessary.

4.3.6.Character 6-Number of megasporophylls on strobili(Fig.2F)

Number of megasporophylls on strobili can be only one (to a few) at base of strobili (Figs.5G and 6D-F) or more than two on dorsal side or/and ventral side of strobili(Figs.3G,4E-G,5G,L,7D,9A-J and 10D-F).Having several megasporophylls on strobili is a synapomorphy ofSelaginella s.l.Only one(to a few)megasporophyll on strobili is a morphological synapomorphy of theSinensisgroup.The ancestral state of the clade containing S.sect.Articulatae+S.sect.Megalosporarum+S.sect.Myosurus+S.sect.Lyalliais ambiguous.

4.3.7.Character 7-Dimorphism of sporophylls on strobili(Fig.2G)

Sporophylls on strobili inSelaginellacan be dimorphic(Figs.8B,D,9A-D,F,G,I,and 10D,E) or monomorphic (Figs.3A,B,D,E,G,4E-G,5G,H,J,L,6D,F,7D,9E,H,J and 10F).Traditionally,based on the dimorphic sporophylls inSelaginella,S.subg.Heterostachyssensu Jermy (1986) was widely accepted.However,phylogenetic analyses showed that these species with dimorphic sporophylls inSelaginellaare polyphyletic (Weststrand and Korall,2016a;Zhou et al.,2016;our Figs.1 and S1-S3).Our morphological analysis suggests that monomorphic sporophylls on strobili are the ancestral state ofSelaginella.Dimorphic sporophylls do not appear in other subgenera or theSinensisgroup but only evolved in the two latest-diverging subgenera,S.subg.HeterostachysandS.subg.Stachygynandrum.Dimorphic sporophylls evolved at least 14 times in these two subgenera,but are dominant inS.subg.Heterostachys.The ancestral state of dimorphism of sporophylls on strobili inS.subg.Heterostachysis ambiguous.

4.3.8.Character 8-Resupination of strobili with dimorphic sporophylls (Fig.2H)

Based on the relative position between dimorphic sporophylls and sterile leaves,strobili with dimorphic sporophylls (those taxa with dimorphic sporophylls in character 7) inSelaginellacan be resupinate(Figs.8B,D,9A-D,I and 10D,E)or non-resupinate(Fig.9F and G).Non-resupinate strobili evolved from monomorphic sporophylls and is a synapomorphy ofS.sect.Homostachys.Selaginellasect.Auriculate,the sister ofS.sect.Homostachys,has monomorphic sporophylls.Resupinate strobili evolved once or twice inS.subg.Heterostachysdepending on the ancestral state and evolved at least 10 times inS.subg.Stachygynandrum.The ancestral state of resupination of strobili inS.subg.Heterostachysis ambiguous.

4.3.9.Character 9-Surfaces of megaspores (Fig.2I)

Based on the main elements of ornamentation,surfaces of megaspores inSelaginellacan be reticulate(Figs.5N-R and 9L-N)or non-reticulate (Figs.3H,L,4J-K,6I-L,7E-H,9K,S-V,X).Our morphological analysis suggests that non-reticulate surfaces are the ancestral state of megaspores inSelaginella.Reticulate megaspores in the genus are not homologous but could have evolved only three times inSelaginellaindependently.Reticulate surfaces of megaspores are unambiguously a synapomorphy ofS.subg.Stachygynandrumand a synapomorphy ofS.subg.Gymnogynumas well.The ancestral state ofS.sect.TetragonostachyaeofS.subg.Heterostachysis ambiguous.Reticulate megaspores reversed to non-reticulate megaspores once inS.subg.Stachygynandrumand at least once inS.sect.Tetragonostachyae.Reticulate megaspores inS.sect.Tetragonostachyaecan be a symplesiomorphy.Based on the morphology and size of muri,five types of reticulate ornamentation can be identified (see below).

4.3.10.Character 10-Reticulate megaspores (Fig.2J)

Based on the density,size,and shape of muri of reticulate ornamentation,reticulate megaspores can be divided into theGymnogynumtype (with often high and strong or wing-like muri and slightly regular meshes: Fig.5N-P,R),theBryodesmatype(with prominent and dense muri: Fig.5Q),theStachygynandrumtype(with more or less open and irregular meshes:Fig.10H,K-N),theTetragonostachyaetype-1 (with fine muri and open meshes:Fig.9M-N),and theTetragonostachyaetype-2 (with fine muri formed by connected verrucate ornamentation,Fig.9L).TheStachygynandrumtype of megaspores is a synapomorphy ofS.subg.Stachygynandrum.TheGymnogynumtype of megaspores is shared byS.subg.GymnogynumexcludingS.sect.Homoeophyllae,which supports a close relationship,not resolved by our molecular data,amongS.sect.Articulatae,S.sect.Lyallia,S.sect.Megalosporarum,andS.sect.Myosurus.TheBryodesmatype of megaspores is a synapomorphy ofS.sect.Homoeophyllae(Bryodesma).TheTetragonostachyaetype-1 and -2 of megaspores characterizeS.sect.Tetragonostachyaebut the ancestral state of the section is ambiguous.

Fig.10.Morphology of Chuselaginella 朱氏卷柏属in subfamily Selaginelloideae(Selaginella subg.Stachygynandrum;Zhou and Zhang,2015).A,K,and O.Chuselaginella petelotii(A.Habit;K.Megaspore;O.Microspore);B,H,M,and Q.C.doederleinii (B.Habit;H.Strobili;M.Megaspore;Q.Microspore);C,D,I,and J.C.frondosa (C.Habit;D.Sterile leaves on stem;I.Megaspore;J.Microspore);E.C.intermedia (Habit);F,G,L,and P.C.rolandi-principis (F.Habit;G.Sterile leaves and strobili;L.Megaspore;P.Microspore);N and R,C.trachophylla (N.Megaspore;R.Microspore).

4.4.Splitting Selaginella into seven subfamilies and 19 genera

With our new molecular and morphological results and results from previous studies (Zhou and Zhang,2015;Weststrand and Korall,2016a,2016b;Zhou et al.,2016),it appears that the major monophyletic clades in the currently definedSelaginella s.l.or Selaginellaceae can be morphologically defined by some characters alone or combinations of characters.We propose to recognize these major clades as genera.The question is how many genera should be recognized.Jermy (1986) recognized four subgenera,Zhou and Zhang (2015) accepted six subgenera excluding theSinensisgroup and three subgenera were further classified into six,five,and seven sections,respectively,and Weststrand and Korall (2016b) recognized seven subgenera with 600+species in one subgenus and excluding theSanguinolentagroup.The major differences between the two recent classifications by Zhou and Zhang (2015) and by Weststrand and Korall (2016b) lie mainly in how to treatS.subg.Stachygynandrum.Weststrand and Korall(2016b)combinedS.subg.Stachygynandrumsensu Zhou and Zhang (2015),S.subg.Heterostachyssensu Zhou and Zhang (2015),S.subg.Pulviniella,and theSinensisgroup based on their largely unresolved relationships.We prefer to divide theirS.subg.Stachygynandrumfurther for two reasons: (a) theirS.subg.Stachygynandrumcontains 600+species with heterogeneous morphologies and distributions and is not informative;and (b) the relationships within it are now well resolved and strongly supported(at least in the nuclear phylogeny)in our current study (Figs.1 and S3).

Here we propose to recognize our six earlier subgenera and theSinensisgroup as seven subfamilies and the 19 major clades in our phylogenies as 19 genera in Selaginellaceae(list of subfamilies and genera below)for the following reasons:

(1) Although recognizing 19 genera for 700-800 species in Selaginellaceae/Selaginellales may be viewed as dramatic deviation from the current single-genus classification,there is a history of dividing Selaginellaceae into more than one genus(Palisot de Beauvois,1804;Kuntze,1891;Börner,1912;Rothmaler,1944;Sakurai,1943;Kung,1988;Soják,1992;Satou,1997;Tzvelev,2004;Weakley,2012,2022;Weakley and Southeastern Flora Team,2022).Schuettpelz et al.(2018) argued that there are too few fern genera.Our current classification is also in agreement with recent classifications of pteridophytes,e.g.,25 genera forca.265 species in Blechnaceae (PPG I,2016;de Gasper et al.,2016;González et al.,2020),17 genera forca.380 species in Lycopodiales(Øllgaard,2012;PPG I,2016;Chen et al.,2022;Zhang and Zhou,2022),37 genera forca.1200 species in Thelypteridaceae (Fawcett et al.,2021;Fawcett and Smith,2021),and 42 genera forca.900 species in Polypodiaceae subfam.Grammitidoideae(Yang et al.,2023;Zhou et al.,2023).Many of these recent generic segregates are not necessarily clearer phylogenetically or morphologically than the genera of Selaginellaceae we recognize here.

(2) The relationships in Selaginellaceae,especially those withinSelaginellasubg.Stachygynandrumsensu Weststrand and Korall (2016b),which were not well resolved in Weststrand and Korall (2016a),have been well resolved (Zhou et al.,2016),even with theSinensisgroup sampled (Figs.1 and S3).The genus-level units are well-supported phylogenetically,such that the circumscription of the genera won't need to change in the near future as more phylogenetic information becomes available.

(3) These 19 genera are all deeply diverged lineages in Selaginellaceae and mostly among the oldest lineages in vascular plants.Molecular dating by Klaus et al.(2017)estimated the 15 out of the 19 genera to have diverged from their sisters(stem ages)about 120-373 Ma and the remaining four about 75-80 Ma.The stem ages of the 15 genera are older than any polypod families and genera and the 19 genera we recognize here are older than those of most genera of pteridophytes as estimated by Schuettpelz and Pryer (2009) and Du et al.(2021).Notably,all 42 grammitid fern genera recognized in recent studies diverged from their sisters in the past 30 million years (Sundue et al.,2014;Yang et al.,2023;Zhou et al.,2023).Lu et al.(2020),Zhang et al.(2020),and Zhang and Zhang (2022) used relatively old ages as an argument for generic recognition in Woodsiaceae,Lepisoreae (Polypodiaceae),and Ophioglossaceae,respectively.

(4) Most of the 19 genera and seven subfamilies are morphologically definable,although combinations of morphological characters are sometimes needed (Zhou and Zhang,2015;Weststrand and Korall,2016a,b;Zhou et al.,2016,Figs.2-10;Tables 4 and 5).In fact,as shown below,five out of the seven subfamilies(except Lycopodioidoideae and Selaginelloideae)and 16 out of the 19 genera(exceptDidiclis,Hypopterygiopsis,Selaginella s.s.) can be distinguished from one another using macromorphological features only.Spore features and distribution information are useful in distinguishing Lycopodioidoideae from Selaginelloideae,and are critical in distinguishingDidiclisfromHypopterygiopsisand in distinguishingSelaginella s.s.fromChuselaginellaandKungiselaginella.

(5) Most of the 19 genera are geographically coherent:Afroselaginellais endemic to Africa;Austroselaginellais endemic to Australia;Boreoselaginella,Chuselaginella,andKungiselaginellaoccurs in the Old World;Bryodesmaoccurs in Africa,East Asia,and North America;Didiclisoccurs in Africa,Asia,and Australasia;Ericetorumoccurs in Africa and Australia;Gymnogynumoccurs in North to South Americas,and rarely in Asia and Africa;Hypopterygiopsisoccurs in Asia and Pacific islands and a few in Africa and Madagascar;Koralliais endemic to Madagascar and adjacent islands;Lepidoselaginellais endemic to North America;Lycopodioidesoccurs in Eurasia;Megaloselaginellais endemic to Central to South America;Pulviniellaoccurs in Africa and Asia and rarely in Americas;Selaginella s.s.is nearly endmics to Americas except one or a few species in Africa;Selaginoidesis endemic to the boreal area and Hawaii;Sinoselaginellaoccurs in China extending to the Arabic area and eastern Africa(Ethiopia,Kenya,Somalia);andValdespinoais endemic to North America.

(6) This 19-genus classification is largely in line with our earlier classification (Zhou and Zhang,2015),albeit the taxa have different ranks (Fig.1).We recognized six subgenera and three of them were further divided into five,six,and seven sections,respectively(Zhou and Zhang,2015).Fifteen out of our 21 subgenera/sections are recognized as genera here,whereas ourS.sect.HeterostachysandS.sect.Tetragonostachyaeare combined to formHypopterygiopsis,andS.subg.Stachygynandrumsensu Zhou and Zhang (2015) collectively forms Selaginelloideae.The major difference is that theSinensisgroup was not treated by Zhou and Zhang(2015)and is divided into three genera here.This 19-genus classification is in fact not contradictory with the one proposed by Weststrand and Korall (2016b) in which seven subgenera were recognized.Five of their subgenera,S.subg.Ericetorum,S.subg.Gymnogynum,S.subg.Lepidophyllae,S.subg.Rupestrae,S.subg.Selaginella,are circumscribed exactly the same as the corresponding five of our 19 genera.TheirS.subg.Exaltataeis divided into ourAfroselaginellaandMegaloselaginella.Their unplacedSanguinolentagroup is ourBoreoselaginella.The biggest difference is that we recognize 11 genera in theirS.subg.Stachygynandrum,which formed an unresolved polytomy in their phylogeny (Weststrand and Korall,2016a),but is well resolved in our current study,especially in the nuclear phylogeny (Figs.1 and S1-S3).

(7) Although segregating these 19 clades will cause disruption of a number of species names,in parallel with this paper we have published a proposal to conserveSelaginellawith a new type (S.flabellata(L.) Spring) in the most species-rich major clade inStachygynandrum(Wan et al.,2023),which will largely reduce nomenclatural instability.In fact,most American species belong to the newly definedSelaginella s.s.The current type ofSelaginella,S.selaginoides,is resolved in theSelaginoidesclade which contains only two species.There is already a generic name available,Selaginoides,for this clade.In addition to the namesSelaginellaandSelaginoides,five generic names,Bryodesma,Didiclis,Gymnogynum,Hypopterygiopsis,andLycopodioides,are available and a number of species have already been transferred to these genera(Kuntze,1891;Sakurai,1943;Rothmaler,1944;Kung,1988;Soják,1992;Satou,1997;Tzvelev,2004;Weakley,2022).Three generic names,Boreoselaginella,Ericetorum,andPulviniella,are raised from existing subgeneric names(Warburg,1900;Jermy,1986;Zhou and Zhang,2015;Weststrand and Korall,2016b).Therefore,only nine new generic names are needed and they mostly contain only one to a few species and will not cause mass disruption of names:AfroselaginellaforS.sect.Myosuruswith about four species,AustroselaginellaforS.australiensisand additional three species,ChuselaginellaforS.sect.Ascendenteswithca.70 species,KungiselaginellaforS.sect.CircinataeandS.sect.Plagiophyllaewith about two dozen species,LepidoselaginellaforS.sect.Lepidophyllaewith about three species,MegaloselaginellaforS.sect.Megalosporarumwith about two species,KoralliaforS.sect.Fissidentoideswith about 15 species,andSinoselaginellafor portion of theSinensisgroup with five species,andValdespinoaforS.sect.Auriculatewith about one species.

(8) SubdividingSelaginella s.l.into homogeneous and manageable genera is good for the purposes of analysis,communication,and conservation.It will be easier for botanists to undertake monographic work,for example,on the subunits,and to identify and communicate meaningful phylogenetic/evolutionary changes.TreatingSelaginellaas a single genus devalues it as a triage level for judging biodiversity value and ignores the ancient diversity of its deeply diverged clades.It has been argued that there are too few genera in pteridophytes,which does not foster more precise and efficient communication,promote additional research,or facilitate herbarium curation(Schuettpelz et al.,2018).

In a very parallel case,ThelypterisSchmidel was reduced to a tiny(2 species) genus,with all other species being placed in “new”genera (Fawcett et al.,2021;Fawcett and Smith,2021).The same could be done inSelaginella,but we prefer to conserveSelaginellawith a new type to maximally stabilize the names (Wan et al.,2023).Thelypterishad been to a much greater degree already treated and split into many genera,whereas,despite some movement over the centuries towards splittingSelaginella,much less had been done and segregates had been much less discussed,used,combinations made,inSelaginellauntil the modern era of gene sequencing and phylogenetic trees.

Treating these 19 genera at the rank of subgenus or section withinSelaginellais an alternative (Jermy,1986,1990;Zhou and Zhang,2015;Weststrand and Korall,2016b),but those ranks are almost always ignored except by professional plant systematists,and there is nothing like the name of a species (incorporating the genus) to emphasize less relationship (different genus names) or more relationship (same genus name;A.Weakley,pers.comm.).

It will be more inconvenient for today's botanists to adopt a dramatically different classification than to use the existing one with all 750+species together in one genus,but in the long run the next-generation botanists will find it informative to distinguish the deeply diverged clades at the generic rank because of their molecular,morphological,and geographical distinctiveness.Treating all 750+species in a single genus is comparable to lumping all then 33 genera of Polypodiaceae subfam.Grammitidodeae intoGrammitisSw.or lumping all 24 genera of cheilanthoid ferns intoHemionitisL.(Pteridaceae) as done by Christenhusz et al.(2018).

5.Taxonomic treatment

Our current phylogeny based on the largest taxon and character(nuclear and plastid) sampling corroborated earlier findings(Weststrand and Korall,2016a;Zhou et al.,2016)thatSelaginellais well resolved into a number of major clades.Our morphological analysis of 10 important characters showed that the major phylogenetic clades can be defined morphologically (Weststrand and Korall,2016a;our Fig.2).Our earlier classification (Zhou and Zhang,2015) recognized six subgenera and three of them further into five,six,and seven sections,respectively,excluding theSinensisgroup.Our new results confirm the monophyly of all but two sections (the relationships betweenS.sect.CircinataeandS.sect.Plagiophyllaeare not well resolved but they together are monophyletic).Considering all evidence available,earlier classifications ofSelaginella(Mirbel(in Lamark and Mirbel,1803);Kuntze,1891;Walton and Alston,1938;Sakurai,1943;Rothmaler,1944;Jermy,1986,1990;Kung,1988;Soják,1992;Satou,1997;Tzvelev,2004;Weakley,2012,2022;Zhou and Zhang,2015;Weststrand and Korall,2016b),and recent trend of pteridophyte classifications,we here propose to split the currently definedSelaginella s.l.into seven subfamilies corresponding our earlier six subgenera(Zhou and Zhang,2015)and theSinensisgroup,three of which are monogeneric and four of which are further divided into 3,3,4,and 6 genera,respectively.The 7 subfamilies and 19 genera include:Boreoselaginelloideae (Boreoselaginella),Gymnogynoideae (Afroselaginella,Bryodesma,Ericetorum,Gymnogynum,Lepidoselaginella,Megaloselaginella),Lycopodioidoideae (Didiclis,Hypopterygiopsis,Lycopodioides,Valdespinoa),Pulvinielloideae (Pulviniella),Selaginelloideae (Chuselaginella,Kungiselaginella,Selaginoides s.s.),Selaginoidoideae (Selaginoides),and Sinoselaginelloideae (Austroselaginella,Korallia,Sinoselaginella).A comparison of our current and earlier classifications is listed in Table 3 and Figs.1 and S1-S3.

We provide a key to subfamilies and genera below.For each subfamily and genus,we provide morphological,phylogenetic,and geographical synopses,constituent species,and the necessary new combinations.For these,we provide basionyms and other recent combinations.For additional synonymy see Tryon (1955),Kung(1988),Soják (1992),Valdespino (1993),Jermy and Holmes(1998),Mickel and Smith (2004),Chu (2006),Roux (2009),Zhang et al.(2013),Fraser-Jenkins et al.(2015,2017),Valdespino and Zimmer (2016),and Hassler (2022).We list all species of each genus currently known to us.Species lacking sufficient evidence are listed at the end of “Taxonomic treatment” without combinations pending further studies.We provide Chinese vernacular names for all subfamilies,genera,and sections in the family and for all species occurring in China.

5.1.Key to subfamilies and genera of Selaginellaceae

1.Rhizophore absent;sterile leaves loosely spirally arranged;strobili cylindric (circumboreal areas and Hawaii;2n=18)[subfam.I.Selaginoidoideae圆穗卷柏亚科] ................................................................................................Selaginoides圆穗卷柏属

1.Rhizophore present;sterile leaves densely and spirally,in four rows,or decussately arranged;strobili tetragonal or dorsiventrally complanate (2n=16-60) ..................................2

2.Rhizophores borne on dorsal side of stems and/or branches;if rhizophores strictly restricted to the base of stem,sterile leaves at least decussately arranged on stem or throughout plants ......................................................................................................3

2.Rhizophores borne on ventral side of stems and/or branches;if rhizophores strictly restricted to the base of stem,sterile leaves on stem never decussately arranged..............................9

3.Plants xerophytic,non-rosette-forming,creeping;sterile leaves in 4 rows,strobili with many megasporangia on dorsal and/or ventral side;megaspore surfaces densely contiguoustuberculate,non-reticulate [subfam.II.Boreoselaginelloideae北方卷柏亚科]....................Boreoselaginella北方卷柏属

3.Plants non-xerophytic(if xerophytic,plants rosette-forming or with spirally monomorphic leaves on sterile stem and branch),erect,suberect,ascending,scandent,or creeping (if creeping,plants with monomorphic leaves throughout the plant or strobili with only one megasporangium at base);megaspore surfaces reticulate(theGymnogynumtype and theBryodesmatype)[subfam.III.Gymnogynoideae关节卷柏亚科]................4

4.Sterile leaves strictly monomorphic throughout the plant or at least on stem....................................................................................5

4.Sterile leaves dimorphic or slightly monomorphic throughout the plant.........................................................................6

5.Sterile leaves monomorphic and spirally arranged throughout the plant..............................Bryodesma同形卷柏属

5.Sterile leaves monomorphic and decussately arranged throughout the plant or at least at the base of stems..........................................................................................Ericetorum对叶卷柏属

6.Plants rosette-forming .............Lepidoselaginella鳞叶卷柏属

6.Plants non-rosette-forming ............................................................7

7.Stems inarticulate;microspores with an equatorial flange and surfaces verrucate ..................Afroselaginella非洲卷柏属

7.Stems often articulate;microspores without an equatorial flange and surfaces echinate or pillared.....................................8

8.Plants extremely large (up to 1 m tall);stem’s vascular system actino-plectostele;megasporesca.1.5 mm;microspore surfaces pillared........................Megaloselaginella大孢卷柏属

8.Plants normally shorter than 80 cm;stem's vascular system haplostele,actinostele,or plectostele;megaspores smaller than 1 mm;microspore surfaces often echinate...............................................................................................Gymnogynum关节卷柏属

9.Plants xerophytic;often forming rosettes,rarely stems erect,rhizophores strictly restricted to the base of stem and stems tufted and curling inward when dry;megaspore surfaces coarse without clear ornamentation;microspores spherical,surface scabrate or verrucate[subfam.V.Pulvinielloideae垫状卷柏亚科] ..................................................Pulviniella垫状卷柏属

9.Plants growing in wet habitats or rarely slightly xerophytic;stems erect,suberect,ascending,creeping,or scandent,rarely rosette-forming but megaspore surfaces reticulate,and microspores often hemispherical.......................................10

10.Sporophylls monomorphic [if dimorphic,then megaspore surfaces with typical reticulate ornamentation(only in Americas)];megaspore surfaces reticulate (theStachygynandrumtype) [if non-reticulate,megaspores larger than 500 μm;Sinoselaginelloideae];microspore surfaces often verrucate to baculate on distal surfaces(Old and New World).......................16

10.Sporophylls dimorphic [if monomorphic,megaspore surfaces verrucate,low-rugate,tuberculate,finely reticulate(or fenestrate)],megaspore surfaces often non-reticulate [if reticulate,megaspores are of theTetragonostachyaetype-1&-2];microspore surfaces verrucate,blunt spiny,lamellate,cristate,plain but with globules or spherules,or rarely baculate on distal surfaces (Old World,onlyDidiclis hoffmanniiin the New World][subfam.VI.Lycopodioidoideae异穗卷柏亚科]..........................................................................................11

11.Sporophylls dimorphic,non-resupinate or resupinate (if monomorphic,strobili strongly loose and not distinct from sterile parts)........................................................................................12

11.Sporophylls monomorphic or submonomorphic,and strobili distinct from sterile parts................................................................14

12.Strobili non-resupinate (sporophylls dimorphic) or rarely slightly lax (monomorphic);sporophylls loosely arranged...................................................................Lycopodioides疏穗卷柏属

12.Strobili resupinate (sporophylls dimorphic);sporophylls densely arranged..............................................................................13

13.Dorsal leaves obovate...............................................................................................................Didiclis(theD.bisulcatagroup)瘤孢卷柏属

13.Dorsal leaves non-obovate........Hypopterygiopsis异穗卷柏属

14.Plants small,ca.10 cm,strictly creeping forming loose mats......................................................................Valdespinoa韦氏卷柏属

14.Plants medium-to large-sized,more than 20 cm,erect,scandent,rarely creeping,not forming loose mats................15

15.Megaspore surfaces tuberculate and usually interconnected to form reticula-like structure;microspore surfaces blunt spiny,laminate,or cristate............................Didiclis瘤孢卷柏属

15.Megaspore surfaces finely reticulate(Tetragonostachyaetype-1 and -2);microspore surfaces low-verrucate or smooth with irregular spherules............Hypopterygiopsis异穗卷柏属

16.Strobili with several megasporophylls (sporangia) on the ventral and/or dorsal side;megaspores smaller than 400 μm,surfaces reticulate (theStachygynandrumtype)[subfamily VII.Selaginelloideae同穗卷柏亚科]......................17

16.Strobili with only one (to a few) megasporophyll (sporangium) at the base;megaspores larger than 500 μm,surfaces non-reticulate[subfamily IV.Sinoselaginelloideae中华卷柏亚科].......................................................................................................19

17.Microspore surfaces coarse,baculate,blunt spiny,papillate,or verrucate,rarely ridged (Americas,exceptS.cathedrifoliain Africa)...............................................Selaginella s.s.同穗卷柏属

17.Microspore surfaces often baculate or with blunt spines(Old World)...................................................................................................18

18.Plants often ascending to suberect,a few strictly erect;sides of veins of ventral leaves with two light-colored bands;megaspores without a zonal structure on proximal surface; microspore surfaces blunt spiny (Asia)..............................................................Kungiselaginella孔氏卷柏属

18.Plants often strictly erect,a few creeping;sides of veins of ventral leaves without light-colored bands;megaspores with a zonal structure on proximal surface;microspore surfaces baculate with extending tips (Asia,Pacific islands to Africa)..................................................................Chuselaginella朱氏卷柏属

19.Plants xerophytic;microspore surfaces prominently tuberculate (East and South Asia extending to Arabic area and northern Africa).................................Sinoselaginella中华卷柏属

19.Plants non-xerophytic;microspore surfaces verrucate or coarse(Australia,Madagascar and adjacent islands)..........20

20.Ventral leaves usually auriculate at basiscopic and acroscopic bases;microspores with three holes on proximal surfaces (Madagascar and adjacent islands)..........................................................................Korallia科氏卷柏属

20.Ventral leaves non-auriculate at base;microspore surfaces without holes on proximal surfaces (Australia)............................................................Austroselaginella澳洲卷柏属

5.2.Subfamily I.Selaginoidoideae

Subfamily I.SelaginoidoideaeLi Bing Zhang &X.M.Zhou,subfam.nov.圆穗卷柏亚科(新拟) -Type:SelaginoidesSég.

Plants erect,with no rhizophores;sterile leaves monomorphic and spirally arranged;sporophylls monomorphic;megaspore surfaces blunt echinate to tuberculate;microspore surfaces with echinate(Fig.3);2n=18 (Takamiya,1993).

This subfamily corresponds toSelaginellasubg.Selaginellasensu Zhou and Zhang (2015) and Weststrand and Korall (2016b).

It contains one genus:Selaginoides,occurring in circumboreal areas and Hawaii.

5.2.1.Selaginoides

SelaginoidesSég.,Pl.Veron.3: 51.1754.圆穗卷柏属(新拟) ≡MirmauAdans.,Fam.Pl.(Adanson) 2: 491.(1763) ≡PolycoccaHill,Gener.Nat.Hist.,ed.2,2(Hist.Pl.): 116 (1773)-Type:Lycopodium selaginoidesL.≡Selaginella selaginoides(L.).P.Beauv.ex Schrank &Mart.,Hort.Reg.Monac.3(1829)=Selaginoides spinulosa(A.Braun ex Döll)Li Bing Zhang&X.M.Zhou ≡Selaginella spinulosaA.Braun ex Döll,Rhein.Fl.: 38 (1843) ≡Lycopodina spinulosa(A.Braun ex Döll)Bubani,Fl.Pyren.4: 445 (1901)=Selaginellasubg.HomoeophyllumHieron.&Sadeb.,Engler&Prantl,Nat.Pflanzenfam.l(4):669.1902-Type:Selaginella selaginoides(L.)P.Beauv.ex Mart.&Schrank.

Selaginoidesis resurrected here.Selaginellahas been proposed to be conserved with a conserved type,Selaginella flabellata(L.)Spring(Wan et al.,2023;also see below).Selaginella spinosaP.Beauv.(Prodr.Aethéogam.:112.1805)is an illegitimate name-the epithet‘selaginoides’ (fromLycopodium selaginoidesL.,Sp.Pl.: 1101.1753)was available for use by Palisot de Beauvois inSelaginella.The epithet ‘selaginoides’ fromL.selaginoideswould create a tautonym inSelaginoides(Art.23.4 of the ICN).The earliest available epithet inSelaginoidesis ‘spinulosa’.

Selaginoidesis the earliest diverging lineage in Selaginellaceae(Korall et al.,1999;Korall and Kenrick,2004;Zhou et al.,2016,2022;Weststrand and Korall,2016a).

Morphologically,this genus differs from the rest of species in the family in having no rhizophores and monomorphic and spirally arranged leaves (Fig.2A,E).

The genus contains two species:Selaginoides spinulosa(circumboreal area) andS.deflexa(Hawaii) (Fig.3).

Members:

Selaginoides deflexa(Brack.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella deflexaBrack.,U.S.Expl.Exped.,Filic.16:332,no.3,t.45,f.3 (1854) ≡Lycopodioides deflexa(Brack.) Kuntze,Rev.Gen.Pl.1: 826 (1891).

Selaginoides spinulosa(A.Braun ex Döll) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella spinulosaA.Braun ex Döll,Rhein.Fl.: 38 (1843) ≡L.spinulosa(A.Braun ex Döll) Bubani,Fl.Pyren.4: 445 (1901).

5.3.Subfamily II.Boreoselaginelloideae

Subfamily II.BoreoselaginelloideaeLi Bing Zhang&X.M.Zhou,subfam.nov.北方卷柏亚科(新拟) -Type:Boreoselaginella(Warb.)Li Bing Zhang &X.M.Zhou.

Plants xerophytic,creeping (Fig.4A-C);sterile leaves in four rows,nearly monomorphic(Fig.4D)or dimorphic(Fig.4H),axillary leaves present;rhizophores borne on dorsal side of stem(Fig.4D);sporophylls monomorphic;megaspore surfaces with dense contiguous tubercles;microspore surfaces rugate to verrucate;2n=30 (Takamiya,1993).

This subfamily corresponds toSelaginellasubg.Boreoselaginellasensu Zhou and Zhang(2015).It is likely the second earlier diverging lineage in the family(Zhou et al.,2016;our Figs.1 and S3).

Plastome conformation of Boreoselaginelloideae known so far(Zhou et al.,2022)was DR structure with only one conformation.In addition,these genes (psaM/rpl20/rpl21/rpl33) tend to be lost or pseudogenized,but are preserved in other taxa of Selaginellaceae.Furthermore,genes (psaM/trnI-CAU/trnP-GGG/trnS-GCU/trnS-UGA)only existed in Boreoselaginelloideae.ExceptndhE,allndhgenes were lost or pseudogenized in Boreoselaginelloideae (Zhang et al.,2022;Zhou et al.,2022).

It contains one genus:Boreoselaginella,occurring from eastern to western Asia,and extending to Russia.

5.3.1.Boreoselaginella

Boreoselaginella(Warb.)Li Bing Zhang&X.M.Zhou,stat.nov.北方卷柏属(新拟) -Basionym:Selaginellasubg.BoreoselaginellaWarb.,Monsunia 1:100.1900-Lectotype(designated by Zhou and Zhang,2015: 1129):Boreoselaginella borealis(Kaulf.) Li Bing Zhang&X.M.Zhou ≡Selaginella borealis(Kaulf.) Spring (=Selaginella sanguinolenta(L.) Spring).

Boreoselaginellahas rhizophores borne on the dorsal side of the stem,monomorphic and spirally arranged sterile leaves throughout,and non-reticulate megaspores.This combination of the morphology is unique in the family.

It contains about six species distributed from eastern to western Asia and extending to Russia.

Members:

Boreoselaginella aitchisonii(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella aitchisoniiHieron.,Engl.&Prantl,Nat.Pfl.1(4)674(1902)≡Lycopodioides aitchisonii(Hieron.)Tzvelev,Novosti Sist.Vyssh.Rast.36: 25 (2004) ≡Selaginella sanguinolenta f.aitchisonii(Hieron.)Alston,Proc.Nat.Inst.Sci.India 11:215 (1945).

Boreoselaginella borealis(Kaulf.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium borealeKaulf.,Enum.Fil.17-18(1842) ≡Selaginella borealis(Kaulf.) Spring,Bull.Acad.Roy.Soc.Bruxelles 10(1):141,no.40(1843)≡Lycopodioides borealis(Kaulf.)Kuntze,Rev.Gen.Pl.1: 826 (1891).北方卷柏(新拟)

Boreoselaginella jacquemontii(Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella jacquemontiiSpring,Bull.Acad.Roy.Soc.Bruxelles 10: 226,no.104 (1843).

Boreoselaginella nummularifolia(Ching)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella nummularifoliaChing,Fl.Xizangica 1: 21 (1983).钱叶北方卷柏(新拟)

Boreoselaginella rossii(Baker) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella mongholicavar.rossiiBaker,J.Bot.21: 45 (1883) ≡Selaginella rossii(Baker) Warb.,Monsunia 1: 101(1900) ≡Lycopodioides rossii(Baker) Tzvelev,Novosti Sist.Vyssh.Rast.36: 25 (2004).鹿角北方卷柏(新拟)

Boreoselaginella sanguinolenta(L.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Lycopodium sanguinolentumL.,Sp.Pl.2:1100(1753) ≡Stachygynandrum sanguinolentum(L.) P.Beauv.,Prodr.aethéogam.114 (1805) ≡Selaginella sanguinolenta(L.) Spring,Bull.Acad.Roy.Soc.Bruxelles 10:135,no.2(1843)≡Lycopodioides sanguinolenta(L.)Kuntze,Rev.Gen.Pl.1:827(1891).红枝北方卷柏(新拟)

5.4.Subfamily III.Gymnogynoideae

Subfamily III.GymnogynoideaeLi Bing Zhang &X.M.Zhou,subfam.nov.关节卷柏亚科(新拟)-Type:GymnogynumP.Beauv.

Plants erect,ascending,or creeping,rarely scandent or rosette-forming (Fig.5A-E),with rhizophores borne on dorsal side of stems and/or branches (Fig.5F,H-L);a few species with rhizophores strictly restricted to the base of stem(Fig.5C);sterile leaves monomorphic and decussately arranged at least in lower portion of plants (Ericetorum) (Fig.5C,K);sporophylls monomorphic,with only one megasporophyll at the base of strobili (Fig.5J);reticulate megaspore surfaces with wing-like and highly convoluted laesurae forming a complex mass close to the pole (Ericetorum) (Fig.5P);megaspore surfaces reticulate (mainlyGymnogynumtype with high or winglike muri and often closed meshes) (Fig.5N-R);microspore surfaces verrucate to rugate or echinate (Fig.5S-W;Zhou et al.,2015);2n=18,20 (Jermy et al.,1967;Takamiya,1993;Marcon,2003).

The plastome conformations of Gymnogynoideae known so far(Zhou et al.,2022) have all four types of structures (IR,DR,IR-DR coexisting,and NR).In addition,genesndhandmatKtend to lose or pseudogenize except inGymnogynum.Therpl14(except pseudogenetization inLepidoselaginella) andycf3 intron2 (except preserved inMegaloselaginella) have been lost in all genera.

This subfamily corresponds to “Selaginellasubg.Ericetorum”(=S.subg.Gymnogynum)sensu Zhou and Zhang(2015).Weststrand and Korall (2016b) recognized five subgenera in this clade.

It contains about 130 species in six genera,Afroselaginella,Bryodesma,Ericetorum,Gymnogynum,Lepidoselaginella,andMegaloselaginella,occurring in Africa,Australasia,the Caribbeans,and Americas.

5.4.1.Afroselaginella

AfroselaginellaLi Bing Zhang &X.M.Zhou,gen.nov.非洲卷柏属(新拟) -Type:Afroselaginella myosurus(Sw.) Li Bing Zhang &X.M.Zhou [Lycopodium myosurusSw.;Selaginella myosurus(Sw.)Alston].

=Selaginellasect.MyosurusLi Bing Zhang&X.M.Zhou,Taxon 64(6): 1133(2015) -Type:Selaginella myosurus(Sw.) Alston.

Etymology: “Afro-” derived fromAfrica,referring to the African distribution of the genus.

Plants creeping,stems inarticulate,with a single large megasporangium per strobilus,reticulate megaspores with extremely wide and high muri(Fig.5N),microspores with an equatorial ring(Fig.5S).

Afroselaginellahas the smallest plastome size (ca.100 kb) in Gymnogynoideae known so far (Zhou et al.,2022).Plastome structure ofAfroselaginellahas only SC region (single copy region)but lacking repeat region and has one ribosomal operon (Zhou et al.,2022).

Afroselaginellacircumscribed here corresponds toSelaginellasect.Myosurussensu Zhou and Zhang (2015) and part ofS.subg.Exaltataesensu Weststrand and Korall (2016b).

It contains about four species in Africa.

Members:

Afroselaginella chevalieri(Hieron.ex Bonap.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella chevalieriHieron.ex Bonap.,Notes pterid.1: 124(1915).

Afroselaginella congoensis(Alston)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella congoensisAlston,Mém.Inst.Fr.Afr.Noire 50: 30,t.6,f.1-8 (1957).

Afroselaginella myosurus(Sw.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium myosurusSw.,Schrad.,J.Bot.(1800)(2):118(1801)≡Selaginella myosurus(Sw.)Alston,J.Bot.70:64,no.6 (1932).J.Bot.70:64,no.6 (1932).

Afroselaginella volubilis(Alston) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella volubilisAlston,Bol.Soc.Broter.Ser.2,30: 25 (1956).

5.4.2.Bryodesma

BryodesmaSoják,Preslia 64(2): 154(1993) 同形卷柏属(新拟)-Type:Bryodesma rupestre(L.) Soják,Preslia 64(2): 155 (1993) ≡Lycopodium rupestreL.,Sp.Pl.2:1101(1753)≡Selaginella rupestris(L.) Spring,Flora 21(1): 182.1838.

=Selaginellasect.HomoeophyllaeSpring in Martius,Fl.Bras.1(2): 118 (1840) ≡Selaginellasect.HomotropaeA.Broun,App.Ind.Sem.Hort.Berol.11.([1857]1858)-Type:Selaginella rupestris(L.)Spring.

=Selaginellasubg.TetragonostachysJermy,Fern Gaz.13: 118.1986,non Hook.&Grev.in Bot.Misc.2: 382 (1831) -Type:Selaginella rupestris(L.) Spring.

Bryodesmacorresponds toSelaginellasect.Homoeophyllaesensu Zhou and Zhang(2015)and toS.subg.Rupestraesensu Weststrand and Korall (2016b),and approximates “S.subg.Tetragonostachys”sensu Jermy (1986).It also approximatesS.sect.Tetragonostachyssensu Tryon (1955) who recognized four series (S.ser.ArenicolaeTryon,S.ser.SartoriiTryon,S.ser.RupestresTryon,andS.ser.EremophilaeTryon) based on the habit and leaf morphology.The relationships of the four series in our phylogeny are unresolved(Fig.S3).Two plastid genomes of species from this genus [Selaginella vardeiH.Lév.andS.indica(Milde) R.M.Tryon] were studied and extremely rare short dispersed repeats have been found(Zhang et al.,2019).

Morphologically,Bryodesmacan be easily distinguished in having xerophytic habit (Fig.5A),rhizophores borne on dorsal side of the stem (Fig.5H),sterile leaves spirally arranged throughout (Fig.5A-H),and axillary leaves absent (Fig.5H).The proximal surfaces of microspores always present rough and irregularly rugate ornamentation(Fig.5V),which is also found in these xerophytic species ofS.subg.Boreoselaginella(Zhou et al.,2015).

Plastome structures of nearly all species inBryodesmaknown so far (Zhou et al.,2022) are DR (direct repeats) structure and without small or medium repeats existed in SC (single-copy region).

Bryodesmacontains about 60 species,about 40 species in Americas (nearly all in North America),about 12 species in Africa and Madagascar and adjacent islands,and about five species in Asia (Tryon and Lugardon,1991;Valdespino,1993a;Roux,2009;Weakley,2012,2022;Arrigo et al.,2013;Fraser-Jenkins et al.,2015,2017;Zhou and Zhang,2015;Weststrand and Korall,2016b;Hassler,2022).Bryodesma corallinum(Riddell) Weakley has recently been transferred to the genus(Weakley,2022).

Members:

Bryodesma acanthonota(Underw.) Skoda,Preslia 68(4): 343(1997).Basionym:Selaginella acanthonotaUnderw.,Torreya 2: 172(1902).

Bryodesma aethiopicum(Bizzarri) Li Bing Zhang &X.M.Zhou,comb.&stat.nov.Basionym:Selaginella caffrorumvar.aethiopicaBizzarri,Webbia 29: 556 (1975).

Bryodesma arenaria(Baker)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella arenariaBaker,J.Bot.21: 82,no.23(1883).

Bryodesma arenicola(Underw.)Soják;Preslia 64(2):154(1992 publ.1993).Basionym:Selaginella arenicolaUnderw.,Bull.Torrey Bot.Club 25: 541 (1898).

Bryodesma arizonicum(Maxon)Soják;Preslia 64(2):154(1992 publ.1993).Basionym:Selaginella arizonicaMaxon,Smithsonian Misc.Collect.72: 5-6 (1920).

Bryodesma arsenei(Weath.) Soják;Preslia 64(2): 154 (1992 publ.1993).Basionym:Selaginella arseneiWeath.,J.Arnold Arbor.25: 417,t.2,f.8 (1944).

Bryodesma asprellum(Maxon) Soják;Preslia 64(2): 154 (1992 publ.1993) Basionym:Selaginella asprellaMaxon,Smithsonian Misc.Collect.72: 6 (1920).

Bryodesma balansae(A.Braun)Soják;Preslia 64(2):154(1992 publ.1993).Basionym:Selaginella balansae(A.Braun) Hieron.,Hedwigia 39: 318 (1900).

Bryodesma bigelovii(Underw.) Soják;Preslia 64(2): 154 (1992 publ.1993).Basionym:Selaginella bigeloviiUnderw.,Bull.Torrey Bot.Club 25: 130,no.6 (1898).

Bryodesma caffrorum(Milde) Soják;Preslia 64(2): 15,4 (1992 publ.1993).Basionym:Selaginella caffrorum(Milde) Hieron.,Hedwigia 39: 313 (1900).

Bryodesma basipilosum(Valdespino) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella basipilosaValdespino,Brittonia 44(3): 314(1992 publ.1993).

Bryodesma carinatum(R.M.Tryon)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella carinataR.M.Tryon,Ann.Mo.Bot.Gard.42: 50,f.25-26,map 31 (1955).

Bryodesma carnerosanum(T.Reeves) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella carnerosanaT.Reeves,Brittonia 32(3): 365 (1980).

Bryodesma cinerascens(A.A.Eaton) Soják;Preslia 64(2): 154(1992 publ.1993).Basionym:Selaginella cinerascensA.A.Eaton;Fern Bull.7: 33(1899).

Bryodesma corallinum(Riddell)Weakley;J.Bot.Res.Inst.Texas 16(2): 405 (2022).Basionym:Lycopodium corallinumRiddell;New Orleans Med.Surg.J.9: 617 (1853).

Bryodesma densum(Rydb.) Soják;Preslia 64(2): 154 (1992 publ.1993).Basionym:Selaginella densaRydb.;Mem.New York Bot.Gard.1: 7 (1900).

Bryodesma dregei(C.Presl)Soják;Preslia 64(2):154(1992 publ.1993).Basionym:Lycopodium dregeiC.Presl;Abh.Königl.Böhm.Ges.Wiss.,Math.-Naturw.Cl.V,3(1845),reimpr.in Bot.Bemerk.(C.Presl)153 (1846).

Bryodesma echinatum(Baker) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella echinataBaker,J.Linn.Soc.Bot.22: 536 (1887).

Bryodesma emodi(Fraser-Jenk.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella emodiFraser-Jenk.,in Fraser-Jenkins,Kandel &Pariyar,Ferns Fern-Allies Nepal 1: 67-68(2015).印度同形卷柏(新拟)

Bryodesma eremophilum(Maxon) Soják;Preslia 64(2): 154(1992 publ.1993).Basionym:Selaginella eremophilaMaxon;Smithsonian Misc.Collect.72(5): 3,t.2 (1920).

Bryodesma extensum(Underw.)Soják;Preslia 64(2):154(1992 publ.1993).Basionym:Selaginella extensaUnderw.;Bull.Torrey Bot.Club 25: 131 (1898).

Bryodesma griseum(Alston)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella griseaAlston,J.Bot.77: 222 (1939).

Bryodesma hansenii(Hieron.) Soják;Preslia 64(2): 154 (1992 publ.1993).Basionym:Selaginella hanseniiHieron.,Hedwigia 39:301 (1900).

Bryodesma landii(Greenm.&N.Pfeiff.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella landiiGreenm.&N.Pfeiff.,Ann.Mo.Bot.Gard.5: 205 (1918).

Bryodesma macratherum(Weath.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella macratheraWeath.,J.Arnold Arbor.24:326 (1943).

Bryodesma niveum(Alston)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella niveaAlston,Dansk Bot.Ark.7: 194,t.78(1-4) (1932),nom.nud.,et Cat.Pl.Madagasc.,Pterid.71(1932).

Bryodesma humbertii(Stefanov.&Rakotondr.)Li Bing Zhang&X.M.Zhou,comb.&stat.nov.Basionym:Selaginella niveasubsp.humbertiiStefanov.&Rakotondr.,Novon 6(2): 207(1996).

Bryodesma leucobryoides(Maxon) Skoda;Preslia 68(4): 343(1997).Basionym:Selaginella leucobryoidesMaxon,Smithsonian Misc.Collect.72(5): 8,t.5 (1920).

Bryodesma limitaneum(Weath.) Li Bing Zhang &X.M.Zhou,comb.&stat.nov.Basionym:Selaginella muticavar.limitaneaWeath.,J.Arnold Arbor.25: 414 (1944) ≡Bryodesma muticumvar.limitaneum(Weath.) Skoda,Preslia 68(4): 343 (1997).

Bryodesma muticum(D.C.Eaton ex Underw.) Soják;Preslia 64(2): 154 (1992 publ.1993).Basionym:Selaginella muticaD.C.Eaton ex Underw.,Bull.Torrey Bot.Club 25: 128 (1898).

Bryodesma×neomexicanum(Maxon)Skoda;Preslia 68(4):343(1997).Basionym:Selaginella×neomexicanaMaxon,Smithsonian Misc.Collect.72: 2 (1920).

Bryodesma njamnjamense(Hieron.) Soják;Preslia 64(2): 154(1992 publ.1993).Basionym:Selaginella njamnjamensisHieron.,Hedwigia 39: 312 (1900).

Bryodesma oreganum(D.C.Eaton) Soják;Preslia 64(2): 154(1992 publ.1993).Basionym:Selaginella oreganaD.C.Eaton;Wats.,Bot.California 2: 350(1880).

Bryodesma parishii(Underw.) Soják;Preslia 64(2): 154 (1992 publ.1993).Basionym:Selaginella parishiiUnderw.,Bull.Torrey Bot.Club 33:202 (1906).

Bryodesma peruvianum(Milde)Soják;Preslia 64(2):155(1992 publ.1993).Basionym:Selaginella peruviana(Milde) Hieron.,Hedwigia 39: 307(1900).

Bryodesma phillipsianum(Hieron.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella wightiivar.phillipsianaHieron.,Hedwigia 39:320(1900)≡Selaginella phillipsiana(Hieron.)Alston,J.Bot.77: 222 (1939).

Bryodesma proximum(R.M.Tryon)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella proximaR.M.Tryon,Ann.Mo.Bot.Gard.42: 56,f.33-35,map 37 (1955).

Bryodesma rupestre(L.) Soják;Preslia 64(2): 155 (1992 publ.1993).Basionym:Selaginella rupestris(L.)Spring;Flora 21:149 and 182,no.12 (1838).

Bryodesma rupincola(Underw.)Soják;Preslia 64(2):155(1992 publ.1993).Basionym:Selaginella rupincolaUnderw.;Bull.Torrey Bot.Club 25: 129 (1898).

Bryodesma sartorii(Hieron.) Soják,Preslia 64(2): 155 (1992 publ.1993).Basionym:Selaginella sartoriiHieron.;Hedwigia 39:304 (1900).

Bryodesma scopulorum(Maxon)Skoda&Holub,Preslia 68(4):343 (1997).Basionym:Selaginella scopulorumMaxon;Am.Fern J.11: 36 (1921).

Bryodesma sellowii(Hieron.)Soják,Preslia 64(2):155(1992 publ.1993).Basionym:Selaginella sellowiiHieron.,Hedwigia 39:306(1900).

Bryodesma shabaense(Bizzarri)LiBing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella shabaensisBizzarri,Webbia 36(1): 204(1982).

Bryodesma shakotanense(Franch.ex Takeda) Soják,Preslia 64(2): 155 (1992 publ.1993).Basionym:Selaginella shakotanensis(Franch.ex Takeda)Miyabe&Kudô;Fl.Hokkaido 63(1930).北海道同形卷柏(新拟)

Bryodesma sibiricum(Milde) Soják,Preslia 64(2): 155 (1992 publ.1993).Basionym:Selaginella sibirica(Milde) Hieron.;Hedwigia 39: 290,no.1 (1900).西伯利亚同形卷柏(新拟)

Bryodesma standleyi(Maxon)Skoda;Preslia 68(4):343(1997).Basionym:Selaginella standleyiMaxon;Smithsonian Misc.Collect.72(5): 9 (1920).

Bryodesma steyermarkii(Alston) Soják,Preslia 64(2): 155(1992 publ.1993).Basionym:Selaginella steyermarkiiAlston,Ann.Mag.Nat.Hist.XII,7: 638,t.XII.9,f.a-c.(1954).

Bryodesma tortipilum(A.Braun)Soják,Preslia 64(2):155(1992 publ.1993).Basionym:Selaginella tortipilaA.Braun,Ann.Sci.Nat.,Bot.,sér.V,3: 271 (1865).

Bryodesma underwoodii(Hieron.) Soják,Preslia 64(2): 155(1992 publ.1993).Basionym:Selaginella underwoodiiHieron.in Engl.&Prantl,Nat.Pflanzenfam.1: 714 (1901).

Bryodesma utahense(Flowers) Skoda &Holub,Preslia 68(4):343(1997).Basionym:Selaginella utahensisFlowers,Am.Fern J.39:83 (1949).

Bryodesma vardei(H.Lév.)Soják,Preslia 64(2):155(1992 publ.1993).Basionym:Selaginella vardeiH.Lév.,Cat.Pl.Yun-Nan 172,f.41 (1915)-17.细瘦同形卷柏(新拟)

Bryodesma viridissimum(Weath.) Soják,Preslia 64(2): 155(1992 publ.1993).Basionym:Selaginella viridissimaWeath.,J.Arnold Arbor.24: 326 (1943).

Bryodesma wallacei(Hieron.) Soják,Preslia 64(2): 155 (1992 publ.1993).Basionym:Selaginella wallaceiHieron.,Hedwigia 39:297 (1900).

Bryodesma watsonii(Underw.) Soják;Preslia 64(2): 155 (1992 publ.1993).Basionym:Selaginella watsoniiUnderw.,Bull.Torrey Bot.Club 25: 127 (1898).

Bryodesma weatherbianum(R.M.Tryon) Soják,Preslia 64(2):155 (1992 publ.1993).Basionym:Selaginella weatherbianaR.M.Tryon,Am.Fern J.40: 69 (1950).

Bryodesma wightii(Hieron.) Soják,Preslia 64(2): 155 (1992).Basionym:Selaginella wightiiHieron.,Hedwigia 39: 319 (1900).

Bryodesma wrightii(Hieron.) Soják;Preslia 64(2): 155(1992).Basionym:Selaginella wrightiiHieron,Hedwigia 39:298-299(1900).

5.4.3.Ericetorum

Ericetorum(Jermy)Li Bing Zhang&X.M.Zhou,stat.nov.对叶卷柏属(新拟) -Basionym:Selaginellasubg.EricetorumJermy,Fern Gaz.13(2): 117.1986 -Type:Ericetorum uliginosa(Labill.) Li Bing Zhang &X.M.Zhou (≡Selaginella uliginosa(Labill.) Spring).

=Didiclissect.LyalliaRothm.,Feddes Repert.Spec.Nov.Regni Veg.54:70(1944)≡Selaginellasect.Lyallia(Rothm.)Li Bing Zhang&X.M.Zhou,Taxon 64(6): 1133 (2015) -Type:Selaginella lyallii(Hook.&Grev.) Spring.

=Selaginella(unranked)TetrastichaeA.Braun,App.Ind.Sem.Hort.Berol.11.[1857] 1858 -Lectotype (designated by Zhou and Zhang 2015: 1133):Selaginella uliginosa(Labill.) Spring.

=Selaginellasubser.PleiostelicaeHieron.&Sadeb.in Engler &Prantl,Nat.Pflanzenf.L(4): 707.1902,non Hieronymus and Sadebeck,1901 (1902: 700,710)-Lectotype (designated by Zhou and Zhang 2015: 1133):Selaginella lyallii(Hook.&Grev.) Spring.

Ericetorumcircumscribed here corresponds toSelaginellasect.Lyallia(Rothm.)Li Bing Zhang&X.M.Zhou(2015)and toS.subg.Ericetorumsensu Weststrand and Korall(2016b).Its geographical distribution and constituent species are same as those reported in previous studies(Zhou and Zhang,2015;Weststrand and Korall,2016b).

Morphologically,Ericetorumcan be easily identified in having erect plant,solenostelic rhizome (Weststrand and Korall,2016a),rhizophores strictly restricted to the base of stem (Fig.5C),sterile leaves monomorphic and decussately arranged(or at least on stem)(Fig.5K;Stefanović et al.,1997;Schulz et al.,2013).Megaspore surfaces are reticulate and have convoluted laesurae forming a complex mass close to the pole on proximal surfaces (Fig.5P;Stefanović et al.,1997;Schulz et al.,2013).Microspore surfaces are gemmate,foveolate,verrucate to rugate ornamentation or scabrate often covered with microstructure being spines or gemmae(Fig.5U;Stefanović et al.,1997;Schulz et al.,2013).

The plastome ofEricetorumknown so far(Zhou et al.,2022)has DR structure,the lowest GC content(ca.50%),and the smallest LSC(ca.45 kb) in Gymnogynoideae.

Eight species are currently known in this genus containing four Australasian and four Afro-Malagasy species(Stefanović et al.,1997;Roux,2009;Schulz et al.,2013).

Members:

Ericetorum aboriginale(C.Schulz&Homberg)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella aboriginalisC.Schulz&Homberg,Syst.Bot.38(1): 11 (2013).

Ericetorum gracillimum(Kunze) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium gracillimumKunze,Lehm.,Pl.Preiss.2:109(1846-47)≡Lycopodioides gracillima(Kunze)Kuntze,Rev.Gen.Pl.1:825(1891)≡Selaginella gracillima(Kunze)Spring ex Salomon;Nomencl.353 (1883).

Ericetorum lyallii(Hook.&Grev.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium lyalliiHook.&Grev.,Enum.Fil.J.Bot.(Hook.) Kew Misc.2: 387,no.120(1831)≡Selaginella laevigatavar.lyallii(Hook.&Grev.)Baker,J.Bot.23:116(1885)≡S.lyallii(Hook.&Grev.)Spring;Bull.Acad.Roy.Soc.Bruxelles 10:146,no.84(1843).

Ericetorum moratii(W.Hagemann &Rauh) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella moratiiW.Hagemann&Rauh;Pl.Syst.Evol.176(3-4): 205 (1991).

Ericetorum pygmaeum(Kaulf.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium pygmaeumKaulf.,Enum.Fil.9(1824) ≡Lycopodium pumilumvar.pygmaeum(Kaulf.) Schltdl.,Adumbr.Fl.Aethiop.6 (1825) ≡Lycopodioides pygmaeum(Kaulf.)Kuntze,Rev.Gen.Pl.1: 825 (1891) ≡Selaginella pygmaea(Kaulf.)Alston;J.Bot.(London) 69: 257 (1931).

Ericetorum pectinatum(Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella pectinataSpring,Bull.Acad.Roy.Sci.Bruxelles 10(1):146.1843,nom.nov.forLycopodium pectinatumWilld.,Sp.Pl.,ed.4.5(1):44.1810,nom.illeg.,non Lamarck (1792:651).-Selaginella polymorphaBadre’,Fl.Madagasc.fam.14:25.1997,a nom.nov.but superfl.forL.pectinatumWilld.-Lectotype(designated by Smith et al.,2016): “America meridionale?” [protologue],“Habitat in Madagascar” [specimens],B-W 19400-02 0.

Ericetorum royenii(Alston) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella royeniiAlston;Nova Guinea ser.2,7: 2(1956).

Ericetorum uliginosum(Labill.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium uliginosumLabill.,Nov.Holl.pl.spec.2: 154,t.251,f.1 (1806) ≡Lycopodioides uliginosa(Labill.)Kuntze,Rev.Gen.Pl.1: 827 (1891) ≡L.uliginosumLabill.,Nov.Holl.pl.spec.2: 154,t.251,f.1 (1806) ≡Selaginella uliginosa(Labill.) Spring,Bull.Acad.Roy.Soc.Bruxelles 10: 136 (1843).

5.4.4.Gymnogynum

GymnogynumP.Beauv.,Mag.Encycl.9(5):480(1804)关节卷柏属(新拟)≡Lycopodiumsubg.Gymnogynum(P.Beauv.)Rchb.,Consp.Regn.Veg.: 78 (1828) ≡Didiclissect.Gymnogynum(P.Beauv.)Rothm.,Feddes Repert.Spec.Nov.Regni Veg.54:70(1944)-Type:Gymnogynumdomingense P.Beauv.,Mag.Encycl.9(5): 480 (1804)[=Selaginellaplumosa (L.) C.Presl ≡Lycopodium plumosumL.].

=Selaginellasect.Articulatae(Spring) Li Bing Zhang &X.M.Zhou,Taxon 64(6):1132(2015)≡Selaginella(unranked)ArticulataeSpring in Mém.Acad.Sci.Belg.24: 53 (1850) ≡S.ser.Articulatae(Spring) Hieron.&Sadeb.in Engler &Prantl,Nat.Pflanzenf.1(4):707 (1902) ≡Didiclissect.Articulatae(Spring) Rothm.in Feddes Repert.Spec.Nov.Regni Veg.54:71.(1944)-Lectotype(designated by Rothmaler,1944: 71):Didiclis sulcata(Desv.ex Poir.) Rothm.(=Selaginella sulcata(Desv.ex Poir.) Spring ex Mart.).

=Selaginella(unranked)AscendentesA.Braun in App.Ind.Sem.Hort.Berol.11 ([1857] 1858) non A.Braun (1858: 11) -Lectotype(designated by Zhou and Zhang,2015: 1132):Selaginella galeottiiSpring (=Selaginella stellataSpring).

=Selaginella(unranked)RepentesA.Braun in App.Ind.Sem.Hort.Berol.12.[1857] 1858,non A.Braun (1858: 11) -Lectotype(designated by Zhou and Zhang 2015: 1132):Selaginella hortensisMett.(=Selaginella kraussiana(G.Kunze) A.Braun).

=Selaginella(unranked)CaulescentesA.Braun in App.Ind.Sem.Hort.Berol.12 ([1857] 1858),non A.Braun (1858: 11) -Type:Selaginella asperulaSpring.

Selaginellasubser.PleiostelicaeHieron.&Sadeb.in Engler.&Prantl,Nat.Pflanzenf.1(4):710(1902)non Hieronymus and Sadebeck,1901(1902:700,707)-Type:Selaginella kraussiana(G.Kunze)A.Braun.

=Selaginellasubser.MonostelicaeHieron.&Sadeb.in Engler &Prantl,Nat.Pflanzenf.1(4): 708 (1902) non Hieronymus and Sadebeck,1901(1902:673,704)-Type:Selaginella remotifoliaSpring.

Gymnogynumcorresponds toSelaginellasect.Articulataesensu Zhou and Zhang (2015) and toS.subg.Gymnogynumsensu Weststrand and Korall (2016b).Gymnogynumis always well supported as monophyletic in morphology (Mickel et al.,2004) and phylogeny (Weststrand and Korall,2016a;Zhou et al.,2016;our Figs.1 and S1-S3).

Morphologically,Gymnogynumhas articulation (presenting swellings or nodules: Fig.5I) on stems and branches (Lopes et al.,2020),strobilus with only one large megasporophyll at the base(Fig.5J),rhizophores borne on dorsal side of stem or/and branch(Fig.5B,F-I),megaspores large and with reticulate ornamentation on surfaces(Korall and Taylor,2006;Zhou et al.,2015;Fig.5R),and microspore surfaces usually spiny or echinate(Fig.5M;Zhou et al.,2015;Valdespino and L‵opez,2020).

Thendhgene of plastomes inGymnogynumis preserved (Zhou et al.,2022).

This genus contains about 50 species mainly in North to South Americas and a few in Africa and Asia.Gymnogynum kraussianumis native to Africa and has been cultivated throughout the world and has been recently transferred toGymnogynumby Weakley (2022)asG.kraussianum(Kunze) Weakley.

Members:

Gymnogynum atirrense(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella atirrensisHieron.,Nat.Pflanzenf.1(4): 711 (1901).

Gymnogynum anaclastum(Alston ex Crabbe &Jermy) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella anaclastaAlston ex Crabbe &Jermy,Am.Fern J.63(3): 135(1973).

Gymnogynum angustifolium(A.Braun) Li Bing Zhang &X.M.Zhou,comb.&stat.nov.Basionym:Selaginella sericeavar.angustifoliaA.Braun;Ann.Sci.Nat.,Bot.,sér.V,3: 299 (1865).

Gymnogynum arthriticum(Alston)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella arthriticaAlston,Arch.Bot.(Forli)11: 43 (1935).

Gymnogynum articulatum(Kunze)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Lycopodium articulatumKunze,Linn.9: 10(1834 publ.1835) ≡Selaginella articulata(Kunze) Spring,Flora 21:182 (1838) ≡Lycopodioides articulata(Kunze) Kuntze,Rev.Gen.Pl.1: 825 (1891).

Gymnogynum asperulum(Mart.ex Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium asperulumC.Mart.ex Spring in Mart.Fl.Bras.1(2): 127,no.15 (1840) ≡Lycopodioides asperula(Mart.ex Spring) Kuntze,Rev.Gen.Pl.1: 825 (1891) ≡Selaginella asperulaMart.ex Spring,Mart.,Fl.Bras.1(2):127(1840).

Gymnogynum asplundii(Alston ex Crabbe &Jermy) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella asplundiiAlston ex Crabbe &Jermy,Fern Gaz.11(4): 257 (1976).

Gymnogynum buchtienii(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella buchtieniiHieron.,Med.Rijks Herb.Leiden,no.27: 2 (1915).

Gymnogynum caluffii(Shelton) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella caluffiiShelton,Willdenowia 33(1):159 (2003).

Gymnogynum coarctatum(Mart.ex Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella coarctataMart.ex Spring,Mart.,Fl.Bras.1(2): 126,no.14 (1840) ≡Lycopodioides coarctata(Mart.ex Spring) Kuntze,Rev.Gen.Pl.1: 826 (1891).

Gymnogynum conduplicatum(Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella conduplicataSpring,Mart.,Fl.Bras.1(2):129(1840)≡Selaginella geniculatasubsp.conduplicata(Spring) Hieron.in Engl.&Prantl,Nat.Pflanzenf.1(4): 712 (1901 publ.1902) ≡Selaginella geniculatavar.conduplicata(Spring) A.Braun,Ann.sc.nat.V,3: 303 (1865).

Gymnogynum decompositum(Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella decompositaSpring,Mart.,Fl.Bras.1(2): 123,no.10 (1840).

Gymnogynum diffusum(C.Presl) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium diffusumC.Presl,Rel.Haenk.1(1): 78 (1825) (nom.illeg.,non R.Br.(1810)) ≡Selaginella diffusa(C.Presl) Spring,Bull.Acad.Roy.Soc.Bruxelles 10 (1): 143(1843).

Gymnogynum eurynotum(A.Braun)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella eurynotaA.Braun,Ann.sc.nat.V,3:293(1865)≡Lycopodioides eurynota(A.Braun)Kuntze,Rev.Gen.Pl.1: 826 (1891).

Gymnogynum expansum(Sodiro) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella expansaSodiro,Rec.Cr.vasc.quit.95,no.12(1883)≡Lycopodioides expansa(Sodiro)Kuntze,Rev.Gen.Pl.1: 826 (1891).

Gymnogynum flabellum(Desv.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella flabellum(Desv.) Spring,Bull.Acad.Roy.Soc.Bruxelles 10: 234,no.157 (1843).

Gymnogynum fragile(A.Braun) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella fragilisA.Braun,Ann.sc.nat.V,5,3: 305 (1865).

Gymnogynum filicaule(Sodiro) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:SelaginellafilicaulisSodiro,Rec.Cr.vasc.quit.600 (1893),et Anal.Univ.Centr.Ecuador 12: 413 (1895).

Gymnogynum fuertesii(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella fuertesiiHieron.,Urban,Symb.Ant.7: 164 (1912).

Gymnogynum geniculatum(C.Presl)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Lycopodium geniculatumC.Presl,Rel.Haenk.1(1):80(1825)≡Selaginella geniculata(C.Presl)Spring,Bull.Acad.Roy.Soc.Bruxelles 10(1):230(1843) ≡Lycopodioides geniculata(C.Presl)Kuntze,Rev.Gen.Pl.1: 826 (1891).

Gymnogynum germinans(Valdespino&C.López)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella germinansValdespino &C.López,Botany Letters 165(3-4): 488 (2018).

Gymnogynum horizontale(C.Presl)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Lycopodium horizontaleC.Presl,Rel.Haenk.1(1): 78 (1825) ≡Selaginella horizontalis(C.Presl) Spring;Bull.Acad.Roy.Soc.Bruxelles 10(1): 226 (1843).

Gymnogynum humboldtianum(A.Braun)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella humboldtianaA.Braun;Ann.Sci.Nat.,Bot.,sér.V,5,3: 293 (1865).

Gymnogynum ivanii(Shelton &Caluff) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella ivaniiShelton &Caluff,Willdenowia 33(1):162 (2003).

Gymnogynum kraussianum(Kunze) Weakley;J.Bot.Res.Inst.Texas 16(2): 405 (2022).Basionym:Lycopodium kraussianumKunze;Linnaea 18: 114 (1844).小翠云(栽培)

Gymnogynum kunzeanum(A.Braun) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella kunzeanaA.Braun,Ann.sc.nat.V,3: 296-297,no.35 (1865) ≡Lycopodioides kunzeana(A.Braun)Kuntze,Rev.Gen.Pl.1: 826 (1891).

Gymnogynum lingulatum(Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella lingulataSpring,Mém.Acad.Roy.Sci.Belg.24: 224 (1849) ≡Lycopodioides lingulata(Spring) Kuntze,Rev.Gen.Pl.1: 826 (1891).

Gymnogynum marginatum(Humb.&Bonpl.ex Willd.) Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Lycopodium marginatumHumb.&Bonpl.ex Willd.,Sp.Pl.5: 41,no.58 (1810) ≡Selaginella marginata(Humb.&Bonpl.ex Willd.)Spring,Flora 21:194(1838)≡Lycopodioides marginata(Kunth)Kuntze,Rev.Gen.Pl.1:826(1891).

Gymnogynum microtus(A.Braun) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella microtusA.Braun,Ann.sc.nat.V,3: 293,no.32 (1865).

Gymnogynum moranianum(Valdespino &C.López) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella moranianaValdespino &C.López,Brittonia 72(1): 24(2019).

Gymnogynum parkeri(Hook.&Grev.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium parkeriHook.&Grev.,Enum.Fil.J.Bot.(Hook.) Kew Misc.2: 388,no.123 (1831) ≡Selaginella parkeri(Hook.&Grev.) Spring,Bull.Acad.Roy.Sci.Bruxelles 10:146(1843)≡Lycopodioides parkeri(Hook.&Grev.)Kuntze,Rev.Gen.Pl.2: 827 (1891).

Gymnogynum parviarticulatum(W.R.Buck) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella parviarticulataW.R.Buck,Brittonia 38(1): 45 (1986).

Gymnogynum pedatum(Klotzsch) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella pedataKlotzsch,Linnaea 17: 521(1844).

Gymnogynum plumosum(L.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Lycopodium plumosumL.,Sp.Pl.2: 1100 (1753) ≡Stachygynandrum plumosum(L.)P.Beauv.,Fl.D'Oware,1:10(1805)≡Lycopodioides plumosa(L.) Kuntze,Rev.Gen.Pl.1: 825 &827(1891)≡Selaginella plumosa(L.)C.Presl,Abh.(K.)Böhm.Ges.Wiss.,Math.-Naturw.Cl.V,3: 583 (1845).

Gymnogynum poeppigianum(Spring)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Lycopodium poeppigianumHook.&Grev.,Enum.Filic.J.Bot.(Hook.) Kew Misc.2: 392-393,no.143 (1831).≡Selaginella poeppigiana(Spring)Spring,Flora 21:185(1838).≡Lycopodioides poeppigiana(Spring.)Kuntze,Revis.Gen.Pl.2:827(1891).

Gymnogynum remotifolium(Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella remotifoliaSpring,Miq.Pl.Jungh.3:276,no.5(1854)≡Lycopodioides remotifolia(Spring)H.S.Kung,Fl.Sichuan.6:65-67,t.19,10-15(1988).疏叶关节卷柏(新拟)

Gymnogynum schizobasis(Baker) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella schizobasisBaker,J.Bot.21: 333,no.86(1883)≡Lycopodioides schizobasis(Baker)Kuntze,Rev.Gen.Pl.1: 827 (1891).

Gymnogynum sericeum(A.Braun) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella sericeaA.Braun,Ann.sc.nat.V,3:298-299,no.37 (1865) ≡Lycopodioides sericea(A.Braun) Kuntze,Rev.Gen.Pl.1: 827 (1891).

Gymnogynum sertatum(Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella sertataSpring,Mém.Acad.Roy.Sci.Belg.24: 104 (1849) ≡Lycopodioides sertata(Spring) Kuntze,Rev.Gen.Pl.1: 827 (1891).

Gymnogynum silvestre(Aspl.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella silvestrisAspl.,Ark.för Bot.20A(7):30,f.3-5 (1926).

Gymnogynum stellatum(Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella stellataSpring,Flora 21:188(1838)≡Selaginella parkerivar.stellata(Spring)Baker,J.Bot.23:120(1885)≡Selaginella stoloniferavar.stellataSpring,Flora 21:194(1838).

Gymnogynum suave(Spring)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella sulcatassp.suavisSpring,Flora 21: 185(1838) ≡Selaginella suavis(Spring) Spring,Bull.Acad.Roy.Soc.Bruxelles 10: 229 (1843) ≡Lycopodioides suavis(Spring) Kuntze,Rev.Gen.Pl.1: 827 (1891).

Gymnogynum sulcatum(Desv.ex Poir.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium sulcatumDesv.ex Poir.,Lam.,Encycl.suppl.3: 549,no.74.(1813 publ.1814) ≡Selaginellasulcata(Desv.ex Poir.) Spring ex Mart.,Flora 20 (2): 126 (1837) ≡D.sulcata(Desv.)Rothm.,Fedde,Repert.Spec.Nov.54:71(1944)≡Lycopodioides sulcata(Desv.) Kuntze,Rev.Gen.Pl.1: 827 (1891).

Gymnogynum tardum(Mickel &Beitel) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella tardaMickel&Beitel,Mem.New York Bot.Gard.46: 353,f.128H-N (1988).

Gymnogynum tenuissimum(Fée) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella tenuissimaFée,Cr.vasc.Br.2: 98,no.6/2,t.108 (1873) ≡Lycopodioides tenuissima(Fée) Kuntze,Rev.Gen.Pl.1: 827 (1891).

Gymnogynum tomentosum(Spring)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella tomentosaSpring,Mém.Acad.Roy.Sci.Belg.24:231(1849)≡Selaginella geniculatavar.tomentosa(Spring) Baker,J.Bot.23: 121 (1885).

Gymnogynum trisulcatum(Aspl.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella trisulcataAspl.,Ark.för Bot.20A(7): 34,f.6 (1926).

Gymnogynum versatile(A.R.Sm.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella versatilisA.R.Sm.,Ann.Mo.Bot.Gard.77(2): 270(1990).

5.4.5.Lepidoselaginella

LepidoselaginellaLi Bing Zhang&X.M.Zhou,gen.nov.鳞叶卷柏属(新拟) -Type:Lepidoselaginella lepidophylla(Hook.&Grev.) Li Bing Zhang&X.M.Zhou ≡Lycopodium lepidophyllumHook.&Grev.,Icon.Filic.2(9):162(1830)≡Selaginella lepidophylla(Hook.&Grev.)Spring,Fl.Bras.(Martius) 1(2): 126 (1840).

=Selaginellasect.LepidophyllaeLi Bing Zhang&X.M.Zhou,Taxon 64(6): 1133 (2015) -Type:Selaginella lepidophylla(Hook.&Grev.)Spring.

Etymology:“Lepido-”derived fromlepidophylla,referring to the scales-looking leaves of the type species.

Lepidoselaginellacorresponds toSelaginellasect.Lepidophyllaesensu Zhou and Zhang (2015) andS.subg.Lepidophyllaesensu Weststrand and Korall(2016b).It is strongly supported as sister toBryodesma(Weststrand and Korall,2016a;Zhou et al.,2016;our Figs.1 and S1-S3).

Morphologically,Lepidoselaginellahas rosette habit(Fig.5D)and rhizophores borne on dorsal side of the stem.Although some taxa in other genera (e.g.,Selaginella pallescens,S.nothohybrida,and most members ofPulviniella) also have rosette habit,they have rhizophores borne on ventral side of the stem.Lepidoselaginellahas megaspores with ridges on the distal surface (Korall and Tryon,2006) and microspores with striped ornamentation which are different from rest of species in the family (Fig.5W).

The plastome ofLepidoselaginellahas IR structure (Zhou et al.,2022).

Lepidoselaginellacontains about three species occurring in Mexico and USA (Mickel and Smith,2004;Weststrand and Korall,2016a;Zhou et al.,2016).

Members:

Lepidoselaginella lepidophylla(Hook.&Grev.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Lycopodium lepidophyllumHook.&Grev.,Icon.Fil.2(9):162(1830),et Enum.Fil.J.Bot.(Hook.)Kew Misc.3:106(1833)≡Selaginella lepidophylla(Hook.&Grev.)Spring,Mart.,Fl.Bras.1(2):126(1840)≡Lycopodioides lepidophylla(Hook.)Kuntze,Rev.Gen.Pl.1: 826 (1891).

Lepidoselaginella novoleonensis(Hieron.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella novoleonensisHieron.in Engl.&Prantl,Nat.Pflanzenf.1(4):676,no.56.(1901 publ.1902).

Lepidoselaginella ribae(Valdespino) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella ribaeValdespino,Mem.New York Bot.Gard.88 (Pterid.Mexico): 591-592 (2004).

5.4.6.Megaloselaginella

MegaloselaginellaLi Bing Zhang&X.M.Zhou,gen.nov.大孢卷柏属(新拟)-Type:Megaloselaginella exaltata(Kunze)Li Bing Zhang&X.M.Zhou (≡Lycopodium exaltatumKunze ≡Selaginella exaltata(Kunze)Spring).

=Selaginellasect.MegalosporarumLi Bing Zhang &X.M.Zhou,Taxon 64(6): 1133(2015 -Type:S.exaltata Kunze.

Etymology: “Megalo-” derived fromMegalosporarum,referring to the extremely large megaspores (ca.1.5 mm).

Megaloselaginellacorresponds toSelaginellasect.Megalosporarumsensu Zhou and Zhang (2015) and part ofS.subg.Exaltataesensu Weststrand and Korall (2016b).

Megaloselaginellais very similar toGymnogynumin having rhizophores borne on dorsal side of the stem and only one megasporophyll at the base of strobili (Fig.5F).However,Megaloselaginellahas large erect plants(up to 1 m)(Fig.5E),several steles in a special actino-plectoste (a sort of three-lobed plectostele) (Lopes et al.,2020),largest megaspores in the family (ca.1.5 mm;Mickel and Hellwig,1969,Fig.5O),and pillared to baculate ornamentation on microspore surfaces (Fig.5T).

The plastome ofMegaloselaginellaknown so far (Zhou et al.,2022) has DR-IR coexisting structure and the intron2 ofycf3(Zhou et al.,2022).

This genus contains about three species in Central and South Americas and Africa (Steyermark et al.,1986;Zhou et al.,2016;Hassler,2022).

Members:

Megaloselaginella exaltata(Kunze)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Lycopodium exaltatumKunze,Linn.9:8(1834[1835]) ≡Selaginella exaltata(Kunze) Spring,Bull.Acad.Roy.Soc.Bruxelles 10: 234 (1843) ≡Lycopodioides exaltata(Kunze) Kuntze,Rev.Gen.Pl.1: 826 (1891).

Megaloselaginella gigantea(Steyerm.&A.R.Sm.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella giganteaSteyerm.&A.R.Sm.,Ann.Mo.Bot.Gard.73(1): 209 (1986).

Megaloselaginella grallipes(Alston)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella grallipesAlston,Mem.Soc.Linn.Normandie Bot.1(3): 80 (1938).

5.5.Subfamily IV.Sinoselaginelloideae

Subfamily IV.SinoselaginelloideaeLi Bing Zhang &X.M.Zhou,subfam.nov.中华卷柏亚科(新拟) -Type:SinoselaginellaLi Bing Zhang &X.M.Zhou.

Plants erect,ascending,or creeping (Fig.6A,G),with ventral rhizophores(Fig.6B);sterile leaves dimorphic(Fig.6A-C,E,G,H);sporophylls monomorphic (Fig.6D,F);strobili with only one (to a few) megasporophyll at the base (Quanansah,1988;Zhao et al.,2001);megaspore surfaces verrucate,prominent elements composed by long curved baculate(Fig.6P)or rope microstructure(Zhou et al.,2015);microspore surfaces rugate (Fig.6M-O),coarse (Fig.6Q-T),sometimes with holes on proximal surfaces(Fig.6Q-S).

The plastomes of Sinoselaginelloideae known so far (Zhou et al.,2022) have the smallest size (78-90 kb) in Selaginellaceae and the plastome structure is more like “network” rather than looping.Exceptrrn16andrrn23,most of the plastid tRNA and rRNA genes are lost in Sinoselaginelloideae.Sinoselaginelloideae have the least protein-coding genes in Selaginellaceae (ca.27)(Zhou et al.,2022).

Some species of this subfamily like most of land plant lineages have a low GC content (ca.30%).

This subfamily corresponds to theSinensisgroup sensu Korall and Kenrick (2004) and Weststrand and Korall(2016a).

It contains about 24 species in three genera:Austroselaginella,Korallia,andSinoselaginella,occurring in Africa,Asia,Australia,and Indian Ocean islands.

5.5.1.Austroselaginella

AustroselaginellaLi Bing Zhang&X.M.Zhou,gen.nov.澳洲卷柏属(新拟)-Type:Austroselaginellaaustraliensis(Baker)Li Bing Zhang&X.M.Zhou (≡Selaginella australiensisBaker,J.Bot.21: 144,no.55(1883)).

Austroselaginellahas creeping plants,monomorphic strobili,and only one (to a few) megasporophyll at the base of strobili (Fig.6F).A.australiensisis very similar toGymnogynum remotifoliumandG.kraussianum,but the former has the rhizophores borne on ventral side and non-articulate stems.Austroselaginellais also similar to some species with creeping plants inKorallia,but the former has the base of ventral leaves not auriculate(Fig.5E vs.5H)and microspores without holes onproximal surfaces(Fig.5T vs.5Q-S).Austroselaginellahasvery large megaspores(600-750 μm in diam.).Although Korall and Taylor(2006) reported that the megaspore surfaces ofA.australiensiswere strongly reticulate,their recent study (Weststrand and Korall,2016)clarified that the “reticulate surfaces of megaspores” was probably based on a misidentified specimen.Jermyand Holmes(1998)reported that the megaspore surfaces ofA.australiensisare faintly reticulate.We did not examine the megaspores ofA.australiensisbut its microspore surfaces are somewhat smooth(Fig.6T).

It contains about four species distributed in rain-forested regions of north-eastern Queensland in Australia(Jermy and Holmes,1998).

Members:

Austroselaginella andrewsii(Jermy &J.S.Holmes) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella andrewsiiJermy &J.S.Holmes,Fl.Australia 48: 705(1998).

Austroselaginella australiensis(Baker) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella australiensisBaker,J.Bot.21:144,no.55(1883)≡Lycopodioides australiensis(Baker)Kuntze,Rev.Gen.Pl.1: 825 (1891).

Austroselaginella brisbanensis(F.M.Bailey) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella brisbanensisF.M.Bailey,Queensl.Fl.Suppl.62 (1886) ≡Lycopodioides brisbanensis(F.M.Bailey) Kuntze,Rev.Gen.Pl.1: 826 (1891).

Austroselaginella leptostachya(F.M.Bailey) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella leptostachyaF.M.Bailey,Proc.Roy.Soc.Queensl.1: 13 (1884) ≡Selaginella australiensisvar.leptostachya(F.M.Bailey) Domin,Bibl.Bot.85: 237(1913).

5.5.2.Korallia

KoralliaLi Bing Zhang&X.M.Zhou,gen.nov.科氏卷柏属(新拟)-Type:Korallia fissidentoides(Hook.&Grev.) Li Bing Zhang &X.M.Zhou (≡Lycopodium fissidentoidesHook.&Grev.,Enum.Fil.Hook.Bot.,Misc.2: 305,no.151 (1831)).

=Selaginellasect.OligomacrosporangiataeHieron.&Sadeb.in Engler &Prantl,Nat.Pflanzenf.1(4): 704 (1902) ≡Didiclissubg.Oligomacrosporangiatae(Hieron.&Sadeb.) Rothm.in Feddes Repert.Spec.Nov.Regni Veg.54: 70 (1944) ["Oligomacrosporangiata"] -Lectotype (designated by Rothmaler,1944: 70):Selaginella fissidentoides(Hook.&Grev.) Spring.

=Selaginellasubser.MonostelicaeHieron.&Sadeb.in Engler &Prantl,Nat.Pflanzenf.1(4):704(1902)non Hieronymusand Sadebeck,1901 (1902: 673,708) -Lectotype (designated by Zhou and Zhang,2015:1135):Selaginella fissidentoides(Hook.&Grev.)Spring.

Etymology:-Fromkorall-,inhonor of Prof.Petra Korall of Uppsala University,Sweden,for her contributions to the study of ferns in general and that ofSelaginellain particular (e.g.,Korall et al.,1999;Korall and Kenrick,2002,2004;Weststrand and Korall,2016a,b).

In our earlier classification(Zhou and Zhang,2015),we thoughtSelaginellafissidentoidesshould be resolved inDidiclisand misappliedS.sect.OligomacrosporangiataetoDidiclis.Koralliais a member of theSinensisgroup and sister toAustroselaginella(Weststrand and Korall,2016;our Figs.S1-S3).

Species ofKoralliahave erect,creeping or ascending plants and often a biauriculate base of ventral leaves(Fig.6H).The megaspore surfaces are often covered with dense and thin thorns (Fig.6P).Microspores have a special hole on each proximal district of the proximal surfaces (Fig.6Q-S)-such structure was never found or reported in other genera in the family.

The master plastome ofKoralliaknown so far(Zhou et al.,2022)can mediate 12 isomers through repeats in SC.Almost all genes do not have copies (exceptrrn16/23andndhE) and the GC content is larger than 50% inKorallia.Koralliais the only genus with two copies ofndhEin Selaginellaceae.

Koralliacontains about 15 species distributed in Madagascar and adjacent Indian Ocean islands (e.g.,Mauritius,Mayotte,Reunion,and Seychelles;Badré,2008).

Members:

Korallia amphirrhizos(A.Braun ex Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella amphirrhizosA.Braun ex Hieron.,Nat.Pflanzenf.1(4): 705 (1902) ≡S. fissidentoidesvar.amphirrhizos(A.Braun ex Hieron.) Stefanov.&Rakotondr.,Novon 6(2): 208 (1996).

Korallia balfourii(Baker)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella balfouriiBaker,Fl.Maurit.522,no.2(1877)≡Lycopodioides balfourii(Baker) Kuntze,Rev.Gen.Pl.1: 826 (1891).

Korallia cataphracta(Willd.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Lycopodium cataphractumWilld.,Sp.Pl.5:43,no.62(1810)≡Selaginella cataphracta(Willd.)Spring,Flora 21:209(1839).

Korallia concinna(Sw.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium concinnumSw.,Syn.Fil.182,408,no.52(1806)≡Selaginella concinna(Sw.)Spring,Flora 21:188,no.15(1838).

Korallia distachya(Cordem.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella distachyaCordem.,Bull.Soc.Sc.Arts La Réunion,104 (1890-91).

Korallia fissidentoides(Hook.&Grev.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium fissidentoidesHook.&Grev.,Enum.Fil.Hook.Bot.,Misc.2:305,no.151(1831)≡Selaginella fissidentoides(Hook.&Grev.)Spring,Bull.Acad.Roy.Soc.Bruxelles 8(12): 142 (1841) ≡Didiclis fissidentoides(Hook.&Grev.) Rothm.,Fedde,Repert.Spec.Nov.54: 70 (1944) ≡Lycopodioides fissidentoides(Hook.&Grev.) Kuntze,Rev.Gen.Pl.1: 826 (1891).

Korallia fruticulosa(Bory ex Willd.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Lycopodium fruticulosumBory ex Willd.,Sp.Pl.5: 41,no.59 (1810) ≡Selaginella fruticulosa(Bory) Spring,Flora 21: 202 (1838).

Korallia obtusa(P.Beauv.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium obtusum(P.Beauv.)Desv.ex Poir.,Lam.,Encycl.suppl.3: 548.(1813 publ.1814) ≡Selaginella obtusa(P.Beauv.) Spring,Bull.Acad.Roy.Soc.Bruxelles 10: 228 (1843) ≡Lycopodioides obtusa(P.Beauv.)Kuntze,Rev.Gen.Pl.1:827(1891).

Korallia rodriguesiana(Baker) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella rodriguesianaBaker,Fl.Maurit.523,no.4(1877)≡Lycopodioides rodriguesiana(Baker)Kuntze,Rev.Gen.Pl.1: 827 (1891).

Korallia sechellara(Baker) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella sechellarumBaker,Fl.Maurit.523,no.523(1877)≡Lycopodioides sechellarum(Baker)Kuntze,Rev.Gen.Pl.1: 827 (1891).

Korallia serrulata(Desv.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium serrulatumDesv.ex Poir.,Lam.,Encycl.suppl.3:550,no.78.(1813 publ.1814)≡Selaginella serrulata(Desv.)Spring,Bull.Acad.Roy.Soc.Bruxelles 10: 228,no.113 (1843).

Korallia sinuosa(Desv.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Lycopodium sinuosumDesv.ex Poir.,Lam.,Encycl.suppl.3:558.(1813 publ.1814)≡Selaginella sinuosa(Desv.)Alston,J.Bot.72: 230 (1934).

Korallia sparsifolia(Desv.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium sparsifoliumDesv.ex Poir.,Lam.,Encycl.suppl.3: 553,no.95.(1813 publ.1814) ≡Selaginella sparsifolia(Desv.) Badré,Fl.Mascar.Ptérid.1: 45 (2008).

Korallia tereticaulis(Desv.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium tereticaulonDesv.ex Poir.,Lam.,Encycl.suppl.3: 551,no.85.(1813 publ.1814) ≡Selaginella tereticaulis(Desv.) Spring,Flora 21: 210 (1838).

Korallia viridula(Bory) Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Lycopodium viridulumBory ex Willd.,Sp.Pl.5:37,no.50(1810) ≡Selaginella viridula(Bory) Spring,Flora 21: 190 (1838).

5.5.3.Sinoselaginella

SinoselaginellaLi Bing Zhang &X.M.Zhou,gen.nov.中华卷柏属(新拟) -Type:S.sinensis(Desv.) Li Bing Zhang &X.M.Zhou [≡Lycopodium sinenseDesv.,Mém.Soc.Linn.Paris 6: 189 (1827) ≡Selaginella sinensis(Desv.) Spring,Bull.Acad.Roy.Sci.Bruxelles 10(1): 137 (1843)].

Etymology:-“Sino-” derived fromsinensis,referring to the Chinese distribution of the type species.

Sinoselaginellais part of theSinensisgroup.It is sister toAustroselaginella+Korallia.Morphologically,Sinoselaginellais xerophytic and has creeping stems(Fig.6A),rhizophores borne on ventral side in axils of branches (Fig.6B),intervals throughout the main stem(Fig.6A and B),only one megasporophyll on the ventral side of the base of strobili(Fig.6F),large and usually verrucate and tuberculate ornamentation on megaspore surfaces and this prominent elements comprised by ropes (Fig.6I-K;Zhou et al.,2015),and microspore surfaces with prominent verrucae,and/or somewhat ridges (Fig.6M-N).Species ofSinoselaginellaare very similar to some species ofBoreoselaginella,especiallySinoselaginella albocinctaandB.sanguinolenta,two species with extremely similar appearance(Kung,1981)and overlapping distribution(Zhang et al.,2013),butB.sanguinolentahas strobili with several megasporophylls rather than one (Fig.4E-G) and rhizophores borne on the dorsal side of stems (Fig.4D).

Nearly all species inSinoselaginellaknown so far (Zhou et al.,2022) have protein-coding genes with two or three copies.GC content of the plastome isca.30% inSinoselaginellawhich is different from other taxa in Selaginellaceae (Zhou et al.,2022).

Sinoselaginellacontains about five species (Sinoselaginella adunca,S.albocincta,S.chuweimingii,S.sinensis,andS.yemensis)distributed in China extending to the Arabic area and eastern Africa(Somalia,Ethiopia,Kenya)(Zhang et al.,2013;Fraser-Jenkins et al.,2015,2017;Zhou et al.,2015).

Members:

Sinoselaginella adunca(A.Braun ex Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella aduncaA.Braun ex Hieron.in Engl.&Prantl,Nat.Pflanzenf.1(4):674(1901 publ.1902).

Sinoselaginella albocincta(Ching ex H.S.Kung) Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella albocinctaChing ex H.S.Kung,Acta Bot.Yunnan.3(2): 251-252,f.1: 9-14 (1981)≡Selaginella aduncassp.albocincta(Ching) Fraser-Jenk.,Indian Fern J.25(1-2): 8 (2008 publ.2009) ≡Lycopodioides albocincta(Ching ex H.S.Kung) H.S.Kung,Fl.Sichuan.6: 61 (1988).白边中华卷柏(新拟)

Sinoselaginella chuweimingii(X.M.Zhou,Z.R.He,Liang Zhang&Li Bing Zhang)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella chuweimingiiX.M.Zhou,Z.R.He,Liang Zhang &Li Bing Zhang,Phytotaxa 231(3): 284 (2015).维明中华卷柏(新拟)

Sinoselaginella sinensis(Desv.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium sinenseDesv.,Mém.Soc.Linn.Paris 6: 189 (1827) ≡Selaginella sinensis(Desv.) Spring,Bull.Acad.Roy.Soc.Bruxelles 10: 137,no.19 (1843) ≡Lycopodioides sinensis(Desv.) Satou,Hikobia 12(3): 269 (1997).中华卷柏(新拟)

Sinoselaginella yemensis(Sw.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium yemenseSw.,Syn.Fil.182,407,no.49,t.4,f.4(1806)≡Selaginella yemensis(Sw.)Spring ex Decne.,Arch.Mus.Paris 2: 191 (1941-42) ≡Lycopodioides yemensis(Sw.)Kuntze,Rev.Gen.Pl.1: 827 (1891).

5.6.Subfamily V.Pulvinielloideae

Subfamily V.PulvinielloideaeLi Bing Zhang&X.M.Zhou,subfam.nov.垫状卷柏亚科(新拟) -Type:Pulviniella(Li Bing Zhang &X.M.Zhou) Li Bing Zhang &X.M.Zhou.

Plants often rosette-forming(Fig.7A and B),or erect and tufted when dry (Fig.7C),with ventral rhizophores;sterile leaves dimorphic,four rows(Fig.7A);sporophylls monomorphic(Fig.7D);megaspores surface coarse,usually without obvious ornamentation(Fig.7E-H);microspores globose,surfaces coarse or nonprominent verruca and granule (Fig.7I-L);2n=20.

The plastome of Pulvinielloideae known so far(Zhou et al.,2022)has DR structure and only one conformation.Genesrpl16 tend to pseudogenize,andtrnC-GCA andrps18are lost in Pulvinielloideae.Allndhgenes are lost or pseudogenized in Pulvinielloideae.

This subfamily corresponds toSelaginellasubg.Pulviniellasensu Zhou and Zhang (2015).

It contains about 17 species in one genus,Pulviniella,occurring in Africa,Asia,and North and Central Americas.

5.6.1.Pulviniella

Pulviniella(Li Bing Zhang &X.M.Zhou) Li Bing Zhang &X.M.Zhou,stat.nov.垫状卷柏属(新拟) -Basionym:Selaginellasubg.PulviniellaLi Bing Zhang&X.M.Zhou,Taxon 64(6):1133(2015)-Type:Pulviniella pulvinata(Hook.&Grev.) Li Bing Zhang &X.M.Zhou ≡S.pulvinata(Hook.&Grev.) Maxim.

Pulviniellacorresponds toSelaginellasubg.PulviniellaLi Bing Zhang&X.M.Zhou and our data resolved it as sister to theSinensisgroup (Figs.1 and S3).However,Weststrand and Korall's (2016a)study found it to be sister to a clade includingS.subg.Heterostachyssensu Zhou and Zhang(2015)andS.subg.Stachygynandrumsensu Zhou and Zhang (2015).

Most species ofPulviniellahave rosette habit.The rosette habit evolved at least three times in Selaginellaceae: once inPulviniella(e.g.,S.pulvinata),once inLepidoselaginella(e.g.,L.lepidophyllaandL.novoleonensis),and once inSelaginella s.s.(e.g.,theS.pallescens+S.nothohybridaclade),respectively.However,except those inPulviniella,all species with rosette habit in Selaginellaceae have reticulate ornamentation.2n=20 or 24 (Jermy et al.,1967;Takamiya,1993;Marcon et al.,2005).

Pulviniellaand some species ofSinoselaginellaoften overlap in distribution and share xerophyte habit in China.For example,theS.chuweimingii-P.pulvinatapairand theS.sinensis-P.tamariscanapair,respectively,usually co-occur in a same habitat in China(Chu et al.,2006;Zhang et al.,2013;our field observations),but they are not sister to each other,which is consistent with morphology.

Pulviniellacontainsca.17 species occurring through Africa,Asia,and North &Central Americas (Zhang et al.,2013;Fraser-Jenkins et al.,2015,2017;Zhou et al.,2015;Yang et al.,2023a).

Members:

Pulviniella algida(Jie Yang bis &X.C.Zhang) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella algidaJie Yang bis &X.C.Zhang,Taxon 72: 14-15 (2023).高寒垫状卷柏(新拟)

Pulviniella bryopteris(L.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium bryopterisL.,Sp.Pl.2: 1103 (1753) ≡Selaginella bryopteris(L.) Baker,J.Bot.22: 376 (1884) ≡Lycopodioides bryopteris(L.) Kuntze,Rev.Gen.Pl.1: 825 (1891) ≡Stachygynandrum bryopterisP.Beauv.,Prodr.aethéogam.109 (1805).

Pulviniella convoluta(Arn.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium convolutumArn.,Mem.Wern.Nat.Hist.Soc.5:199(1824)≡Lycopodioides convoluta(Arn.)Kuntze,Rev.Gen.Pl.1:826(1891)≡Selaginella convoluta(Arn.)Spring,Mart.,Fl.Bras.1 (2): 131 (1840).

Pulviniella digitata(Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella digitataSpring,Mém.Acad.Roy.Sci.Belg.24:75(1850)≡Lycopodioides digitata(Spring)Kuntze,Rev.Gen.Pl.1: 826 (1891).

Pulviniella graniticola(Jie Yang bis&X.C.Zhang)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella graniticolaJie Yang bis &X.C.Zhang,Taxon 72: 15.2023.花岗岩垫状卷柏(新拟)

Pulviniella gypsophila(A.R.Sm.&T.Reeves) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella gypsophilaA.R.Sm.&T.Reeves,Sida 10(3): 211 (1984).

Pulviniella helioclada(Alston)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella helicocladaAlston,Dansk Bot.Ark.7:195,t.78(10-12)(1932),nom.nud.,et ex C.Chr.,Perrier Cat.71(1932).

Pulviniella imbricata(Forssk.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium imbricatumForssk.,Fl.Aegypt.-Arab.125: 1 (1775) ≡Selaginella imbricata(Forssk.) Spring ex Decne.,Arch.Mus.Hist.Nat.2: 193,t.7 (1841) ≡Lycopodioides imbricata(Forssk.) Kuntze,Rev.Gen.Pl.1: 826 (1891).

Pulviniella iridescens(X.C.Zhang&Y.R.Wang)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella iridescensX.C.Zhang&Y.R.Wang,Taxonomy 1: 305.(2021).彩虹垫状卷柏(新拟)

Pulviniella nubigena(J.P.Roux) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella nubigenaJ.P.Roux,Bothalia 38(2):154 (2008).

Pulviniella orientali-chinensis(Ching&C.F.Zhang ex Hao Wei Wang &W.B.Liao) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella orientali-chinensisChing &C.F.Zhang ex Hao Wei Wang &W.B.Liao,Acta Sci.Nat.Univ.Sunyatsenia 61(2): 305(2022).华东垫状卷柏(新拟)

Pulvinae pilifera(A.Braun) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella piliferaA.Braun,Sitz.Ges.Naturf.Freunde,Jan.1857 ex Index 1866,p.2 ≡Lycopodioides pilifera(A.Braun)Kuntze,Rev.Gen.Pl.1: 827 (1891).

Pulviniella pseudotamariscina(X.C.Zhang &C.W.Chen) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella pseudotamariscinaX.C.Zhang &C.W.Chen,Guihaia: 42 (10): 1635(2022).

Pulviniella pulvinata(Hook.&Grev.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium pulvinatumHook.&Grev.,J.Bot.(Hook.)Kew Misc.2:381(1831)≡Selaginella pulvinata(Hook.&Grev.) Maxim.,Mém.Ac.Imp.Sci.St.Pétersb.9: 335 (1859) ≡Lycopodioides pulvinata(Hook.&Grev.)H.S.Kung,Fl.Sichuan.6:64,t.18,1-3 (1988) ≡Selaginella tamariscinavar.pulvinata(Hook.&Grev.) Alston,Bull.Fan Mem.Inst.Biol.Bot.5: 271 (1934).垫状卷柏(新拟)

Pulviniella qinbashanica(Jie Yang bis &X.C.Zhang) Li Bing Zhang &X.M.Zhou,comb.&stat.nov.Basionym:Selaginella pulvinatasubsp.qinbashanicaJie Yang bis &X.C.Zhang,Taxon 72:16-17 (2023).秦巴垫状卷柏(新拟)

Pulviniella stauntoniana(Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella stauntonianaSpring,Mém.Acad.Roy.Sci.Belg.24: 71 (1850) ≡Lycopodioides stauntoniana(Spring)Kuntze,Rev.Gen.Pl.1: 827 (1891).旱生垫状卷柏(新拟)

Pulviniella tamariscina(P.Beauv.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Stachygynandrum tamariscinumP.Beauv.,Mag.Encycl.9(5):483(1804),also Prodr.aethéogam.106(1805)≡Lycopodium tamariscinum(P.Beauv.) Desv.ex Poir.,Lam.,Encycl.suppl.3:540.(1813 publ.1814)≡Selaginella tamariscina(P.Beauv.)Spring,Bull.Acad.Roy.Soc.Bruxelles 10: 136,no.9 (1843) ≡Lycopodioides tamariscina(P.Beauv.)H.S.Kung,Fl.Sichuan.6:62-64,t.18,4-6 (1988).卷柏

5.7.Subfamily VI.Lycopodioidoideae

Subfamily VI.LycopodioidoideaeLi Bing Zhang &X.M.Zhou,subfam.nov.异穗卷柏亚科(新拟) -Type:LycopodioidesBoehm.

Plants creeping (Fig.8B,D-H,K),prostrate,ascending(Fig.8E),scandent (Fig.8L) or erect (Fig.8A,I,J),with ventral rhizophores(Fig.8D,G);sterile leaves dimorphic (Fig.8),four rows;sporophylls monomorphic(Fig.9E,H,J)or dimorphic[nonresupinate (Fig.9F and G) or resupinate (Fig.9B-D,I)];megaspore surfaces often verrucate to tuberculate,few reticulate,smooth (Fig.9K-N,S-V);microspore surfaces can be cristate,verrucate,granulate,laminate,blunt spiny,baculate,or smooth,depending on different genera (Fig.9O-R,W,Y,Z);2n=16,18,20,32 (Jermy,1967;Loyal and Kumar,1984;Takamiya,1993;Mukhopadhyay and Goswami,1996;Marcon et al.,2005).

This subfamily corresponds toSelaginellasubg.Heterostachyssensu Zhou and Zhang (2015).

Lycopodioidoideae comprise those species with dimorphic or slightly dimorphic and resupinate sporophylls (Hypopterygiopsis),dimorphic and non-resupinate sporophylls (most species ofLycopodioides),or monomorphic sporophylls(Valdespinoaand nearly all species ofDidiclis).

Traditionally,Selaginellasubg.Heterostachysonly contains those species with dimorphic and resupinate sporophylls inSelaginella(e.g.,Baker,1883,1887;Warburg,1900;Walton and Alston,1938;Jermy,1986,1990)before our classification(Zhou and Zhang,2015).However,morphological analysis shows that species with dimorphic sporophylls are not monophyletic (Zhou et al.,2016) and dimorphic sporophylls have evolved at least 13 times in the family(Fig.2G).

Based on our study on the phylogeny,gross morphology,spore morphology,and geographic distribution,all species with dimorphic sporophylls and resupinate strobili in the Old World(Asia,Pacific islands,Africa,and Madagascar) should belong to Lycopodioidoideae.However,based on the morphology of megaspores and microspores and molecular data,those species with dimorphic sporophylls and resupinate strobili in the New World (mainly in South and Central Americas) are members of Selaginelloideae (see below).The dimorphic sporophylls and resupinate strobili evolved many times independently in Lycopodioidoideae and Selaginelloideae (Fig.2G and H).

Lycopodioidoideae known so far (Zhou et al.,2022) have the most complicated and diverse plastome structures in Selaginellaceae.Two genera (LycopodioidesandValdespinoa)have three ribosomal operon copies and the largest plastome size in Selaginellaceae.All of known plastome structures of Lycopodioidoideae are DR-IR coexisting structure (Zhou et al.,2022).

Lycopodioidoideae contain over 260 species in the Old World except one American species (Didiclis hoffmannii) in four genera,Didiclis,Hypopterygiopsis,Lycopodioides,andValdespinoa,occurring in the Old World and New World.

5.7.1.Didiclis

DidiclisP.Beauv.ex Mirb.,Hist.Nat.Vég.3:477,4:314(1802)瘤孢卷柏属(新拟) -Type:Didiclis ornithopodioides(L.) P.Beauv.ex J.St.-Hil(Selaginella ornithopodioides(L.).Spring,Flora 21(14):216(1838)).

=Selaginellaser.SuberosaeWarb.in Monsunia 1: 110 (1900) -Type:Selaginella suberosaSpring.

=Selaginella(unranked)ErectaeA.Braun in App.Ind.Sem.Hort.Berol.11([1857]1858)-Type:Selaginella inaequalifoliaSpring.

=Selaginella(unranked)ScandentesA.Braun in App.Ind.Sem.Hort.Berol.11([1857]1858)≡Selaginellaser.Scandentes(A.Braun)Baker in J.Bot.Lond.21: 4 (1883) -Lectotype (designated here):Selaginella laevigataSpring(=S.willdenowii(Desv.ex Poir.)Baker).

=Selaginellaser.SarmentosaeBaker in J.Bot.Lond.21:4(1883)-Lectotype (designated by Zhou and Zhang,2015: 1135):Selaginella inaequalifoliaSpring.

=Selaginellasubser.PleurophyllaeWarb.in Monsunia 1: 106(1900) -Lectotype (designated by Zhou and Zhang,2015: 1135):Selaginella willdenowii(Desv.ex Poir.) Baker.

=Selaginellaser.BisulcataeWarb.in Monsunia 1: 108 (1900) -Type:Selaginella bisulcataSpring.

=Selaginellaser.PleiostelicaeHieron.&Sadeb.in Engler &Prantl,Nat.Pflanzenf.1(4): 700.(1902) ≡Lycopodioidessect.Pleiostelica(Hieron.&Sadeb.)Rothm.in Feddes Repert.Spec.Nov.Regni Veg.54: 69 (1944),“Pleiostele” -Lectotype (designated by Rothmaler,1944: 69):Selaginella uncinata(Desv.ex Poir.) Spring.

Didicliscorresponds to “Selaginellasect.Oligomacrosporangiatae”sensu Zhou and Zhang(2015).The type ofSelaginellasect.Oligomacrosporangiatae,S.fissidentoides,turned out to be a member of theSinensisgroup (nowKorallia) shown by Weststrand and Korall (2016a).The type shares one megasporangium only at the base of strobili with theSinensisgroup(Quansah,1988;Stefanović et al.,1997;also see above).

Although species ofDidiclisshow high diversity of gross morphology,they share tuberculate or verrucate megaspores and blunt-spiny to lamellate or cristate microspores(Zhou et al.,2015;Wang et al.,2018).

Comparing with other subfamilies and genera,Didiclisknown so far has the most diverse master conformations ranging from 2 to 10 in their plastomes(Zhou et al.,2022).

Didicliscontainsca.80 species from Asia and some species from Africa (e.g.,Didiclis pervillei) and one species from America(D.hoffmannii) in the subfamily (Fig.S3).

Based on the morphology and phylogeny,Didicliscan be divided into six well identifiablegroups(theBrauniigroup,theBisulcatagroup,theDelicatulagroup,thePervilleigroup,theSiamensisgroup,and theWilledenowiigroup)corresponding to six subclades in the phylogeny.These six groups each are distinguishable fromHypopterigyopsis.

(1) TheBrauniigroup:This group is characterized by xerophytic and erect plants with monomorphic sterile leaves on stem,broadly ovate sporophylls,and sterile leaves usually involute when dry.Based on our field observations and specimen examination,this group contains about five species (Didiclis braunii,D.fulcrata,D.mairei,D.ostenfieldii,andD.pubescens)in South and Southeast Asia.

(2) TheDelicatulagroup:This group is similar to theWilledenowiigroup in having nearly entire leaves (exceptD.hoffmanniifrom America),erect and suberect plants,but has blunt-spiny ornamentation on microspores.This group contains more than seven species (Didiclis caudata,D.delicatula,D.picta,D.plana,D.mayeri,D.stipulata,andD.wallichii).

(3) TheBisulcatagroup: This group is characterized by resupinate and extremely complanate strobili with dimorphic sporophylls,obovate dorsal leaves,and nearly smooth surfaces on megaspores.Dimorphic sporophylls and resupinate strobili evolved independently or reversed in this group.This group contains about five species (Didiclis bisulcata,D.obovata,D.opaca,D.pennata,andD.soyauxii)in South and Southeast Asia.

(4) ThePervilleigroup: This group is very similar to theBrauniigroup in having xerophytic habit,erect and pubescent plants,and involute leaves when dry,but species of this group have verrucate ornamentation on microspores and are only distributed in Madagascar and South Africa.This group contains about three species(Didiclis eublepharis,D.pervillei,andD.vogelii).

(5) TheSiamensisgroup: This group has long-creeping,ascending or scandent plants.It is similar to some species in theWilledenowiigroup,but has leaf margins ciliolate (vs.entire in theWilledenowiigroup),only one vascular bundle(vs.usually 3 vascular bundles in theWilledenowiigroup),and slightly xerophytic habit.This group contains only one species,Didiclis siamensis,in Southeast Asia.

(6) TheWilledenowiigroup:This group is easily distinguished in having a large plant size (up to 1 m),scandent habit,entire leaves,and lamellate to cristate ornamentation on microspores.This group containsca.60 species,the most of species inDidiclis(e.g.,D.bamleri,D.helferi,D.limbata,D.pseudopaleifera,D.uncinata,D.viridangula,D.willdenowii,etc.) (Chen et al.,2017;Zhang et al.,2013).

These groups might deserve some taxonomical rank,but more samples need to be included and detailed morphological studies are necessary.

Members:

Didiclis axillifolia(Alderw.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella axillifoliaAlderw.,Bull.Jard.Bot.Buitenz.II,11: 37 (1913).

Didiclis bamleri(Hieron.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella bamleriHieron.,Engl.Bot.Jahrb.56:239,no.51 (1920).

Didiclis bisulcata(Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella bisulcataSpring,Mém.Acad.Roy.Sci.Belg.24:259(1850)≡Lycopodioides bisulcata(Spring)Kuntze,Rev.Gen.Pl.1: 826 (1891).双沟瘤孢卷柏(新拟)

Didiclis braunii(Baker) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella brauniiBaker,Gard.Chr.(1867):783,1120,no.24 ≡Lycopodioides braunii(Baker) Kuntze,Rev.Gen.Pl.1: 826(1891).布朗瘤孢卷柏(新拟)

Didiclis buergersiana(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella buergersianaHieron.,Engl.Bot.Jahrb.56: 231,no.30 (1920).

Didiclis canaliculata(L.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium canaliculatumL.,Sp.Pl.2:1105(1753)≡Stachygynandrum canaliculatum(L.) P.Beauv.,Prodr.aethéogam.110(1805)≡Selaginellacanaliculata(L.)Spring,Flora21:201,no.21(1838)≡Lycopodioides canaliculata(L.)Kuntze,Rev.Gen.Pl.1:824(1891).

Didiclis caudata(Desv.S)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Lycopodium caudatumDesv.,Lam.,Encycl.suppl.3: 558(1813 publ.1814) ≡Selaginella caudata(Desv.) Spring,Bull.Acad.Roy.Soc.Bruxelles 10: 144 (1843) ≡Selaginella canaliculatavar.caudata(Desv.) Warb.,Monsunia 1: 107,122 (1900).

Didiclis commersonii(Spring)Li Bing Zhang&X.M.Zhou,comb.&stat.nov.Basionym:Selaginella caudatavar.commersoniiSpring;Mém.Acad.Roy.Sci.Belgique 24[Monogr.Lyc.2]: 140 (1849).

Didiclis conferta(T.Moore) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella confertaT.Moore,Proc.Roy.Hort.Soc.1:133(1861)≡Lycopodioides conferta(T.Moore)Kuntze,Rev.Gen.Pl.1: 826 (1891)

Didiclis decurrens(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella decurrensHieron.,Engl.&Prantl,Nat.Pfl.1(4): 703,no.7 (1901 publ.1902).

Didiclis delicatula(Desv.ex Poir.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium delicatulumDesv.ex Poir.,Lam.,Encycl.suppl.3: 554,no.99.(1813 publ.1814) ≡Selaginella delicatula(Desv.ex Poir.) Alston,J.Bot.70(838): 282 (1932) ≡Lycopodioides delicatula(Desv.ex Poir.)H.S.Kung,Fl.Sichuan.6:67-68,t.20,1-6 (1988).薄叶瘤孢卷柏(新拟)

Didiclis dixitii(Madhus.&S.Nampy)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella dixitiiMadhus.&S.Nampy,Nordic J.Bot.14(5): 527 (1994).

Didiclis dolichoclada(Alston)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella dolichocladaAlston,J.Bot.70: 64,no.5(1932).

Didiclis engleri(Hieron.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella engleriHieron.in Engl.&Prantl,Nat.Pfl.1(4):704,no.374.(1901 publ.1902).

Didiclis eublepharis(A.Braun ex Hieron.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella eublepharisA.Braun ex Hieron.,Nat.Pfl.1 (4): 677 (1902).

Didiclis finium(Alderw.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:SelaginellafiniumAlderw.,Bull.Jard.Bot.Buitenz.II,16:52 (1914).

Didiclis fulcrata(Buch.-Ham.ex D.Don) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium fulcratumBuch.-Ham.ex D.Don,Prodr.Fl.Nepal.17(1825)≡Selaginella fulcrata(Buch.-Ham.ex D.Don) Spring,Bull.Acad.Roy.Soc.Bruxelles 10: 231 (1843) ≡Lycopodioides fulcrata(Buch.-Ham.ex D.Don)Kuntze,Rev.Gen.Pl.1: 826 (1891).

Didiclis fulvicaulis(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella fulvicaulisHieron.,Hedwigia 50: 29,no.21 (1910).

Didiclis furcillifolia(Hieron.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella furcillifoliaHieron.,Hedwigia 50:31,no.22 (1910).

Didiclis gastrophylla(Warb.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella gastrophyllaWarb.,Monsunia 1:107,121,no.103,t.4,f.D (1900).

Didiclis gaudichaudiana(Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella gaudichaudianaSpring,Bull.Acad.Roy.Soc.Bruxelles 10: 145,no.72 (1843).

Didiclis gracilis(T.Moore) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella gracilisT.Moore,Gard.Chr.25,1:752,2:407 (1886).

Didiclis hainanensis(X.C.Zhang&Noot.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella hainanensisX.C.Zhang &Noot.,Bot.J.Linn.Soc.148(3): 323 (2005).琼海瘤孢卷柏(新拟)

Didiclis helferi(Warb.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella helferiWarb.,Monsunia 1: 107,121,no.106(1900).攀缘瘤孢卷柏(新拟)

Didiclis hewittii(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella hewittiiHieron.,Hedwigia 51:262,no.12(1911 publ.1912).

Didiclis hindsii(Hieron.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella hindsiiHieron.,Engl.Bot.Jahrb.50:2,43,no.18 (1913).

Didiclis hochreutineri(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella hochreutineriHieron.,Ann.Jard.Cons.Bot.Genève 15-16: 228-230 (1912).

Didiclis hoffmannii(Hieron.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella hoffmanniiHieron.,Hedwigia 41: 184(1902).

Didiclis inaequalifolia(Hook.&Grev.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium inequalifoliumHook.&Grev.,J.Bot.(Hook.) Kew Misc.2.391 (1831) ≡Selaginella inaequalifolia(Hook.&Grev.)Spring,Bull.Acad.Roy.Soc.Bruxelles 10:145 (1843) ≡Lycopodioides inaequalifolia(Hook.&Grev.) Kuntze,Rev.Gen.Pl.1: 826 (1891).

Didiclis ingens(Alston) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella ingensAlston,J.Bot.72: 229 (1934).

Didiclis kittyae(Alderw.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella kittyaeAlderw.,Bull.Jard.Bot.Buitenz.II,7:35(1912).

Didiclis lacerata(Warb.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella lacerataWarb.,Monsunia 1: 106,120,no.97(1900).

Didiclis latupana(Alderw.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella latupanaAlderw.,Bull.Jard.Bot.Buitenz.II,16: 54 (1914).

Didiclis limbata(Alston)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella limbataAlston,J.Bot.70(831):62(1932).具边瘤孢卷柏(新拟)

Didiclis lobbii(Veitch ex A.Braun) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella lobbiiVeitch ex A.Braun,Ind.Sem.Hort.Berol.,20 (1858) ≡Lycopodioides lobbii(Veitch ex A.Braun)Kuntze,Rev.Gen.Pl.1: 826 (1891).

Didiclis mairei(H.Lév.) Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella maireiH.Lév.,Sert.Yunnan.299(1916).狭叶瘤孢卷柏(新拟)

Didiclis maxima(Alderw.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella maximaAlderw.,Bull.Jard.Bot.Buitenz.II,16: 53 (1914).

Didiclis mayeri(Hieron.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella mayeriHieron.in Engl.&Prantl,Nat.Pfl.1(4):700,no.343.(1901 publ.1902).

Didiclis megalura(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella megaluraHieron.in Engl.&Prantl,Nat.Pfl.1(4): 702,no.358(1901 publ.1902).

Didiclis megastachya(Baker)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella megastachyaBaker,J.Bot.23:20,no.202(1885)≡Lycopodioides megastachya(Baker)Kuntze,Rev.Gen.Pl.1:826 (1891).

Didiclis obovata(S.Y.Dong) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella obovataS.Y.Dong,Syst.Bot.47(1): 88(2022).倒卵叶瘤孢卷柏(新拟)

Didiclis opaca(Warb.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella opacaWarb.,Monsunia 1: 108,122,no.112(1900).

Didiclis ornithopodioides(L.) Spring Basionym:Lycopodium ornithopodioidesL.,Sp.Pl.2: 1105,no.23 (1753) ≡Selaginella ornithopodioides(L.) Spring,Flora 21: 216,no.25 (1838) ≡Lycopodioides ornithopodioides(L.) Kuntze,Rev.Gen.Pl.1: 824 &827(1891) ≡Stachygynandrum ornithopodioides(L.) P.Beauv.,Prodr.aethéogam.110 (1805).

Didiclis ostenfeldii(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella ostenfeldiiHieron.,Bull.Herb.Boiss.II,5:721,no.98 (1905).

Didiclis padangensis(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella padangensisHieron.,Hedwigia 50:34,no.23 (1910).

Didiclis pennata(D.Don)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Lycopodium pennatumD.Don,Prodr.Fl.Nepal.18(1825)≡Selaginella pennata(D.Don)Spring,Bull.Acad.Roy.Soc.Bruxelles 10: 232 (1843) ≡Lycopodioides pennata(D.Don) Kuntze,Rev.Gen.Pl.2: 827 (1891).拟双沟瘤孢卷柏(新拟)

Didiclis permutata(Hieron.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella permutataHieron.,Hedwigia 50:24,no.19 (1910).

Didiclis pervillei(Spring)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella pervilleiSpring,Mém.Acad.Roy.Sci.Belg.24:169(1850)≡Lycopodioides pervillei(Spring)Kuntze,Rev.Gen.Pl.1:825 (1891).

Didiclis picta(Griff.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium pictumGriff.ex Baker,J.Bot.23:19(1885)≡Selaginella picta(Griff.)A.Braun ex Baker,J.Bot.23(265):19(1885)≡Lycopodioides picta(Griff.)Kuntze,Rev.Gen.Pl.1:827(1891).黑顶瘤孢卷柏(新拟)

Didiclis plana(Desv.ex Poir.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Lycopodium planumDesv.ex Poir.,Lam.,Encycl.suppl.3:554,no.98.(1813 publ.1814)≡Selaginella plana(Desv.ex Poir.) Hieron.,Nat.Pfl.1(4): 703 (1901).

Didiclis polystachya(Warb.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella wallichiivar.polystachyaWarb.,Monsunia 1: 106 (1900) ≡Selaginella polystachya(Warb.) Hieron.in Engl.&Prantl,Nat.Pfl.1(4): 702,no.362.(1901 publ.1902).

Didiclis praetermissa(Alston)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella praetermissaAlston,J.Bot.70:65(1932).

Didiclis pseudopaleifera(Hand.-Mazz.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella pseudopaleiferaHand.-Mazz.,Sitzungsber.Akad.Wiss.Wien,61: 82 (1924).耳叶瘤孢卷柏(新拟)

Didiclis pubescens(Wall.ex Hook.&Grev.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Lycopodium pubescensWall.,Num.List no.123 (1829),nomen,ex Hook.&Grev.,Enum.Fil.J.Bot.(Hook.)Kew Misc.1: 383,no.103 (1831) ≡Selaginella pubescens(Wall.ex Hook.&Grev.)Spring,Bull.Acad.Roy.Soc.Bruxelles 10:225(1843).二歧瘤孢卷柏(新拟)

Didiclis rechingeri(Hieron.ex Rech.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella rechingeriHieron.ex Rech.,Denkschr.Kaiserl.Akad.Wiss.,Wien.Math.-Naturwiss.Kl.89:486(1914).

Didiclis retroflexa(v.A.v.R.)Li Bing Zhang&X.M.Zhou,comb.&stat.nov.Basionym:Selaginella axillifoliavar.retroflexav.A.v.R.,Bull.Jard.Bot.Buit.II,11: 38.(1913).

Didiclis schlechteri(Hieron.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella schlechteriHieron.,Bot.Jahrb.Syst.50:2,41,no.17 (1913).

Didiclis semicordata(Wall.ex Hook.&Grev.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium semicordatumWall.,(Cat.n.126-3,11 &13 (1829),nom.nud.) ex Hook.&Grev.,J.Bot.(Hook.)Kew Misc.2:396(1831)≡Lycopodioides semicordata(Wall.ex Hook.&Grev.) Kuntze,Rev.Gen.Pl.1: 827 (1891) ≡Selaginella semicordata(Wall.ex Hook.&Grev.) Spring,Mart.,Fl.Bras.1 (2):122 (1840).

Didiclis siamensis(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella siamensisHieron.,Bot.Tidsskr.24: 113(1901).泰国瘤孢卷柏(新拟)

Didiclis soyauxii(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella soyauxiiHieron.,Engl.&Prantl,Nat.Pfl.1(4): 697,no.305.(1901 publ.1902).

Didiclis stipulata(Blume) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium stipulatumBlume,Enum.Pl.Jav.2:268,no.18(1830)≡Selaginella stipulata(Blume)Spring,Bull.Acad.Roy.Soc.Bruxelles 10: 145,no.70(1843).

Didiclis subbiflora(Hieron ex Brause) Li Bing Zhang &X.M.Zhou,comb.&stat.nov.Basionym:Selaginella gracilisvar.subbifloraHieron ex Brause,Engl.Bot.Jahrb.56: 238,no.47.(1920).

Didiclis trichoclada(Alston)Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella trichocladaAlston,J.Bot.70: 63,no.3(1932).毛枝瘤孢卷柏(新拟)

Didiclis tylophora(Alderw.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella tylophoraAlderw.,Bull.Jard.Bot.Buitenz.II,16: 50 (1914).

Didiclis uncinata(Desv.ex Poir.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium uncinatumDesv.ex Poir.,Lam.,Encycl.Suppl.3: 558 (1814) ≡Lycopodioides uncinata(Desv.) Kuntze,Rev.Gen.Pl.1:825(1891)≡Selaginella uncinata(Desv.ex Poir.)Spring,Bull.Acad.Roy.Soc.Bruxelles 10: 141 (1843).翠云草

Didiclis usterii(Hieron.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella usteriiHieron.,ex Usteri,Beitr.Phil.Veg.,135(1905),et Leafl.Phil.Bot.6: 2055 (1913).

Didiclis velutina(Ces.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella velutinaCes.,Rec.Ac.Sci.Nap.16:28,31(1877).

Didiclis victoriae(J.W.Moore)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella victoriaeJ.W.Moore,Cat.n.143: 8,t.6(1878)≡Lycopodioides victoriae(J.W.Moore)Kuntze,Rev.Gen.Pl.1:827 (1891).

Didiclis viridangula(Spring)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella viridangulaSpring,v.Heurck,Pl.Nov.,29(1870)≡Lycopodioides viridangula(Spring) Kuntze,Rev.Gen.Pl.1:827 (1891).

Didiclis vogelii(Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella vogeliiSpring,Monogr.Lyc.2: 170,no.111(1850).

Didiclis wallichii(Hook.&Grev.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium wallichiiHook.&Grev.,Enum.Fil.J.Bot.(Hook.) Kew Misc.2: 384,no.106 (1831) ≡Selaginella wallichii(Hook.&Grev.)Spring,Mart.,Fl.Bras.1(2):124(1840).瓦氏瘤孢卷柏(新拟)

Didiclis willdenowii(Desv.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium willdenoviiDesv.ex Poir.,Lam.,Encycl.suppl.3: 540,552,no.87.(1813 publ.1814) ≡Selaginella willdenowii(Desv.) Baker,Gard.Chr.783,950 (1867) ≡Lycopodioides willdenowii(Desv.)Kuntze,Rev.Gen.Pl.1: 827(1891).藤瘤孢卷柏(新拟)

5.7.2.Hypopterygiopsis

HypopterygiopsisSakurai,Bot.Mag.(Tokyo)57:255(1943)异穗卷柏属(新拟) -Type:Hypopterygiopsis reptansSakurai,Bot.Mag.(Tokyo)57: 255 (1943) (≡Selaginella sakuraiiH.A.Mill.).

=Lycopodium(unranked)PlatystachyaHook.&Grev.in Bot.Misc.2:380(1831)≡Selaginella(unranked)Platystachyae(Hook.&Grev.) Spring in Mém.Acad.Sci.Belg.24: 53 (1850) -Lectotype(designated by Zhou and Zhang,2015: 1134):Selaginella chrysocaulos(Grev.&Hook.) Spring.

=Selaginellaser.BrachystachyaeWarb.in Monsunia 1: 110(1900) -Type:Selaginella brachystachya(Hook.&Grev.)Spring.

=Lycopodium(unranked)PlanifoliaHook.&Grev.,Bot.Misc.2:382 (1831) ≡Selaginella(unranked)Planifoliae(Hook.&Grev.)Spring in Martius,Fl.Bras.1(2):118.(1840)-Lectotype(designated by Zhou and Zhang,2015: 1135):Lycopodium tetragonostachyumWall.ex Grev.&Hook.

=Selaginella(unranked)PronaeA.Braun,App.Ind.Sem.Hort.Berol.12([1857]1858)-Lectotype(designated by Zhou and Zhang,2015: 1135):Selaginella ciliarisSpring.

=Selaginellaser.PronifloraeWarb.,Monsunia 1: 108 (1900) -Type:Selaginella proniflora(L.) Baker.

The type ofHypopterygiopsiswas described as a“moss”species(Sakurai,1943) but it turned out to be a Selaginellaceae (Miller,1967).Our examination of the type confirmed this.The type is not sampled in our molecular work and is assigned here based on the morphology.

Plant without creeping rhizome (vs.with creeping rhizome in theDidiclis brauniigroup and theD.pervilleigroup);stem with single vascular bundle(vs.often with three vascular bundles in theD.willedenowiigroup and theD.delicatulagroup);stem creeping to ascending,sometimes erect,never scandent(vs.scandent in theD.willedenowiigroup and theD.siamensisgroup);dorsal leaves rarely obovate (vs.obovate in theD.bisulcatagroup);sporophylls dimorphic or submonomorphic,resupinate,rarely strictly monomorphic(vs.always strictly monomorphic inDidiclis,except for theD.bisulcatagroup;often dimorphic,non-resupinate inLycopodioides);megaspore surfaces often not interconnected verrucate to tuberculate,or finely reticulate(Tetragonostachyaetype-1 and type-2) (vs.megaspore surfaces often interconnected verrucate to tuberculate and never reticulate inDidiclis);microspore surfaces often verrucate to spherulate,rarely smooth to coarse or baculate(vs.microspore surfaces blunt-spiny,lamellate,or cristate in theD.brauniigroup,theD.delicatulagroup,theD.siamensisgroup,and theD.willedenowiigroup).

This genus contains about 170 species in Africa and Asia.Based on the synapomorphy of strobili with dimorphic sporophylls,we combinedSelaginellasect.HeterostachysandS.sect.Tetragonostachyaesensu Zhou and Zhang (2015) to formHypopterygiopsishere.WithinHypopterygiopsis,we recognize two sections.

Hypopterygiopsissect.Hypopterygiopsis藓状异穗卷柏组(新拟)

=Selaginella(unranked)Tetragonostachyae(Hook.&Grev.)Spring in Mém.Acad.Sci.Belg.24: 53 (1850).Basionym:Lycopodium(unranked)TetragonostachyaHook.&Grev.in Bot.Misc.2:382 (1831) ≡S.sect.Tetragonostachyae(Hook.&Grev.) Hieron.&Sadeb.in Engler&Prantl,Nat.Pflanzenf.1(4):669(1902)≡S.sect.Tetragonostachyae(Hook.&Grev.) Rothm.in Feddes Repert.Spec.Nov.Regni Veg.54: 68 (1944),“Tetragonostachya” ≡Lycopodioidessubg.Tetragonostachya(Hook.&Grev.)Tzvel.in Novosti Sist.Vyssh.Rast.36: 25 (2004) -Type:L.tetragonostachyumWall.ex Grev.&Hook.[≡Hypopterygiopsis tetragonostachya(Wall.ex Grev.&Hook.)Li Bing Zhang and X.M.Zhou].

This section corresponds toSelaginellasect.Tetragonostachyaesensu Zhou and Zhang (2015).

Based on the evolution of strobilus morphology,monomorphic sporophylls are reversed in this section.Although some species from Pacific islands have monomorphic to submonomorphic sporophylls,they have same megaspores as those species with dimorphic sporophylls in Asia.Species of this section have megaspore surfaces reticulate(Tetragonostachyaetype-1 and type-2 with very fine muri and open meshes:Fig.9L-N)and usually microspore surfaces smooth (Fig.9P and Q),verrucate (Fig.9O,R) or baculate(S.ciliaris;Zhou et al.,2015).

This section contains two well-supported clades in our phylogenetic analysis.One clade contains species with monomorphic sporophylls (e.g.,Hypopterygiopsis arbuscula,H.kanehirae,H.whitmeei) mainly from Pacific islands (Kato,2008;Chen et al.,2017),and the other contains species with strongly (H.ciliarisandH.xipholepis) or slightly dimorphic (H.repandaandH.vaginata)sporophylls mainly in Asia and some in Africa and Madagascar(Figs.S1-S3)(Quansah and Thomas,1985;Stefanović,1997;Zhang et al.,2013).

Hypopterygiopsissect.Heterostachys(Baker) Li Bing Zhang &X.M.Zhou,comb.nov.异穗卷柏组(新拟) -Basionym:Selaginellasubg.HeterostachysBaker in J.Bot.Lond.21: 3 (1883)≡Selaginellasect.Heterostachys(Baker)Li Bing Zhang&X.M.Zhou,Taxon 64(6):1134 (2015) -Lectotype (designated by Jermy,1986: 118):Hypopterygiopsis heterostachys(Baker) Li Bing Zhang &X.M.Zhou (≡Selaginella heterostachys(Baker).

This section corresponds toSelaginellasect.Heterostachyssensu Zhou and Zhang(2015).This section contains the most species with resupinate strobili and dimorphic sporophylls from Asia and Africa(Quansah and Thomas,1985;Stefanović,1997;Zhang et al.,2013).

Morphologically,Selaginellasect.Heterostachyshas often creeping (Fig.8B and C),sometimes ascending or erect plants(Fig.8A);resupinate strobili and dimorphic sporophylls (Fig.9A);megaspore surfaces papillate,tuberculate,and verrucate(Fig.9S,U;Zhou et al.,2015),sometimes elements connected (Fig.9U);and microspore surfaces often verrucate,gemmulate,and spherulate(Fig.9W-Y),rarely few tuberculate and rugate (Zhou et al.,2015).

Although dimorphic sporophylls and resupinate strobili also exist inHypopterygiopsissect.Hypopterygiopsisand some America species ofSelaginella s.s.(see below),these species of the latter usually have megaspores with reticulate ornamentation (Figs.9 and 10;Korall and Taylor 2006;Bauer et al.,2016;Valdespino,2017c).

Members of the genus:

Hypopterygiopsis abyssinica(Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella abyssinicaSpring,Monogr.Lyc.2: 99,no.44 (1850) ≡Selaginella goudotianavar.abyssinica(Spring)Bizzarri,Webbia 29:585(1975)≡Lycopodioides abyssinica(Spring) Kuntze,Rev.Gen.Pl.1: 825 (1891).

Hypopterygiopsis albociliata(P.S.Wang) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella albociliataP.S.Wang,J.Arnold Arbor.71(2): 269 (1990).白毛异穗卷柏(新拟)

Hypopterygiopsis alutacea(Spring)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella alutaceaSpring,Bull.Acad.Roy.Soc.Bruxelles 10: 233,no.154 (1843) ≡Lycopodioides alutacea(Spring) Kuntze,Rev.Gen.Pl.1: 825 (1891).

Hypopterygiopsis amblyphylla(Alston) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella amblyphyllaAlston,Bull.Fan Mem.Inst.Biol.Bot.5(6): 287-288 (1934).钝叶异穗卷柏(新拟)

Hypopterygiopsis angustifolia(Hieron.ex Jeanpert) Li Bing Zhang &X.M.Zhou,comb.&stat.nov.Basionym:Selaginella ustavar.angustifoliaHieron.ex Jeanpert,Bull.Mus.Paris 17: 579(nomen) (1911).

Hypopterygiopsis antimonanensis(B.C.Tan &Jermy) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella antimonanensisB.C.Tan &Jermy,Fern Gaz.12(3): 170(1981).

Hypopterygiopsis apoensis(Hieron.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella apoensisHieron.,Elmer,Leafl.Phil.Bot.,6: 2023,no.18 (1913).

Hypopterygiopsis arbuscula(Kaulf.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Lycopodium arbusculumKaulf.,Enum.Fil.19(1824) ≡Selaginella arbuscula(Kaulf.) Spring,Bull.Acad.Roy.Soc.Bruxelles 10:227(1843)≡Lycopodioides arbuscula(Kaulf.)Kuntze,Rev.Gen.Pl.1: 825 (1891).

Hypopterygiopsis arbusculoides(J.W.Moore) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella arbusculoidesJ.W.Moore,Bull.Bernice P.Bishop Mus.102: 15 (1933).

Hypopterygiopsis aristata(Spring)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella aristataSpring,Bull.Acad.Roy.Soc.Bruxelles 10: 232,no.152 (1843) ≡Selaginella circinalisvar.aristataC.Presl,Bot.Bem.153 (1845).

Hypopterygiopsis atimonanensis(B.C.Tan &Jermy) Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella atimonanensisB.C.Tan &Jermy,Fern Gaz.12(3): 170(1981).

Hypopterygiopsis auquieri(Bizzarri) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella auquieriBizzarri,Bull.Jard.Bot.Belg.51(1-2): 220 (1981).

Hypopterygiopsis austro-orientalis(H.J.Wei &X.M.Zhou) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella austroorientalisH.J.Wei&X.M.Zhou;Phytotaxa 579(2):91(2023).东南异穗卷柏(新拟)

Hypopterygiopsis bankii(Alston) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella banksiiAlston;Bull.Bernice P.Bishop Mus.93: 15,83 (1932).

Hypopterygiopsis beccariana(Baker) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella beccarianaBaker,J.Bot.23:154,no.254(1885)≡Lycopodioides beccariana(Baker)Kuntze,Rev.Gen.Pl.1: 826 (1891).

Hypopterygiopsis behrmanniana(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella behrmannianaHieron.,Engl.Bot.Jahrb.56: 233(1920).

Hypopterygiopsis bemarahensis(Stefanović &Rakotondr.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella bemarahensisStefanović &Rakotondr.,Novon 6(2): 203 (1996).

Hypopterygiopsis blepharophylla(Alston)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella blepharophyllaAlston,Mém.Inst.Fr.Afr.Noire 50: 40,t.6,ff.9-15 (1957).

Hypopterygiopsis bodinieri(Hieron.ex Christ)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella bodinieriHieron.ex Christ,Bull.Ac.Géogr.Bot.Mans 11:273,no.109(1902),nomen,et Hedwigia 43: 6,no.25 (1904).大叶异穗卷柏(新拟)

Hypopterygiopsis boninensis(Baker) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella boninensisBaker,J.Bot.23(270): 178 (1885) ≡Lycopodioides boninensis(Baker) Kuntze,Rev.Gen.Pl.1: 826 (1891).小笠原异穗卷柏(新拟)

Hypopterygiopsis brachystachya(Hook.&Grev.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Lycopodium brachystachyumHook.&Grev.,J.Bot.(Hook.) Kew Misc.3:107 (1833) ≡Selaginella brachystachya(Hook.&Grev.)Spring,Bull.Acad.Roy.Sci.Bruxelles 10: 232 (1843) ≡Lycopodioides brachystachya(Hook.&Grev.)Kuntze,Rev.Gen.Pl.1: 826 (1891).

Hypopterygiopsis buchholzii(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella buchholziiHieron.in Engl.&Prantl,Nat.Pfl.1(4): 696,no.272 (1901 publ.1902).

Hypopterygiopsis burbidgei(Baker)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella burbidgeiBaker,J.Bot.23:154,no.253 (1885).

Hypopterygiopsis busuensis(Alderw.) Li Bing Zhang &X.M.Zhou,comb.&stat.nov.Basionym:Selaginella darmandvilleivar.busuensisAlderw.;Bull.Jard.Bot.Buitenzorg,sér.2,16.50 (1914).

Hypopterygiopsis calcicola(Hieron.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella calcicolaHieron.,Hedwigia 51:258,no.9 ((1911) 1912).

Hypopterygiopsis calostachya(Hook.&Grev.) Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Lycopodium calostachyumHook.&Grev.,Enum.Fil.J.Bot.(Hook.)Kew Misc.3:108(1833)≡Selaginella calostachya(Hook.&Grev.) Alston,J.Bot.70:65,no.8 (1932).

Hypopterygiopsis cataractarum(Alston) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella cataractarumAlston,Proc.Nat.Inst.Sci.India 11: 228 (1945).

Hypopterygiopsis chaetoloma(Alston) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella chaetolomaAlston,J.Bot.70:67,no.12 (1932).毛边异穗卷柏(新拟)

Hypopterygiopsis chaii(Jermy) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella chaiiJermy,Kew Bull.41(3): 551(1986).

Hypopterygiopsis chingii(Alston) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella chingiiAlston,J.Bot.70:66,no.10(1932).秦氏异穗卷柏(新拟)

Hypopterygiopsis chrysocaulos(Hook.&Grev.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Lycopodium chrysocaulonHook.&Grev.,Enum.Fil.J.Bot.(Hook.) Kew Misc.2: 401,no.182 (1831) ≡Selaginella chrysocaulos(Hook.&Grev.)Spring,Bull.Acad.Roy.Soc.Bruxelles 10: 232 (1843) ≡Lycopodioides chrysocaulos(Hook.&Grev.) H.S.Kung,Fl.Sichuan.6: 78 (1988).块茎异穗卷柏(新拟)

Hypopterygiopsis chrysorrhizos(Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella chrysorrhizosSpring,Mém.Acad.Roy.Sci.Belg.24 (2): 251 (1850) ≡Lycopodioides chrysorhiza(Spring) Kuntze,Rev.Gen.Pl.1: 826 (1891).

Hypopterygiopsis ciliaris(Retz.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium ciliareRetz.,Obs.Bot.5:(1789)≡Selaginella ciliaris(Retz.) Spring,Bull.Acad.Roy.Soc.Bruxelles 10:231 (1843) ≡Lycopodioides ciliaris(Retz.) Kuntze,Rev.Gen.Pl.1:826 (1891).睫毛异穗卷柏(新拟)

Hypopterygiopsis ciliata(Alston) Li Bing Zhang &X.M.Zhou,comb.&stat.nov.Basionym:Selaginella selangorensisvar.ciliataAlston,Gard.Bull.Straits Settlem.8: 44 (1934).

Hypopterygiopsis cochleata(Hook.&Grev.)Zhang&X.M.Zhou,comb.nov.Basionym:Lycopodium cochleatumHook.&Grev.,Enum.Fil.J.Bot.(Hook.)Kew Misc.2:395,no.153(1831) ≡Lycopodioides cochleata(Hook.&Grev.) Kuntze,Rev.Gen.Pl.1: 826 (1891) ≡Selaginella cochleata(Hook.&Grev.) Spring,Bull.Acad.Roy.Soc.Bruxelles 10: 143,no.55 (1843).

Hypopterygiopsis compta(Hand.-Mazz.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella comptaHand.-Mazz.,Symb.Sin.6: 9,pl.2.(1929).缘毛异穗卷柏(新拟)

Hypopterygiopsis coonooriana(R.D.Dixit)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella coonoorianaR.D.Dixit,Bull.Bot.Surv.India 25(1-4): 223 (1983 publ.1985).

Hypopterygiopsis coriaceifolia(X.M.Zhou,N.T.Lu &Li Bing Zhang)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella coriaceifoliaX.M.Zhou,N.T.Lu &Li Bing Zhang,Phytotaxa 453(2):125 (2020).

Hypopterygiopsis crassipes(Spring)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella crassipesSpring,Monogr.Lyc.2:243,no.181 (1850) ≡Lycopodioides crassipes(Spring) Kuntze,Rev.Gen.Pl.1: 826 (1891).

Hypopterygiopsis curtisii(Ridl.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella curtisiiRidl.,J.Roy.As.Soc.,Str.Br.,No.80: 148,no.2 (1919).

Hypopterygiopsis daozhenensis(Li Bing Zhang,Q.W.Sun&Jun H.Zhao) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella daozhenensisLi Bing Zhang,Q.W.Sun&Jun H.Zhao,Phytotaxa 207(2): 187 (2015).道真异穗卷柏(新拟)

Hypopterygiopsis darmandvillei(Alderw.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella darmandvilleiAlderw.,Bull.Jard.Bot.Buitenz.II,1: 23-24(1911).

Hypopterygiopsis decipiens(Warb.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella decipiensWarb.,Monsunia 1:127(1900).拟大叶异穗卷柏(新拟)

Hypopterygiopsis densiciliata(X.M.Zhou,Liang Zhang &Bo Xu) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella densiciliataX.M.Zhou,Liang Zhang &Bo Xu,PhytoKeys 227:135-149 (2023).密毛异穗卷柏(新拟)

Hypopterygiopsis devolii(H.M.Chang,P.F.Lu &W.L.Chiou) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella devoliiH.M.Chang,P.F.Lu&W.L.Chiou,Blumea 56(1):21(2011).棣氏异穗卷柏(新拟)

Hypopterygiopsis dianzhongensis(X.C.Zhang) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella dianzhongensisX.C.Zhang,PhytoKeys 118: 77 (2019).滇中异穗卷柏(新拟)

Hypopterygiopsis drepanophylla(Alston)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella drepanophyllaAlston,J.Bot.70(831): 66-67(1932).镰叶异穗卷柏(新拟)

Hypopterygiopsis dulongjiangensis(W.M.Chu)Li Bing Zhang&X.M.Zhou,comb.&stat.nov.Basionym:Selaginella effusavar.dulongjiangensisW.M.Chu,Fl.Yunnan.20: 718 (2006).独龙江异穗卷柏(新拟)

Hypopterygiopsis effusa(Alston) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella effusaAlston,J.Bot.70: 65,no.9(1932).疏松异穗卷柏(新拟)

Hypopterygiopsis elegantissima(Warb.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella elegantissimaWarb.,Monsunia 1: 111,no.175 (1900).

Hypopterygiopsis eschscholzii(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella eschscholziiHieron.,Elmer,Leafl.Phil.Bot.6: 2041,no.26 (1913).

Hypopterygiopsis exasperata(Warb.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella exasperataWarb.,Monsunia 1: 109,126,no.139 (1900).

Hypopterygiopsis firmula(A.Braun ex Kuhn) Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:SelaginellafirmulaA.Braun ex Kuhn;Verh.K.K.Zool.-Bot.Ges.Wien 19: 585,no.132(1869).

Hypopterygiopsis goudotiana(Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella goudotianaSpring,Bull.Acad.Roy.Soc.Bruxelles 8 (12): 140 (1841) ≡Lycopodioides goudotiana(Spring) Kuntze,Rev.Gen.Pl.1: 826 (1891).

Hypopterygiopsis heterostachys(Baker) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella heterostachysBaker,J.Bot.23: 177,no.264 (1885) ≡Lycopodioides heterostachya(Baker)Kuntze,Rev.Gen.Pl.1: 826 (1891).异穗卷柏.

Hypopterygiopsis hildebrandtii(A.Braun ex Kuhn) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella hildebrandtiiA.Braun ex Kuhn,v.Decken,Reis.III,3,Bot.:71,no.88(1879).

Hypopterygiopsis hollrungii(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella hollrungiiHieron.Engl.Bot.Jahrb.50: 35,no.14 (1913).

Hypopterygiopsis intertexta(Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella intertextaSpring,Bull.Acad.Roy.Soc.Bruxelles 10: 233,no.156 (1843).

Hypopterygiopsis jainii(R.D.Dixit)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella jainiiR.D.Dixit,Bull.Bot.Surv.India 25: 225,f.2 A-H (1985).

Hypopterygiopsis kaernbachii(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella kaernbachiiHieron.in Engl.&Prantl,Nat.Pfl.1(4): 700,no.335 (1901 publ.1902).

Hypopterygiopsis kalbreyeri(Baker)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella kalbreyeriBaker,J.Bot.22:276,no.157 (1884).

Hypopterygiopsis kanehirae(Alston) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella kanehiraeAlston,J.Bot.72:227 (1934).

Hypopterygiopsis keralensis(R.D.Dixit) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella keralensisR.D.Dixit,Bull.Bot.Surv.India 27 (1-4): 123-124.1987 (“1986”).

Hypopterygiopsis ketra-ayam(Alderw.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella ketra-ayamAlderw.,Bull.Jard.Bot.Buitenz.II,1: 24-25 (1911).

Hypopterygiopsis kivuensis(Bizzarri) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella kivuensisBizzarri,Bull.Jard.Bot.Belg.53(1-2): 174 (1983).

Hypopterygiopsis kouytcheensis(H.Lév.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella kouytcheensisH.Lév.,Fedde,Repert.9 (222-226): 451 (1911).贵州异穗卷柏(新拟)

Hypopterygiopsis kurzii(Baker) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella kurziiBaker,J.Bot.23: 249,no.283 (1885) ≡Lycopodioides kurzii(Baker) Kuntze,Rev.Gen.Pl.1:826 (1891).缅甸异穗卷柏(新拟)

Hypopterygiopsis labordei(Hieron.ex Christ) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella labordeiHieron.ex Christ,Bull.Ac.Inst.Géogr.Bot.III,11: 272,no.168 (1902) ≡Lycopodioides labordei(Hieron.ex Christ) H.S.Kung,Fl.Sichuan.6:77-78 (1988).细叶异穗卷柏(新拟)

Hypopterygiopsis lanceolata(Warb.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella lanceolataWarb.,Monsunia 1: 108,123,no.117 (1900).

Hypopterygiopsis lauterbachii(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella lauterbachiiHieron.,Bot.Jahrb.Syst.50: 31,no.12 (1913).

Hypopterygiopsis laxa(Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella laxaSpring,Bull.Acad.Roy.Soc.Bruxelles 10: 233,no.153 (1843) ≡Lycopodioides laxa(Spring)Kuntze,Rev.Gen.Pl.1: 826 (1891).

Hypopterygiopsis leoneensis(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella leoneensisHieron.,Engl.&Prantl,Nat.Pfl.1(4): 697(1901 publ.1902).

Hypopterygiopsis leptophylla(Baker) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella leptophyllaBaker,J.Bot.23(269): 157 (1885) ≡Lycopodioides leptophylla(Baker) Kuntze,Rev.Gen.Pl.1: 826 (1891).膜叶异穗卷柏(新拟)

Hypopterygiopsis lewalleana(Bizzarri) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella lewalleanaBizzarri,Bull.Jard.Bot.Belg.51(1-2): 222 (1981).

Hypopterygiopsis lindhardtii(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella lindhardtiiHieron.,Bull.Herb.Boiss.II,5: 723,no.99 (1905).

Hypopterygiopsis llanosii(Hieron.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella llanosiiHieron.,Elmer,Leafl.Phil.Bot.6: 2039,no.(1913).

Hypopterygiopsis longiciliata(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella longiciliataHieron.,Engl.,Bot.Jahrb.50: 2,33,no.13 (1913).

Hypopterygiopsis loriai(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella loriaiHieron.,Engl.Bot.Jahrb.50:2,27,no.10 (1913).

Hypopterygiopsis lutchuensis(Koidz.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella lutchuensisKoidz.,Pl.Nov.Amamio-Oshimensis,Ins.Adjasc.,4 (1928),et Acta Phytotax.Geobot.1: 165 (1932).琉球异穗卷柏(新拟)

Hypopterygiopsis macroblepharis(Warb.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella macroblepharisWarb.,Monsunia 1: 108,124,no.126,t.3C (1900).

Hypopterygiopsis mannii(Baker) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella manniiBaker,J.Bot.23: 180,no.277 (1885) ≡Lycopodioides mannii(Baker) Kuntze,Rev.Gen.Pl.1:826 (1891).

Hypopterygiopsis marinii(Stefanović &Rakotondr.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella mariniiStefanović &Rakotondr.,Novon 6(2): 205 (1996).

Hypopterygiopsis megaphylla(Baker) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella megaphyllaBaker,J.Bot.23:180(1885)≡Lycopodioides megaphylla(Baker)Kuntze,Rev.Gen.Pl.1: 826 (1891).宽叶异穗卷柏(新拟)

Hypopterygiopsis menziesii(Hook.&Grev.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium menziesiiHook.&Grev.,Enum.Fil.1: 2,no.131 (1831) ≡Selaginella menziesii(Hook.&Grev.) Spring,Bull.Acad.Roy.Soc.Bruxelles 10: 227,no.109 (1843) ≡Lycopodioides menziesii(Hook.&Grev.) Kuntze,Rev.Gen.Pl.1: 826 (1891) ≡Selaginella arbusculavar.menziesii(Hook.&Grev.) Skottsb.,Medd.Göteborgs Bot.Trädgard,15: 145(1944).

Hypopterygiopsis miniatospora(Dalzell) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium miniatosporumDalzell,J.Bot.(Hook.)4:114(1852)≡Selaginella miniatospora(Dalzell)Baker,J.Bot.23:249(1885)≡Lycopodioides miniatospora(Dalzell)Kuntze,Rev.Gen.Pl.1: 826 (1891).

Hypopterygiopsis minutifolia(Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella minutifoliaSpring,Monogr.Lyc.2: 239,no.176 (1850) ≡Lycopodioides minutifolia(Spring)Kuntze,Rev.Gen.Pl.1: 826 (1891).小叶异穗卷柏(新拟)

Hypopterygiopsis mittenii(Baker) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella mitteniiBaker,J.Bot.(London)21:18 (1883).

Hypopterygiopsis molleri(Hieron.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella molleriHieron.in Engl.&Prantl,Nat.Pfl.1(4): 697,no.292.(1901 publ.1902).

Hypopterygiopsis molliceps(Spring)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella mollicepsSpring,Mém.Acad.Roy.Sci.Belg.24:257(1850)≡Lycopodioides molliceps(Spring)Kuntze,Rev.Gen.Pl.1: 826 (1891).

Hypopterygiopsis monodii(Alston)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella monodiiAlston,Bull.Inst.Franc.Afr.Noire,A,21: 440 (1959).

Hypopterygiopsis monospora(Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium monosporum(Spring)Hook.,Bot.Misc.9: 362 (1857) ≡Selaginella monosporaSpring,Mém.Acad.Roy.Sci.Belg.24:135(1850)≡Selaginella plumosavar.monospora(Spring)Baker,J.Bot.21:145(1883).单子异穗卷柏(新拟)

Hypopterygiopsis morganii(Zeiller)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella morganiiZeiller,Bull.Soc.Bot.France 32: 78 (1985).

Hypopterygiopsis myosuroides(Kaulf.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium myosuroidesKaulf.,Enum.Fil.19 (1824) ≡Selaginella myosuroides(Kaulf.) Spring,Bull.Acad.Roy.Soc.Bruxelles 10: 232,no.150 (1843) ≡Lycopodioides myosuroides(Kaulf.) Kuntze,Rev.Gen.Pl.1: 827 (1891).

Hypopterygiopsis nairii(R.D.Dixit)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella nairiiR.D.Dixit,Bull.Bot.Surv.India 26: 106 (1985).

Hypopterygiopsis namdaphaensis(Sarn.Singh &Panigrahi) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella namdaphaensisSarn.Singh &Panigrahi,Ferns &Fern-Allies of Arunachal Pradesh 1: 64.(2005).

Hypopterygiopsis nana(Desv.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium nanumDesv.ex Poir.,Lam.,Encycl.suppl.3: 554,no.96.(1813 publ.1814) ≡Selaginella nana(Desv.) Spring,Bull.Acad.Roy.Sci.Bruxelles 10: 232,no.151(1843) ≡Lycopodioides nana(Spring) Kuntze,Rev.Gen.Pl.1: 827(1891).

Hypopterygiopsis nayarii(R.D.Dixit) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella nayariiR.D.Dixit,Bull.Bot.Surv.India 27(1-4): 123 (1987).

Hypopterygiopsis neocaledonica(Baker) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella neocaledonicaBaker,J.Bot.22: 245,no.143 (1884) ≡Lycopodioides neocaledonica(Baker) Kuntze,Rev.Gen.Pl.1: 827 (1891).

Hypopterygiopsis nummularia(Warb.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella nummulariaWarb.,Monsunia 1: 108,123,no.121 (1900).

Hypopterygiopsis ornata(Hook.&Grev.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Lycopodium ornatumHook.&Grev.,Enum.Fil.J.Bot.(Hook.) Kew Misc.3: 108 (1883) ≡Selaginella ornata(Hook.&Grev.)Spring,Bull.Acad.Roy.Soc.Bruxelles 10:232(1843)≡Selaginella brachystachyavar.ornata(Hook.&Grev.)Baker,J.Bot.23: 180 (1885).微齿异穗卷柏(新拟)

Hypopterygiopsis panchghaniana(R.D.Dixit) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella panchghanianaR.D.Dixit,Bull.Bot.Surv.India 25: 226 (1985).

Hypopterygiopsis panigrahii(R.D.Dixit) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella panigrahiiR.D.Dixit,Bull.Bot.Surv.India 25: 226,f.4.A-H (1985).

Hypopterygiopsis parachrysocaulos(M.H.Zhang&X.C.Zhang)Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella parachrysocaulosM.H.Zhang &X.C.Zhang,Taxon 71(6): 1167 (2022).拟块茎异穗卷柏(新拟)

Hypopterygiopsis perpusilla(Baker)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella perpusillaBaker,J.Bot.23: 292(1885)≡Lycopodioides perpusilla(Baker)Kuntze,Rev.Gen.Pl.1:827(1891).

Hypopterygiopsis phanotricha(Baker) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella phanotrichaBaker,J.Bot.23:156,no.261 (1885) ≡Lycopodioides phanotricha(Baker) Kuntze,Rev.Gen.Pl.1: 827 (1891).

Hypopterygiopsis philippina(Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella philippinaSpring,Bull.Acad.Roy.Soc.Bruxelles 10: 140 (1843) ≡Lycopodioides philippina(Spring) Kuntze,Rev.Gen.Pl.1: 827 (1891).

Hypopterygiopsis pricei(B.C.Tan&Jermy)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella priceiB.C.Tan&Jermy,Fern Gaz.12(3): 170(1981).

Hypopterygiopsis proniflora(Lam.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Lycopodium proniflorumLam.,Encycl.3:652,no.25.(1789 publ.1791)≡Selaginella proniflora(Lam.)Baker,J.Bot.22: 156 (1885) ≡Stachygynandrum proniflorum(Lam.) P.Beauv.,Prodr.aethéogam.110 (1805) ≡Lycopodioides proniflora(Lam.)Kuntze,Rev.Gen.Pl.1: 827 (1891).

Hypopterygiopsis protensa(Alston)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella protensaAlston,Mém.Inst.Fr.Afr.Noire 50: 41 (1957).

Hypopterygiopsis qingchengshanensis(Li Bing Zhang &X.M.Zhou)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella qingchengshanensisLi Bing Zhang &X.M.Zhou,Phytotaxa 522(4):286 (2021).青城山异穗卷柏(新拟)

Hypopterygiopsis radicata(Hook.&Grev.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Lycopodium radicatumHook.&Grev.,Enum.Fil.J.Bot.(Hook.) Kew Misc.2: 397,no.160 (1831) ≡Selaginella radicata(Hook.&Grev.)Spring,Mém.Acad.Roy.Sci.Belg.24(2) 114 (1850) ≡S.plumosavar.radicata(Hook.&Grev.) Warb.,Monsunia 1: 102 (1900).

Hypopterygiopsis raiateensis(J.W.Moore)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella raiateensisJ.W.Moore,Bull.Bernice P.Bishop Mus.102: 15 (1933).

Hypopterygiopsis rajasthanensis(Gena,Bhardwaja &A.K.Yadav)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella rajasthanensisGena,Bhardwaja &A.K.Yadav,Am.Fern J.69: 119(1979).

Hypopterygiopsis rasoloheryi(Rakotondr.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella rasoloheryiRakotondr.,Candollea 71(1): 144 (2016).

Hypopterygiopsis reineckei(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella reineckeiHieron.in Engl.&Prantl,Nat.Pfl.1(4): 678,no.83.(1901 publ.1902).

Hypopterygiopsis repanda(Desv.&Poir.) Spring Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Lycopodium repandumDesv.ex Poir.,Lam.,Encycl.suppl.3: 558.(1813 publ.1814) ≡Selaginella repanda(Desv.&Poir.) Spring,Gaudich.,Voy.Bonite Bot.1: 329(1846).高雄异穗卷柏(新拟)

Hypopterygiopsis reptansSakurai,Bot.Mag.(Tokyo) 57: 255(1943)Basionym:Selaginella sakuraiiH.Miller,Taxon 16:70(1967).

Hypopterygiopsis reticulata(Hook.&Grev.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium reticulatumHook.&Grev.,J.Bot.(Hook.)Kew Misc.2:402(1831)≡Selaginella reticulata(Hook.&Grev.)Spring,Bull.Acad.Roy.Soc.Bruxelles 10:233(1843).

Hypopterygiopsis robinsonii(Alderw.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella robinsoniiAlderw.,Phil.J.Sci.Bot.,11C: 118 (1916).

Hypopterygiopsis roesickeana(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella roesickeanaHieron.,Engl.Bot.Jahrb.56: 235,no.36 (1920).

Hypopterygiopsis sambiranensis(Stefanov.&Rakotondr.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella sambiranensisStefanov.&Rakotondr.,Adansonia ser.3,3:19(1):166,f.1-2 (1997).

Hypopterygiopsis scabrida(Ridl.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella scabridaRidl.,J.Roy.As.Soc.,Str.Br.,No.80: 159,no.32 (1919) ≡Selaginella alutaceavar.scabrida(Ridl.) Alston,Gard.Bull.Str.Settl.8: 56 (1934).

Hypopterygiopsis schefferi(Hieron.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella schefferiHieron.,Engl.Bot.Jahrb.50: 2,24,no.8 (1913).

Hypopterygiopsis selangorensis(Bedd.ex Ridl.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella selangorensisBedd.ex Ridl.,J.As.Soc.,Str.Br.,No.80: 148,no.3 (1919).

Hypopterygiopsis sespillifolia(Brownlie) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella sespillifoliaBrownlie,Fl.Nouv.-Caléd.3: 32 (1969).

Hypopterygiopsis setchellii(O.C.Schmidt)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella setchelliiO.C.Schmidt,Fedde,Repert,20: 158 (1924).

Hypopterygiopsis sichuanica(H.S.Kung) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella sichuanicaH.S.Kung,Acta Bot.Yunnan.3(2): 252 (1981).四川异穗卷柏(新拟)

Hypopterygiopsis singalanensis(Hieron.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella singalanensisHieron.,Hedwigia 50: 18,no.12 (1910).

Hypopterygiopsis societatis(J.W.Moore) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella societatisJ.W.Moore;Bull.Bernice P.Bishop Mus.102: 13 (1923).

Hypopterygiopsis spinulosovena(G.Q.Gou&P.S.Wang)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella spinulosovenaG.Q.Gou&P.S.Wang,Acta Bot.Yunnan.27(2):145-146(2005).刺脉异穗卷柏(新拟)

Hypopterygiopsis squarrosa(Baker)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella squarrosaBaker,J.Bot.23:180,no.276 (1885).

Hypopterygiopsis stolleana(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella stolleanaHieron.,Engl.Bot.Jahrb.56: 236,no.37 (1920).

Hypopterygiopsis strigosa(Bedd.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella strigosaBedd.,Kew Bull.,192,no.600 (1911).

Hypopterygiopsis subcordata(A.Braun ex Kuhn)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella subcordataA.Braun ex Kuhn;Fil.Afr.193 (1868) ≡Lycopodioides subcordata(A.Braun)Kuntze,Rev.Gen.Pl.1:827(1891)≡Stachygynandrum subcordatum(A.Braun) Carruth.,Cat.Afr.Pl.coll.by Dr.F.Welwitsch in 1853-1861,2(2): 262 (1901).

Hypopterygiopsis subeffusa(Shalimov et X.C.Zhang) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella subeffusaShalimov&X.C.Zhang,Turczaninowia 25(3):58(2022).近疏松异穗卷柏(新拟)

Hypopterygiopsis subdiaphana(Wall.ex Hook.&Grev.)Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium subdiaphanumWall.,(Cat.n.136 (1829),nom.nud.) ex Hook.&Grev.,Enum.Fil.J.Bot.(Hook.) Kew Misc.2: 401,no.183 (1831) ≡Selaginella subdiaphana(Wall.ex Hook.&Grev.) Spring,Bull.Acad.Roy.Sci.Bruxelles 10:232(1843)≡Lycopodioides subdiaphana(Wall.ex Hook.&Grev.) Kuntze,Rev.Gen.Pl.1: 825 (1891).近透明异穗卷柏(新拟)

Hypopterygiopsis subisophylla(Jermy) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella subisophyllaJermy,Brit.Fern Gaz.10: 30,f.1-9 (1968).

Hypopterygiopsis subchaetoloma(X.M.Zhou,Li Bing Zhang &Z.R.He) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella subchaetolomaX.M.Zhou,Li Bing Zhang &Z.R.He,Phytotaxa603(3): 225 (2023).近毛边异穗卷柏(新拟)

Hypopterygiopsis submonospora(Shalimov &X.C.Zhang) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella submonosporaShalimov &X.C.Zhang,Turczaninowia 25(1): 154(2022).拟单子异穗卷柏(新拟)

Hypopterygiopsis subspinulosa(Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella subspinulosaSpring,Miq.Pl.Jungh.3: 277,no.10 (1854).

Hypopterygiopsis subvaginata(X.C.Zhang&Shalimov)Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella subvaginataX.C.Zhang &Shalimov,J.Sp.Res.9(3): 222 (2020).拟鞘舌异穗卷柏(新拟)

Hypopterygiopsis tectissima(Baker)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella tectissimaBaker,J.Bot.22:89,no.119 (1884).

Hypopterygiopsis temehaniensis(Baker) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella temehaniensisJ.W.Moore;Bull.Bernice P.Bishop Mus.102: 13 (1933).

Hypopterygiopsis tenera(Hook.&Grev.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium tenerumHook.&Grev.,J.Bot.(Hook.) Kew Misc.2: 400 (1831) ≡Selaginella tenera(Hook.&Grev.) Spring,Bull.Acad.Roy.Soc.Bruxelles 10: 232 (1843) ≡Lycopodioides tenera(Hook.&Grev.)Kuntze,Rev.Gen.Pl.1:827(1891).

Hypopterygiopsis tenerrima(A.Braun ex Kuhn)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella tenerrimaA.Braun ex Kuhn,Fil.Afr.193 (1868) ≡Lycopodioides tenerrima(A.Braun)Kuntze,Rev.Gen.Pl.1:827(1891) ≡Stachygynandrum tenerrimum(A.Braun) Carruth.,Cat.Afr.Pl.coll.by Dr.F.Welwitsch in 1853-1861,2(2): 262 (1901).

Hypopterygiopsis tenuifolia(Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella tenuifoliaSpring,Mém.Acad.Roy.Sci.Belg.24(2): 253 (1850) ≡Lycopodioides tenuifolia(Spring) Kuntze,Rev.Gen.Pl.1: 827 (1891).

Hypopterygiopsis thomensis(Alston) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella thomensisAlston,Exell,Cat.Vasc.Pl.S.Tomé,97,f.3 (1944).

Hypopterygiopsis trichophylla(K.H.Shing) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella trichophyllaK.H.Shing,Vasc.Pl.Hengduan Mount.1:11(1993),without Latin descr.et Acta Phytotax.Sin.31(6): 569 (1993) ≡Selaginella monosporassp.trichophylla(K.H.Shing) X.C.Zhang,Fl.Reipubl.Popularis Sin.6(3):189 (2004).毛叶异穗卷柏(新拟)

Hypopterygiopsis unilateralis(Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella unilateralisSpring,Bull.Acad.Roy.Soc.Bruxelles 10: 232,no.140 (1843) ≡Lycopodioides unilateralis(Spring) Kuntze,Rev.Gen.Pl.1: 827(1891).

Hypopterygiopsis usta(Vieill.ex Baker) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella ustaVieill.ex Baker,Jour.Bot.23: 23,no.212.1885.

Hypopterygiopsis vanderystii(Bizzarri) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella vanderystiiBizzarri,Bull.Jard.Bot.Belg.53(1-2): 171 (1983).

Hypopterygiopsis vaginata(Spring)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella vaginataSpring,Mém.Acad.Roy.Sci.Belg.24: 87 (1850) ≡Lycopodioides vaginata(Spring) Kuntze,Rev.Gen.Pl.1: 827 (1891).鞘舌异穗卷柏(新拟)

Hypopterygiopsis vieillardii(Warb.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella vieillardiiWarb.,Monsunia 1:125,no.138 (1900).

Hypopterygiopsis wangpeishanii(Li Bing Zhang,H.He &Q.W.Sun) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella wangpeishaniiLi Bing Zhang,H.He &Q.W.Sun,Phytotaxa 164(3):195 (2014).培善异穗卷柏(新拟)

Hypopterygiopsis wattii(Baker) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella wattiiBaker,Handb.Fern-Allies 109 (1887) ≡Lycopodioides wattii(Baker) Kuntze,Rev.Gen.Pl.1:827 (1891).

Hypopterygiopsis weinlandii(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella weinlandiiHieron.,Engl.Bot.Jahrb.50: 21,29,no.11 (1913).

Hypopterygiopsis whitmeei(Baker)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella whitmeeiBaker,J.Bot.23:24,no.215(1885)≡Lycopodioides whitmeei(Baker)Kuntze,Rev.Gen.Pl.1:827 (1891).

Hypopterygiopsis wuyishanensis(K.W.Xu,X.M.Zhou &Y.F.Duan)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella wuyishanensisK.W.Xu,X.M.Zhou&Y.F.Duan,PhytoKeys 202:111(2022).武夷山异穗卷柏(新拟)

Hypopterygiopsis xichouensis(W.M.Chu)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella xichouensisW.M.Chu,Fl.Yunnan.20: 720 (2006).西畴异穗卷柏(新拟)

Hypopterygiopsis xipholepis(Baker)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella xipholepisBaker,J.Bot.23:155,no.258 (1885) ≡Lycopodioides xipholepis(Baker) Kuntze,Rev.Gen.Pl.1: 827 (1891).剑叶异穗卷柏(新拟)

Hypopterygiopsis xishuiensis(G.Q.Gou &P.S.Wang) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella xishuiensisG.Q.Gou&P.S.Wang,Acta Phytotax.Sin.43(1):71(2005).习水异穗卷柏(新拟)

Hypopterygiopsis yunckeri(Alston)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella yunckeriAlston,Bull.Bernice P.Bishop Mus.220: 44 (1959).

Hypopterygiopsis zechii(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella zechiiHieron.,Engl.&Prantl,Nat.Pfl.1(4): 697,no.298 (1901 publ.1902).

Hypopterygiopsis zhangii(S.Y.Dong) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella zhangiiS.Y.Dong,Syst.Bot.47(1): 91 (2022).宪春异穗卷柏(新拟)

Hypopterygiopsis zollingeriana(Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella zollingerianaSpring,Miq.,Pl.Jungh.3: 278,no.11 (1854).

5.7.3.Lycopodioides

LycopodioidesBoehm.,Def.Gen.Pl.(ed.3).485.1760.疏穗卷柏属(新拟) -Isonym:LycopodioidesKuntze,Revis.Gen.Pl.2: 824(1891) ≡TrispermiumHill,Gener.Nat.Hist.,ed.2,2(Hist.Pl.): 112(1773)-Type:Lycopodium denticulatumL.,Sp.Pl.2:1106(1753)≡Selaginella denticulata(L.) Spring,Flora 21(10): 149 (1838) ≡Lycopodioides denticulata(L.) Kuntze,Revis.Gen.Pl.1-2: 824(1891).

=DiplostachyumP.Beauv.,Mag.Encycl.5: 481.(1804) (‘Diplostachium’).Lycopodioidessect.Diplostachyum(P.Beauv.) Rothm.Lycopodiumsect.Diplostachyum(P.Beauv.)Rchb.≡Selaginellasect.Diplostachyum (P.Beauv.) T.Moore ≡HeterophylliumHieron.ex Börner.Lectotype (designated by Pfeiffer,Nom.1: 1100.1874):Lycopodium helveticumL.,Sp.Pl.2: 1104 (1753) ≡Bernhardia helvetica(L.)Gray,Nat.Arr.Brit.Pl.2:23(1821)≡Selaginella helvetica(L.) Spring,Flora 21(1): 149 (1838) ≡Lycopodioideshelvetica (L.)Kuntze,Revis.Gen.Pl.2: 826 (1891) ≡Heterophyllium helveticum(L.) Börner,Fl.Deut.Volk 285 (1912).

Lycopodioidesis the most frequently recognized genus in addition toSelaginellain the family,but our circumscription is different from those in earlier literature and corresponds toSelaginellasect.Homostachyssensu Zhou and Zhang (2015).The latter has been recognized in earlier classifications ofSelaginella(e.g.,Walton and Alston,1938;Warburg,1900).Walton and Alston (1938) includedSelaginella rothertii,a species from high mountain in Java (Alston,1935)as a member ofS.subg.Homostachys.S.rothertiimight be the southernmost species in subg.Homostachys,whereas the remaining species are distributed from East Asia to Europe (Zhang et al.,2021).Warburg (1900) recognizedS.subg.Homostachys,but includedS.ciliarisandS.pallidissimain it.AlthoughS.ciliarishas creeping plant,similar to those species in subg.Homostachys,this species has dimorphic but resupinate strobili and reticulate megaspore and is a member ofHypopterygiopsissect.Hypopterygiopsis.

This genus is characterized by creeping stems (Fig.8G),dimorphic sporophylls (Fig.8F and G) (except forSelaginella denticulataSpring andS.helvetica(L.) Spring with nearly monomorphic but loose sporophylls (Fig.9E)),loose and non-resupinate (with the smaller ones in the same plane as the median leaves and the larger ones in the same plane as the ventral leaves)strobili(Fig.8F and G).Species inLycopodioidesusually grow in wet habitat.Although megaspore surfaces with papillate,tuberculate,and verrucate ornamentation are similar to those ofHypopterygiopsissect.Hypopterygiopsis,but the laesurae of megaspores inLycopodioidesare usually disconnected at the pole (Zhou et al.,2015;Zhou and Zhang,2015).2n=18 (Reeve,1935;Jermy et al.,1967;Loyal et al.,1984;Takamiya,1993).

Lycopodioidesknown so far (Zhou et al.,2022) has large plastome size(ca.187 kb)with the most genes(128)in Selaginellaceae.Lycopodioideshas three ribosomal operon copies,and its master conformation of plastomes can mediate eight isomers(Zhou et al.,2022).

This genus contains about 12 species mainly occurring in Asia and Europe(Zhang et al.,2021).

Members:

Lycopodioides denticulata(L.) Kuntze in Revis.Gen.Pl.2: 824.1891.Basionym:L.denticulatumL.,Sp.Pl.2: 1106 (1753).

Lycopodioides helvetica(L.)Kuntze,Revis.Gen.Pl.2:826.1891.Basionym:L.helveticumL.,Sp.Pl.2: 1104-1105 (1753).小疏穗卷柏(新拟)

Lycopodioides jiulongensisH.S.Kung,Li Bing Zhang &X.S.Guo in Acta Bot.Yunnan.17(4): 420,fig.1 (1995).九龙疏穗卷柏(新拟)

Lycopodioides laxistrobila(K.H.Shing) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella laxistrobilaK.H.Shing,Acta Phytotax.Sin.31(6): 569 (1993).疏穗卷柏(新拟)

Lycopodioides longistrobilina(P.S.Wang,X.Y.Wang &Li Bing Zhang)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella longistrobilinaP.S.Wang,X.Y.Wang&Li Bing Zhang,Novon 22:260.2012.长穗疏穗卷柏(新拟)

Lycopodioides nipponica(Franch.&Sav.)Kuntze,Revis.Gen.Pl.2:827.1891(‘nipponicum’).Basionym:Selaginella nipponicaFranch.&Sav.,Enum.Pl.Jap.2(2): 199,615 (1879).伏地疏穗卷柏(新拟)

Lycopodioides pallidissima(Spring) Kuntze;Revis.Gen.Pl.2:827 (1891).Selaginella pallidissimaSpring in Bull.Acad.Roy.Sci.Bruxelles 10: 231 (1843).平疏穗卷柏(新拟)

Lycopodioides prostrataH.S.Kung,Fl.Sichuan.6:76(1988).地疏穗卷柏(新拟)

Lycopodioides pseudonipponica(Tagawa) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella pseudonipponicaTagawa,Acta Phytotax.Geobot.25: 177 (1973) ≡Selaginella helveticasubsp.pseudonipponica(Tagawa)H.M.Chang,W.L.Chiou&J.C.Wang,Fl.Taiwan,Selaginellaceae 38 (2012).拟伏地疏穗卷柏(新拟)

Lycopodioides rothertii(Alderw.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella rothertiiAlderw.,Bull.Jard.Bot.Buitenz.II,7: 30 (1912).

Lycopodioides shensiensis(Christ) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella shensiensisChrist;Nuov.Giorn.Bot.Ital.n.s.4: 102 (1897).陕西疏穗卷柏(新拟)

Lycopodioides tama-montana(Seriz.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella tama-montanaSeriz.,J.Jpn.Bot.53(8): 242(1978).高山疏穗卷柏(新拟)

5.7.4.Valdespinoa

ValdespinoaLi Bing Zhang&X.M.Zhou,gen.nov.韦氏卷柏属(新拟) -Type:Valdespinoa douglasii(Hook.&Grev.) Li Bing Zhang &X.M.Zhou (≡Selaginella douglasii(Hook.&Grev.) Spring).

Etymology:In honor of Prof.Iván A.Valdespino currently based at PMA for his contributions to the study ofSelaginella(e.g.,Valdespino,1993a,2015,2016,2017b,2020;Valdespino et al.,2015,2018a,b).

Valdespinoacorresponds toSelaginellasect.Auriculatesensu Zhou and Zhang (2015).

Valdespinoais sister toLycopodioideswith strong support.However,Valdespinoahas monomorphic sporophylls and is only distributed in America.We recognize the monospecificValdespinoacontainingV.douglasiionly.Hieronymous and Sadebeck (1902)recognized theDouglasiigroup which containsS.accharata,S.delicatissma,S.douglasii,andS.reflexa.Phylogenetic analyses showed thatS.reflexais member of ourSelaginellas.s(Weststrand and Korall,2016a;Zhou et al.,2016).Based on the morphology,S.delicatissma(includingS.accharata) might be members ofValdespinoa,but more studies are necessary.

Valdespinoahas the largest plastome size (>187 kb) known in Selaginellaceae.Its plastome has three ribosomal operon copies and its master conformation can mediate 24 isomers (Zhou et al.,2022).

Valdespinoacurrently containsV.douglasiiin USA only(Valdespino,1993a).

Member:

Valdespinoa douglasii(Hook.&Grev.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium douglasiiHook.&Grev.Bot.Misc.2: 396 (1831) ≡Selaginella douglasii(Hook.&Grev.)Spring,Bull.Acad.Roy.Soc.Bruxelles 10: 138 (1843).

5.8.Subfamily VII.Selaginelloideae

Subfamily VII.SelaginelloideaeLi Bing Zhang&X.M.Zhou,subfam.nov.同穗卷柏亚科(新拟) -Type:SelaginellaP.Beauv.

Plants erect,suberect,ascending,or creeping,rarely rosetteforming,with ventral rhizophores borne on ventral side of stems and/or branches;sterile leaves dimorphic,decussate;sporophylls monomorphic or dimorphic (resupinate);megaspore surfaces reticulate(Stachygynandrumtype with low and open muri;Fig.2Z),rarely verrucate;microspore surfaces often baculate,sometimes verrucate to blunt echinate;2n=18,20,22,36,36-40,or 50-60(Jermy et al.,1967;Love and Love,1976;Takamiya,1993;Marcon et al.,2005).

This subfamily corresponds toSelaginellasubg.Stachygynandrumsensu Zhou and Zhang (2015).

Morphologically,Selaginelloideae are characterized by creeping,ascending or erect(never scandent)plants(Figs.10A-F and 11A-D),monomorphic sporophylls(Figs.10G,H and 11B,E,F)(but including all these species with dimorphic or subdimorphic sporophylls in new world) (Fig.11G and H).Megaspores usually have more or less a slightly developed zona at the end of laesurae (Figs.10K-N) or obviously equatorial flange (e.g.,Kungiselaginella involvens: Fig.11I)and surfaces with reticulate ornamentation usually with often open mesh(Figs.10I,K-N and 11J,M-P).Microspore surfaces are usually baculate(Fig.10J,O-R),blunt spiny(Fig.10 Q-R),papilary(K-L),or verrucate (Fig.10S and T) (Valdespino,2015,2016,2017b,c,2020;Valdespino et al.,2015,2018b;Zhou et al.,2015).

Following our current classification,nearly all New World species with ventral rhizophores,exceptDidiclis hoffmannii,Pulviniella,andValdespinoa douglasii,should be members of this subfamily.Selaginelloideae are mainly pantropical distribution,and some species can extend to subtropical and temperate Asia and Americas(Valdespino,1993a;Zhang et al.,2013).

In our previous analysis (Zhou et al.,2015,2016) and classification (Zhou and Zhang,2015),megaspore data have been well explored and used and their taxonomic significance was intensively discussed (Korall and Taylor,2006;Zhou and Zhang,2015;Zhou et al.,2015,2016;Weststrand and Korall,2016a).In contrast,the microspore morphology of Selaginelloideae was rarely observed and reported except that of the Old World(especially Asian)species(e.g.,Zhou et al.,2015).In recent years,as a large number of new species from the New World have been described and microspore data have been provided by Valdespino and co-authors (e.g.,Valdespino,2015,2016,2017b,c;2020;Valdespino et al.,2015,2018b),we now know that New-World species have more diverse microspore surfaces than those of the Old-World species (Zhou et al.,2015;unpublished data).

Jermy (1986) definedSelaginellasubg.Stachygynandrumas containing only those species with monomorphic sporophylls,but phylogenetic analyses showed that some species with dimorphic strobili in the New World are also mixed with species with monomorphic sporophylls in the tree and the dimorphism of sporophylls have evolved 10 times (Zhou et al.,2016;Weststrand and Korall,2006a;our Fig.2G).Based on previous studies and our observations,species with dimorphic sporophylls from the New World also have same reticulate ornamentation as those species with monomorphic sporophylls inS.subg.Stachygynandrumsensu Jermy (1986) [e.g.,Selaginella hyalogrammaValdespino (Valdespino,2017a)S.mucronata(Valdespino et al.,2015);S.pellucidopunctataValdespino(Valdespino et al.,2015);and our observations forS.flagellataandS.simplex].These are entirely different from species with dimorphic sporophylls inLycopodioidesandDidicliswhich do not occur in the New World exceptD.hoffmannii.The dimorphism of sporophylls in those species from the Old World and that in those from the New World independently evolved(Fig.2G).Evidence so far showed that the New-World species with dimorphic sporophylls are typically reticulate on megaspores and baculate on microspores.It is clear that reticulate megaspores and baculate or verrucate microspores are diagnosing features of Selaginelloideae newly defined here.

In addition,tuberculate or verrucate microspores were also observed in some New-World species in the subfamily [(e.g.,Selaginella pubimarginataValdespino (2020);S.brigitteanaValdespino (Valdespino,2019);S.neospringianaValdespino (2015c);S.mucugensisValdespino;S.saltuicolaValdespino;S.sematophyllaValdespino (Valdespino et al.,2015)].Also,some morphological characters,e.g.,dimorphic or monomorphic sporophylls,reticulate or non-reticulate megaspores,baculate or non-baculate microspores,etc.,reversed and/or independently evolved many times in the New-World Selaginelloideae (Fig.2J).

The plastomes of Selaginelloideae known so far (Zhou et al.,2022) are extremely conserved and have the same gene order.Species of Selaginelloideae have no more than two pairs of small repeats in SC and their plastomes have up to five conformations.Those ofSelaginella moellendorffiiand its close relatives have IR and a pair of repeats existed in SC with four conformations,whereas those of the remaining species in the subfamily have DR or DR-IR coexisting.

This subfamily contains about 330 species in three genera,Chuselaginella,Kungiselaginella,andSelaginellas.s.,the former two occurring in the Old World only,and the latter in Americas only except one (a few) species in Africa.ChuselaginellaandKungiselaginellaare morphologically distinguishable from each other (see below and our key)and have slightly different distributions:Asia to Pacific islands and Africa forChuselaginella,and Asia forKungiselaginella.K.moellendorffii(=S.moellendorffii) in America is introduced from the Old World.Selaginella s.s.is morphologically nearly indistinguishable from the other two.Alternatively,these three genera could be combined into a broadly definedSelaginella,but we prefer to separate them because of their molecular divergence(especially in nuclear phylogeny;Fig.S2),geographical coherence,and more manageable size.

5.8.1.Chuselaginella

ChuselaginellaLi Bing Zhang &X.M.Zhou,gen.nov.朱氏卷柏属(新拟)-Type:Chuselaginella alopecuroides(Baker)Li Bing Zhang&X.M.Zhou (=Selaginella alopecuroidesBaker,J.Bot.British &Foreign 19: 368,no.83.(1881)).

≡Selaginellasect.Ascendentes(Baker) Li Bing Zhang &X.M.Zhou,Taxon 64(6): 1136.2015 ≡S.ser.AscendentesBaker,J.Bot.Lond.21: 3 (1883) -Lectotype (designated by Zhou and Zhang,2015: 1136):S.alopecuroidesBaker.

≡S.ser.RadicantesWarb.,Monsunia 1: 102 (1900) -Lectotype(designated by Zhou and Zhang,2015:1136):S.alopecuroidesBaker.

Etymology:-FromChu-,in honor of the late Prof.Wei-Ming Chu of Yunnan University(herbarium PYU),China,for his contributions to the study of ferns in general and those ofSelaginellaof Yunnan in particular (Chu et al.,2006).

Chuselaginellacorresponds toSelaginellasect.Ascendentessensu Zhou and Zhang(2015).This genus is supported as monophyletic in all of our analyses (Figs.1 and S1-S3).Species of this genus have plants often ascending (a few strictly erect),microspores always baculate with expanded tips and megaspores with reticulum and a unique zonal structure at the ends of laesurae(Fig.10K-N)(e.g.,Liu et al.,1989 forSelaginella doederleiniiHieron.;Liu et al.,2001 forS.frondosaWarb.;Zhao et al.,2006 forS.commutataAlderw.;Zhou et al.,2015,forS.trichophyllaK.H.Shing andS.scabrifoliaChing &Chu H.Wang;and Korall and Taylor,2006 for additional species).

Chuselaginellacontains more than 70 species from Africa,Asia,Australia,and South Pacific islands.

Members:

Chuselaginella aenea(Warb.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella aenea Warb.,Monsunia 1: 104,115,no.58 (1900).

Chuselaginella agusanensis(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella agusanensisHieron.,Leafl.Philipp.Bot.6: 1998.

Chuselaginella alligans(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella alligansHieron.,Elmer,Leafl.Phil.Bot.6: 1998,2012,no.15 (1913).

Chuselaginella alopecuroides(Baker) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella alopecuroidesBaker,J.Bot.19: 368,no.83 (1881).

Chuselaginella auriculata(Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella auriculataSpring,Bull.Acad.Roy.Soc.Bruxelles 10: 142,no.147 (1843).

Chuselaginella bluuensis(Alderw.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella bluuensisAlderw.,Bull.Jard.Bot.Buitenz.II,11: 29 (1913).

Chuselaginella borneensis(Hieron.)Li Bing Zhang&X.M.Zhou,comb.&stat.nov.Basionym:Selaginella frondosavar.borneensisHieron.,Bot.Jahrb.Syst.44: 511 (1910).

Chuselaginella brachyblepharis(Alderw.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella brachyblepharisAlderw.,Bull.Jard.Bot.Buitenzorg,sér.2,11: 24(1913)

Chuselaginella brevipes(A.Braun) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella brevipesA.Braun,App.Ind.Sem.Hort.Berol.1 (1867)

Chuselaginella breynioides(Baker) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella breynioidesBaker,J.Bot.23: 45,no.222 (1885).

Chuselaginella brooksii(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella brooksiiHieron.,Hedwigia 51:252,no.6 (1912).

Chuselaginella burkei(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella burkeiHieron.,Engl.Bot.Jahrb.50:2,16,no.5 (1913).

Chuselaginella carnea(Alderw.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella carneaAlderw.,Bull.Jard.Bot.Buitenzorg,sér.2,24: 7 (1917).

Chuselaginella cesatii(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella cesatiiHieron.,Hedwigia 50:6,no.5 (1910).

Chuselaginella ciliata(Alderw.) Li Bing Zhang &X.M.Zhou,comb.&stat.nov.Basionym:Selaginella frondosavar.ciliataAlderw.,Bull.Jard.Bot.Buitenzorg,sér.2,16: 48 (1914).

Chuselaginella commutata(Alderw.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella commutataAlderw.,Bull.Jard.Bot.Buitenz.II,28: 46 (1918).长芒朱氏卷柏(新拟)

Chuselaginella copelandii(Hieron.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella copelandiiHieron.Repert.Spec.Nov.Regni Veg.10: 106 (1911).

Chuselaginella cumingiana(Spring)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella cumingianaSpring Bull.Acad.Roy.Soc.Bruxelles 10: 146,no.81 (1843).

Chuselaginella cuprea(Ridl.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella cupreaRidl.;J.Straits Branch Roy.Asiat.Soc.80: 152,no.14 (1919)

Chuselaginella cupressina(Willd.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium cupressinumWilld.,Sp.Pl.5: 43(1810).

Chuselaginella dielsii(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella dielsiiHieron.;Hedwigia 51:254,no.7 (1911 publ.1912).

Chuselaginella distans(Warb.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella distansWarb.,Monsunia 1: 106,120,no.86 (1900).

Chuselaginella doederleinii(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella doederleiniiHieron.,Hedwigia 43(1): 41-42 (1904).深绿朱氏卷柏(新拟)

Chuselaginella dolichocentrus(K.M.Wong) Li Bing Zhang &X.M.Zhou,comb.&stat.nov.Basionym:Selaginella intermediavar.dolichocentrusK.M.Wong,Gard.Bull.Singapore 35(2): 125 (1982 publ.1983)

Chuselaginella elmeri(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella elmeriHieron.,Repert.Spec.Nov.Regni Veg.10: 46,no.5 (1911).

Chuselaginella fenixii(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella fenixiiHieron.Repert.Spec.Nov.Regni Veg.10: 98,no.10 (1911).

Chuselaginella frondosa(Warb.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella frondosaWarb.Monsunia 1: 105,117,no.75 (1900).繁叶朱氏卷柏(新拟)

Chuselaginella grabowskyi(Warb.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella grabowskyiWarb.,Monsunia 1:107,122,no.109a (1900).

Chuselaginella grandis(T.Moore) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella grandisT.Moore,Gard.Chron.18(2): 40,t.8 (1882).

Chuselaginella griffithii(Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella griffithiiSpring Bull.Acad.Roy.Soc.Bruxelles 10: 145,no.80 (1843).

Chuselaginella guihaia(X.C.Zhang)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella guihaiaX.C.Zhang PhytoKeys 80:44 (2017).桂海朱氏卷柏(新拟)

Chuselaginella hordeiformis(Baker)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella hordeiformisBaker J.Bot.23: 47,no.229 (1885).

Chuselaginella hosei(Hieron.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella hoseiHieron.;Hedwigia 51: 243,no.3.(1911 publ.1912).

Chuselaginella intermedia(Blume)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Lycopodium intermediumBlume,Enum.Pl.Javae 2:269,no.20(1830).≡Selaginella intermedia(Blume)Spring Bull.Acad.Roy.Soc.Bruxelles 10: 144 (1843).

Chuselaginella jagorii(Warb.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella jagoriiWarb.Monsunia 1: 104,116,no.68 (1900).

Chuselaginella kerstingii(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella kerstingiiHieron.Engl.Bot.Jahrb.50: 2,21,no.7 (1913).

Chuselaginella kusaiensis(Hosok.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella kusaiensisHosok.Trans.Nat.Hist.Soc.Formosa 25 (147): 440 (1935).

Chuselaginella latifrons(Warb.non Hort.ex Williams) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella latifronsWarb.non Hort.ex Williams Monsunia 1: 106,120,no.89 (1900).

Chuselaginella lebongtandaiana(Alderw.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella lebongtandaianaAlderw.Bull.Jard.Bot.Buitenz.II,23: 23 (1916).

Chuselaginella leveriana(Alston) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella leverianaAlston J.Bot.77: 225(1939).

Chuselaginella longiaristata(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella longiaristataHieron.Hedwigia 50: 16,no.11 (1910).

Chuselaginella longipinna(Warb.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella longipinnaWarb.Monsunia 1:105,119 (1900).

Chuselaginella longirostris(Alderw.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella longirostrisAlderw.Bull.Jard.Bot.Buitenz.II,11: 25 (1913).

Chuselaginella lonko-batu(Hieron.&Alderw.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella lonko-batuHieron.&Alderw.Bull.Jard.Bot.Buitenz.II,16: 42 (1914).

Chuselaginella louisiadensis(Hieron.) Li Bing Zhang &X.M.Zhou,comb.&stat.nov.Basionym:Selaginella burkeivar.louisiadensisHieron.,Bot.Jahrb.Syst.50: 2,18 (1913).

Chuselaginella luzonensis(Hieron.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella luzonensisHieron.Engl.&Prantl,Nat.Pfl.1(4): 681,no.113.(1901 publ.1902).

Chuselaginella magnifica(Warb.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella magnificaWarb.Monsunia 1:103,114,no.45,t.3A (1900).

Chuselaginella melanesica(Kuhn) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella melanesicaKuhn,Forschungsr.Gazelle 4,Bot: 17 (1889).

Chuselaginella moszkowskii(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella moszkowskiiHieron.Bot.Jahrb.Syst.50: 2,14,no.4 (1913).

Chuselaginella opaca(Seriz.)Li Bing Zhang&X.M.Zhou,comb.&stat.nov.Basionym:Selaginella doederleiniivar.opacaSeriz.,J.Phytogeogr.Taxon.30(1): 44 (1982).

Chuselaginella parvifolia(Alderw.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella parvifoliaAlderw.;Bull.Jard.Bot.Buitenzorg,sér.2,11: 30 (1913).

Chuselaginella paxii(Hieron.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella paxiiHieron.;Bot.Jahrb.Syst.44: 512(1910).

Chuselaginella pentagona(Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella pentagonaSpring Mém.Acad.Roy.Sci.Belg.24(2): 150 (1850).

Chuselaginella petelotii(Alston) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella petelotiiAlston H.Lecomte,Fl.Indo-Chine 7: 562,t.66,f.1-5 (1951).

Chuselaginella poperangensis(Hieron.ex Rech.) Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella poperangensisHieron.ex Rech.Denkschr.Kaiserl.Akad.Wiss.,Wien.Math.-Naturwiss.Kl.89: 483,t.7,f.13C (1914).

Chuselaginella posewitzii(Hieron.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella posewitziiHieron.Hedwigia 51:241,no.1.(1911 publ.1912).

Chuselaginella procera(Alston) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella proceraAlston J.Bot.72: 227(1934).

Chuselaginella puberulipes(Alderw.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella puberulipesAlderw.,Nova Guinea,Rés.d.Expéd.á Nouv.Guin.14 (Bot.): 66 (1924).

Chuselaginella quadrivenulosa(Alderw.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella quadrivenulosaAlderw.Nova Guinea,Rés.d.Expéd.á Nouv.Guin.14 (Bot.): 65(1924).

Chuselaginella ramosii(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella ramosiiHieron.Repert.Spec.Nov.Regni Veg.10: 52,no.8 (1911).

Chuselaginella rivalis(Ridl.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella rivalisRidl.Bull.Misc.Inform.Kew 1924:266 (1924).

Chuselaginella rolandi-principis(Alston) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella rolandi-principisAlston J.Bot.72: 228-229 (1934).海南朱氏卷柏(新拟)

Chuselaginella roxburghii((Hook.&Grev.) Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella roxburghii(Hook.&Grev.) Spring Bull.Acad.Roy.Soc.Bruxelles 10: 228,no.115 (1843).

Chuselaginella rugulosa(Ces.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella rugulosaCes.Felci Borneo,Atti della R.Acad.Sc.7 (8): 35(1876).

Chuselaginella sambasensis(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella sambasensisHieron.Hedwigia 50: 9,no.7 (1910).

Chuselaginella sarawakensis(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella sarawakensisHieron.Hedwigia 50: 13,no.10 (1910).

Chuselaginella scabrifolia(Ching&Chu H.Wang)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella scabrifoliaChing &Chu H.Wang Acta Phytotax.Sin.8(2): 157(1959).糙叶朱氏卷柏(新拟)

Chuselaginella schatteburgiana(Hieron.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella schatteburgianaHieron.Engl.Bot.Jahrb.56: 229,no.29 (1920).

Chuselaginella sepikensis(Hieron.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella sepikensisHieron.Bot.Jahrb.Syst.56: 243,no.58 (1920).

Chuselaginella spanielema(Alston)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella spanielemaAlston,J.Bot.72: 229(1934) (spanioclema,sphalm.)

Chuselaginella strigosa(Ridl.)Li Bing Zhang&X.M.Zhou,comb.&stat.nov.Basionym:Selaginella trichobasisvar.strigosaRidl.,J.Straits Branch Roy.Asiat.Soc.80: 153 (1919).≡Selaginella roxburghiivar.strigosa(Ridl.) K.M.Wong,Gard.Bull.Singapore 35(2):corrigendum slip [131 (1982 publ.1983).

Chuselaginella subalpina(Alderw.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella subalpinaAlderw.Bull.Jard.Bot.Buitenz.II,20: 26 (1915).

Chuselaginella subcalcarata(Alderw.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella subcalcarataAlderw.Bull.Jard.Bot.Buitenz.II,16: 49 (1914).

Chuselaginella subpedalis(Alderw.)Li Bing Zhang&X.M.Zhou,comb.&stat.nov.Basionym:Selaginella paxiivar.subpedalisAlderw.,Bull.Jard.Bot.Buitenzorg,sér.2,11: 30 (1913).

Chuselaginella subserpentina(Alderw.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella subserpentinaAlderw.Bull.Jard.Bot.Buitenz.II,1: 17 (1911).

Chuselaginella suffruticosa(Alderw.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella suffruticosaAlderw.;Bull.Jard.Bot.Buitenzorg,sér.2,1: 22 (1911).

Chuselaginella superba(Alston) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella superbaAlston,J.Bot.70:63-64,no.4,t.600,f.A-F (1932).粗茎朱氏卷柏(新拟)

Chuselaginella thurnwaldiana(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella thurnwaldianaHieron.Engl.Bot.Jahrb.56: 227,no.28 (1920).

Chuselaginella trachyphylla(A.Braun) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella trachyphyllaA.Braun Sitz.Ges.Naturf.Freunde Berl.Berl.8 (1863).粗叶朱氏卷柏(新拟)

Chuselaginella versicolor(Spring) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella versicolorSpring Bull.Acad.Roy.Soc.Bruxelles 10: 143,no.57 (1843).

Chuselaginella vestita(Alderw.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella vestitaAlderw.Bull.Jard.Bot.Buitenz.II,28: 54 (1918).

Chuselaginella vonroemeri(Alderw.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella vonroemeriAlderw.Bull.Jard.Bot.Buitenz.II,24: 7 (1917).

Chuselaginella wariensis(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella wariensisHieron.Engl.Bot.Jahrb.50: 2,19,no.6 (1913).

Chuselaginella zahnii(Hieron.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella zahniiHieron.Engl.Bot.Jahrb.50:2,37,no.15 (1913).

5.8.2.Kungiselaginella

KungiselaginellaLi Bing Zhang&X.M.Zhou,gen.nov.孔氏卷柏属(新拟) -Type:Kungiselaginella involvens(Sw.) Li Bing Zhang &X.M.Zhou(=Lycopodium involvensSw.,Syn.Fil.182,no.50(1806)).

≡Selaginellasect.Circinatae(Hook.&Grev.) Li Bing Zhang &X.M.Zhou,Taxon 64(6): 1136.2015 ≡Lycopodium(unranked)CircinataeHook.&Grev.,Bot.Misc.2: 380.1831 ["Circinata"] ≡S.(unranked)Circinatae(Hook.&Grev.)Spring in Mart.,Fl.Bras.1(2):118 (1840) -Lectotype (designated by Zhou and Zhang,2015:1136):S.involvens(Sw.) Spring.

≡S.(unranked)CaulescentesA.Braun,App.Ind.Sem.Hort.Berol.11.[1857]1858 ≡S.ser.Caulescentes(A.Braun)Baker,J.Bot.Lond.21:3(1883)-Type:S.caulescensSpring[=S.involvens(Sw.)Spring].

≡S.(unranked)RosulataeA.Braun,App.Ind.Sem.Hort.Berol.11([1857]1858),nom.illeg.≡S.ser.Rosulatae(A.Braun)Baker,J.Bot.Lond.2: 21: 3 (1883) ≡Lycopodioidessect.Rosulatae(A.Braun)Tzvel.,Novosti Sist.Vyssh.Rast.36: 25 (2004) -Type:S.involvens(Sw.) Spring.

=S.sect.Plagiophyllae(Warb.)Li Bing Zhang&X.M.Zhou,Taxon 64(6):1137(2015)≡S.subser.PlagiophyllaeWarb.,Monsunia 1:103(1900) -Lectotype (designated by Zhou and Zhang,2015: 1137):Selaginella biformisA.Braun ex Kuhn.

Etymology:-FromKung-,in honor of the late Prof.Hsian-Shiu Kung of Chengdu Institute of Biology,Chinese Academy of Sciences (herbarium CDBI),for his contributions to the study of ferns in general and those of Sichuan in particular (e.g.,Kung,1988).He was one of the early authors who recognized more than one genus in Selaginellaceae.

This genus circumscribed here corresponds toSelaginellasect.CircinataeandS.sect.Plagiophyllaesensu Zhou and Zhang (2015)combined.The genus is strongly supported as monophyletic in two of the three analyses based on concatenated data and strongly supported in nuclear tree(Fig.1 and S2).

Plants ofKungiselaginellaare strictly erect (a few creeping).Ventral leaves of members of this genus usually have two lightcolored bands on the sides of veins (Chu,2006).Their microspores are either slightly baculate(e.g.,K.davidii(Franch.)Li Bing Zhang &X.M.Zhou,andK.moellendorffii(Hieron.) Li Bing Zhang&X.M.Zhou) or tuberculate to blunt-spiny (e.g.,K.involvens)(Zhou et al.,2015).Two forms of habit and usually sterile microspores are present inS.biformis(Zhou et al.,2015).

This genus contains more than two dozen species in Asia.

Members:

Kungiselaginella argentea(Wall.ex Hook.&Grev.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium argenteumWall.ex Hook.&Grev.,Bot.Misc.2: 384 (1831).

Kungiselaginella biformis(A.Braun ex Kuhn) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella biformisA.Braun ex Kuhn Fil.Afr.189(1868),nomen,ex Baker,J.Bot.21:145(1883).二形孔氏卷柏(新拟)

Kungiselaginella davidii(Franch.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella davidiiFranch.,Pl.David.1: 344(1884).蔓生孔氏卷柏(新拟)

Kungiselaginella hellwigii(Hieron.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella hellwigiiHieron.,Bot.Jahrb.Syst.50: 2,12,no.3 (1913).

Kungiselaginella hieronymiana(Alderw.)Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella hieronymianaAlderw.Bull.Jard.Bot.Buitenz.II,7: 31 (1912).

Kungiselaginella involvens(Sw.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium involvensSw.,Syn.Fil.182,no.50(1806).兖州孔氏卷柏(新拟)

Kungiselaginella moellendorffii(Hieron.) Li Bing Zhang&X.M.Zhou,comb.nov.Basionym:Selaginella moellendorffiiHieron.in Engl.&Prantl,Nat.Pfl.1(4):680,no.102(1901 publ.1902).江南孔氏卷柏(新拟)

Kungiselaginella muelleri(Baker) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella muelleriBaker,J.Bot.23:122,no.123 (1885).

Kungiselaginella pallida(Hook.&Grev.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Lycopodium pallidumHook.&Grev.;Enum.Filic.J.Bot.(Hook.) Kew Misc.2: 389,no.130 (1831) (non Beyr.) 灰白孔氏卷柏(新拟)

Kungiselaginella peltata(C.Presl) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella peltataC.Presl Abh.Königl.Böhm.Ges.Wiss.,ser.5.

Kungiselaginella polyura(Warb.) Li Bing Zhang &X.M.Zhou,comb.nov.Basionym:Selaginella polyuraWarb.,Monsunia 1: 104,116,no.64,t.VI,f.A.(1900).

5.8.3.Selaginella

SelaginellaP.Beauv.,Mag.Encycl.9(5): 478 (1804),nom.cons.同穗卷柏属(新拟) -Type:S. flabellata(L.) Spring.conserved type.prop(Wan et al.,2023).

=SelagoP.Browne,Civ.Nat.Hist.Jamaica 82.1756-Lectotype(designated by Pichi Sermolli,Webbia 26:158(1971)):Lycopodium serpensDesv.ex Poir.non L.(1753).

≡StachygynandrumP.Beauv.ex Mirb.in Lamarck &Mirbel,Hist.Nat.Vég.3: 477 (1803) ≡Selaginellasubg.Stachygynandrum(P.Beauv.ex Mirb.) Baker,J.Bot.Lond.21: 3.1883 ≡Lycopodiumsubg.Stachygynandrum(P.Beauv.ex Mirb.) Spreng.,Anleit.,ed.2,2(1): 109 (1817) ≡L.(unranked)Stachygynandrum(P.Beauv.ex Mirb.) Hook.&Grev.,Bot.Misc.2: 380 (1831) ≡L.(unranked)Stachygynandrum(P.Beauv.ex Mirb.) Kunze,Linnaea 9: 8.1834,“Stachygynandra” ≡Didiclissubg.Stachygynandrum(P.Beauv.ex Mirb.)Rothm.,Feddes Repert.Spec.Nov.Regni Veg.54:70(1944)-Lectotype (designated by Pichi Sermolli,Webbia 26: 164(1971)):Selaginella flabellata(L.) Spring.

≡Selaginellasubg.HeterophyllumHieron.&Sadeb.in Engler&Prantl,Nat.Pflanzenf.l(4): 673 (1902) -Lectotype (designated by Zhou and Zhang,2015: 1136):Selaginella flabellata(L.)Spring.

Selaginella s.s.in this study corresponds toS.subg.StachygynandrumZhou and Zhang (2015) excludingS.sect.Ascendentes,S.sect.Circinatae,andS.sect.Plagiophyllae.The monophyly of the genus is strongly supported with nuclear data but its relationships are unresolved with plastid data.The combined dataset supported its monophyly strongly in BI analysis but weakly in ML and MP analyses.

Species ofSelaginella s.s.have erect,suberect,creeping,asscending (a few rosette-forming) plants.Megaspore surfaces are often reticulate (Fig.11J,M-P).Microspore surfaces are baculate,blunt spiny,verrucate,papillate,coarse,or ridge (Fig.11Q-T).

The newly defined genusSelaginella s.s.contains about 230 species nearly endemic to Americas except one (to a few) species in Africa.WithinSelaginella s.s.,four sections,S.sect.Austroamericanae,S.sect.Heterophyllae,S.sect.Poceres,andS.sect.Selaginella(=S.sect.Pallescentes sensuZhou and Zhang(2015)),can be recognized.

Section 1.Selaginellasect.Selaginella同穗卷柏组(新拟).

=Selaginellasect.HeterophyllaeSpring in Mart.,Fl.Bras.1(2):118 (1840) -Lectotype (designated by Zhou and Zhang,2015:1137):SelaginellaflexuosaSpring.

=S.(unranked)EnodesSpring in Mart.,Fl.Bras.1(2):118(1840)-Lectotype (designated by Zhou and Zhang,2015: 1136):S. flexuosaSpring.

=S.(unranked)ContinuaeSpring,Mém.Acad.Sci.Belg.24: 53(1850) ≡S.ser.Continuae(Spring) Hieron.&Sadeb.in Engl.&Prantl,Nat.Pflanzenf.1(4): 704 (1902) -Lectotype(designated by Zhou and Zhang,2015: 1137):Selaginella microphylla(Kunth)Spring.

=S.(unranked)PusillaeSpring,Mém.Acad.Sci.Belg.24: 53(1850) -Lectotype (designated by Zhou and Zhang,2015: 1137):S.microphylla(Kunth) Spring.

=S.(unranked)AscendentesA.Braun,App.Ind.Sem.Hort.Berol.11([1857]1858)-Lectotype(designated by Zhou and Zhang,2015:1137):S. flexuosaSpring.

=S.(unranked)PersistentesA.Braun,App.Ind.Sem.Hort.Berol.11([1857]1858)-Lectotype(designated by Zhou and Zhang,2015:1137):S. flexuosaSpring.

=S.sect.DichotropaeA.Braun,App.Ind.Sem.Hort.Berol.,11.([1857] 1858).nom.illeg.-Lectotype (designated by Zhou and Zhang,2015: 1137):S. flexuosaSpring.

=S.(unranked)ResupinataeA.Braun,App.Ind.Sem.Hort.Berol.12.([1857] 1858) -Lectotype (designated by Zhou and Zhang,2015: 1137):Selaginella stenophyllaA.Braun.

=S.ser.DecumbentesBaker,J.Bot.Lond.21: 3 (1883) -Lectotype (designated by Zhou and Zhang,2015: 1137):S.microphylla(Kunth) Spring.

=S.sect.PleiomacrosporangiataeHieron.&Sadeb.in Engl.&Prantl,Nat.Pflanzenf.1(4):673.(1902)-Lectotype(designated by Zhou and Zhang,2015: 1137):S. flexuosaSpring.

=S.ser.MonostelicaeHieron.&Sadeb.in Engl.&Prantl,Nat.Pflanzenf.1(4): 673 (1902) -Lectotype (designated by Zhou and Zhang,2015: 1137):S. flexuosaSpring.

≡S.sect.Stachygynandrum(P.Beauv.ex Mirb.)T.Moore,Ind.Fil.Cxxviii (1857) ≡Lycopodioidessect.Stachygynandrum(P.Beauv.ex Mirb.) Tzvel.,Novosti Sist.Vyssh.Rast.36: 25 (2004) -Type:S. flabellata(L.) Spring.

Because the genus has been proposed to be conserved with a conserved type,Selaginella flabellata(Wan et al.,2023),ourS.sect.Heterophyllaesensu Zhou and Zhang (2015) now becomes a synonym of the autonym.

This section is weakly supported as monophyletic in two but strongly in one of our three analyses based on the concatenated data (Fig.1) and is composed of New-World (South &Central Americas) species only.They are suberect,rarely creeping or erect(if plants erect,main stems with monomorphic leaves),and have dimorphic leaves and various chromosome numbers: 2n=18(S.simplexBaker)(Graustein,1930;Marcon et al.,2005),2n=20[S.erythropusSpring andS. flabellata(L.) Spring] (Jermy et al.,1967).

Section 2.Selaginellasect.AustroamericanaeLi Bing Zhang&X.M.Zhou,Taxon 64(6): 1136 (2015) 南美同穗卷柏组(新拟) -Type:S.hartwegianaSpring.

=S.(unranked)RedivivaeA.Braun,App.Ind.Sem.Hort.Berol.11([1857] 1858) -Lectotype (designated by Zhou and Zhang,2015:1136):S.ciliata(Willd.) A.Braun [=Selaginella novae-hollandiae(Sw.) Spring].

The monophyly of this section is weakly supported (Fig.1).Species of this section have stem nearly erect,different from those species with dimorphic sporophylls from America that often have creeping stems.They have dimorphic sterile leaves on stem,different from those species with erect stem and monomorphic sporophylls species from America that have monomorphic sterile leaves on stem (Valdespino,1993b).Moreover,at leastSelaginella hartwegianatend to have monomorphic leaves below first branch of the stems.

About 13 species of this section are sampled in our phylogenetic analysis (Fig.S3).This section mainly occurs in North to South America and a few possibly occur in Africa (currently onlySelaginella cathedrifoliaknown).

Section 3.Selaginella.sect.PallescentesLi Bing Zhang&X.M.Zhou白变同穗卷柏组(新拟),Taxon 64(6): 1137 (2015) -Type:S.pallescens(C.Presl) Spring.

Members of this section have monomorphic sporophylls[Selaginella apoda(L.)C.Morren has slightly dimorphic sporophylls,but generally its sporophylls are considered to be monomorphic;Valdespino,1993a],but variable numbers of chromosomes,e.g.,S.apoda: 2n=18;S.pallescens(C.Presl) Spring: 2n=20,22;Selaginella pulcherrimaLiebm.: 2n=20 (Tschermak-Woes and Doležal-Janish,1959;Jermy et al.,1967).

About eight species of this section were sampled in our phylogenetic analysis (Fig.S3).However,it is necessary that more morphological and molecular studies are needed to confirm which species in Americas should be placed in this section.

Section 4.Selaginellasect.Proceres(Spring) Li Bing Zhang &X.M.Zhou,Taxon 64(6): 1137 (2015) 高贵同穗卷柏组(新拟) ≡S.(unranked)ProceresSpring,Mem.Acad.Sci.Belg.24: 53 (1850) -Lectotype(designated by Zhou and Zhang,2015:1137):S.oaxacanaSpring.

This section is strongly supported as monophyletic and resolved as sister toSelaginellasect.pallescentes(Fig.1).Species of this section have monomorphic sporophylls and reticulate megaspore surfaces(S.martensiiSpring: Giorgi et al.,1997;Korall and Taylor,2006;S.oaxacanaSpring:Hellwig,1969].Only three species were included in our phylogenetic analysis(Fig.S3).More studies are needed.

Members of the genus:

Selaginella acanthostachysBaker,J.Bot.21: 99,no.40 (1883).

Selaginella achotalensisShelton &Caluff,Willdenowia 33(1):163 (2003).

Selaginella aculeatifoliaValdespino,Nordic J.Bot.34(3): 372(2016).

Selaginella acutifolia(Stolze) Valdespino,Mem.New York Bot.Gard.88 (Pterid.Mexico): 558 (2004).

Selaginella agioneumaValdespino&C.López,PeerJ 6(e4708):4(2018).

Selaginella alampetaM.Kessler &A.R.Sm.,Edinburgh J.Bot.63(1): 86 (2006).

Selaginella albolineataA.R.Sm.,Ann.Mo.Bot.Gard.77: 262,f.7A-E (1990).

Selaginella alstoniiG.Heringer,Salino&Valdespino,PhytoKeys 50: 65 (2015).

Selaginella altheaeValdespino,PhytoKeys 91: 15 (2017).

Selaginella amazonicaSpring,Mart.,Fl.Bras.1(2): 124(1840).

Selaginella anceps(C.Presl) C.Presl,Abh.Königl.Böhm.Ges.Wiss.,Math.-Naturw.Cl.V,3: 581 (repr.151) (1845).

Selaginella apoda(L.) Spring,Mart.,Fl.Bras.1(2): 119 (1840).

Selaginella applanataA.Braun,Ann.Sci.Nat.,Bot.,sér.V,3:274(1865).

Selaginella armataBaker,J.Bot.22: 90,no.125 (1884).

Selaginella arrectaA.R.Sm.,Ann.Mo.Bot.Gard.77(2): 264(1990).

Selaginella arsicladaValdespino,Am.Fern J.84(3): 99 (1994).

Selaginella ayitiensisValdespino,Willdenowia 49(1): 71-80(2019).

Selaginella bahiensisSpring,Mart.,Fl.Bras.1(2): 124,no.12(1840).

Selaginella bahiensissubsp.manausensis(Bautista) Jermy &J.M.Rankin,Bull.Brit.Mus.(Nat.Hist.) Bot.9(4): 260 (1981).

Selaginella barnebyanaValdespino,Mem.New York Bot.Gard.88 (Pterid.Mexico): 560-561 (2004).

Selaginella beiteliiA.R.Sm.,Ann.Mo.Bot.Gard.77(2): 264(1990).

Selaginella bernoulliiHieron.,Hedwigia 41: 192,no.15 (1902).

Selaginella blepharodellaValdespino,PhytoKeys 50: 68(2015).

Selaginella bombycinaSpring,Mém.Acad.Roy.Sci.Belg.24:191(1849).

Selaginella boomiiValdespino,Brittonia 67(4):329 (2015).

Selaginella braceiHieron.ex O.C.Schmidt,Repert.Spec.Nov.Regni Veg.20: 156 (1924).

Selaginella breedloveiValdespino,Mem.New York Bot.Gard.88(Pterid.Mexico): 563 (2004).

Selaginella brevifoliaBaker,J.Bot.21: 21(3): 83 (1883).

Selaginella brewerianaA.R.Sm.,Ann.Mo.Bot.Gard.77(2): 266(1990).

Selaginella breyniiSpring,Mart.,Fl.Bras.1(2): 121 (1840).

Selaginella brigitteanaValdespino,Willdenowia 49(1): 71-80(2019).

Selaginella bryophilaM.Kessler &A.R.Sm.,Edinburgh J.Bot.63(1): 91 (2006).

Selaginella cabrerensisHieron.,Hedwigia 43: 29 (1904).

Selaginella calostichaSpring,Mém.Acad.Roy.Sci.Belgique 24[Monogr.Lyc.2]: 206,no.145 (1849).

Selaginella cardiophyllaValdespino,Brittonia 44(2): 199(1992).

Selaginella carioiHieron.,Engl.&Prantl,Nat.Pflanzenf.1(4):688 (1901 publ.1902).

Selaginella cathedrifoliaSpring,Mém.Acad.Roy.Sci.Belgique 24[Monogr.Lyc.2]: 112,no.58 (1849).

Selaginella cavernariaCaluff &Shelton,Willdenowia 44(3): 311(2014).

Selaginella cavifoliaA.Braun,Ann.Sci.Nat.,Bot.,sér.V,5,3:272(1865).

Selaginella cheiromorphaAlston,Jermy &J.M.Rankin,Bull.Brit.Mus.(Nat.Hist.) Bot.9(4): 257 (1981).

Selaginella chiapensisA.R.Sm.,Am.Fern J.70(1):25 (1980).

Selaginella chionolomaAlston ex Crabbe &Jermy,Am.Fern J.63: 137 (1973).

Selaginella chrysoleucaSpring,Bull.Acad.Roy.Soc.Bruxelles 10(1): 226 (1843).

Selaginella confusaSpring,Flora 21: 218 (1838).

Selaginella contiguaBaker,J.Bot.22: 295,no.162 (1884).

Selaginella cordifolia(Desv.ex Poir.)Spring,Bull.Acad.Roy.Soc.Bruxelles 10: 228 (1843).

Selaginella correaeValdespino,Brittonia 45(4): 315 (1993).

Selaginella corrugisMickel &Beitel,Pterid.Fl.Oaxaca (Mem.New York Bot.Gard.46): 336(1988).

Selaginella crinitaValdespino,PhytoKeys 50: 74 (2015).

Selaginella cristalensisShelton &Caluff,Willdenowia 33(2):435 (2003).

Selaginella cruciformisAlston ex Crabbe &Jermy,Fern Gaz.11(4): 257 (1976).

Selaginella culverwelliiN.R.Crouch,Ferns S.Afr.Compreh.Guide: 746 (2011).

Selaginella cuneataMickel &Beitel,Pterid.Fl.Oaxaca (Mem.New York Bot.Gard.46: 336(1988).

Selaginella cyclophyllaA.R.Sm.,Ann.Mo.Bot.Gard.77(2): 266(1990).

Selaginella dasylomaAlston,J.Bot.72: 228 (1934).

Selaginella delicatissimaLinden,Cat.n.11: 20 (1856),nomen,ex A.Braun,App.Ind.Sem.Hort.Berol.13 (1857).

Selaginella dendricolaJenman,Gard.Chron.,ser.3,2: 99(1887).

Selaginella densifoliaSpruce ex Hook.,Hook.,2nd Cent.,t.85(1861) (non Klotzsch,1844).

Selaginella denudata(Willd.) Spring,Flora 21: 212 (1839).

Selaginella eatoniiHieron.ex Small,Ferns trop.Florida 67(1918).

Selaginella eclipesW.R.Buck,Canad.J.Bot.55: 366 (1977).

Selaginella epipubensCaluff&Shelton,Willdenowia 39(1):116(2009).

Selaginella epirrhizosSpring,Bull.Acad.Roy.Soc.Bruxelles 10:229,no.126 (1843).

Selaginella erectifoliaSpring,Mém.Acad.Roy.Sci.Belgique 24[Monogr.Lyc.2]: 92,no.36 (1849).

Selaginella erythropus(Mart.) Spring,Mart.,Fl.Bras.1(2): 125(1840).

Selaginella euclimaxAlston ex Crabbe &Jermy,Fern Gaz.11:259 (1976).

Selaginella falcata(P.Beauv.) Spring,Bull.Acad.Roy.Soc.Bruxelles 10: 225,no.91 (1843).

SelaginellafinitimaMickel &Beitel,Pterid.Fl.Oaxaca (Mem.New York Bot.Gard.46: 338 (1988).

Selaginellaflabellata(L.) Spring,Flora 21: 198,no.19 (1838).

SelaginellaflaccaAlston,Jermy &J.M.Rankin,Bull.Brit.Mus.(Nat.Hist.) Bot.9(4): 269 (1981).

Selaginellaflagellata(L.) Spring,Bull.Acad.Roy.Soc.Bruxelles 10(1): 228 (1843).

Selaginella flexuosaSpring,Flora 21: 197,no.18 (1838).

Selaginella fragillimaSilveira,Bol.Comm.Geogr.Geol.Est.Minas Gerais 5: 127,t.11 (1898).

Selaginella gioiaeValdespino,PhytoKeys 159: 74 (2020).

Selaginella glossophyllaCrabbe &Jermy,Fern Gaz.11: 259(1976).

Selaginella guatemalensisBaker,J.Bot.21: 243 (1883).

Selaginella gynostachyaValdespino,Fern Gaz.18(2): 42 (2007 publ.2008).

Selaginella haematodes(Kunze) Spring,in Mart.,Fl.Bras.1(2):126 (1840).

Selaginella haenkeanaSpring,Bull.Acad.Roy.Soc.Bruxelles 10:225 (1843).

Selaginella harrisiiUnderw.&Hieron.,Symb.Ant.7:162(1912).

Selaginella hartiiHieron.,Urban,Symb.Ant.3: 525 (1903).

Selaginella hartwegianaSpring,Mém.Acad.Roy.Sci.Belg.24:188 (1849).

Selaginella hemicardiaValdespino,Brittonia 44(2):201(1992).

Selaginella heterodonta(Desv.ex Poir.) Hieron.,Symb.Ant.9:392 (1925).

Selaginella hirsutaAlston ex Crabbe &Jermy,Am.Fern J.63:138 (1973).

Selaginella hirtifoliaValdespino,Mem.New York Bot.Gard.88(Pterid.Mexico): 573-574 (2004).

Selaginella hispida(Willd.)A.Braun ex Urb.,Symb.Ant.9: 394(1925).

Selaginella homaliaeA.Braun,Ann.Sci.Nat.,Bot.,sér.V,3:274(1865).

Selaginella huehuetenangensisHieron.,Hedwigia 43:32(1904).

Selaginella hyalogrammaValdespino,Am.Fern J.107(2): 74(2017).

Selaginella idiosporaAlston,Bull.Brit.Mus.(Nat.Hist.) Bot.1(8): 246,t.6,f.A-E (1955).

Selaginella illecebrosaAlston,Bull.Brit.Mus.(Nat.Hist.) Bot.1(8): 239,t.5,f.A-E(1955).

Selaginella imbricansA.R.Sm.,Ann.Mo.Bot.Gard.77: 266,f.9K-N (1990).

Selaginella jungermannioides(Gaudich.) Spring,Bull.Acad.Roy.Soc.Bruxelles 10: 143 (1843).

Selaginella karowtipuensisValdespino,Fern Gaz.18(2): 46(2007 publ.2008).

Selaginella kochiiHieron.,Koch-Grünberg Reisen in Nordwest-Brasilien,2: 361 (1910).

Selaginella kriegerianaL.A.Góes,Syst.Bot.43(4): 921 (2018).

Selaginella krugiiHieron.,Symb.Ant.3: 526 (1903).

Selaginella laxifoliaBaker ex Krug ex Urb.,Bot.Jahrb.Syst.24:151 (1897).

Selaginella lechleriHieron.,Engl.&Prantl,Nat.Pflanzenf.1(4):683 (1901).

Selaginella leonardiiO.C.Schmidt,Fedde,Repert.20: 155(1924).

Selaginella leucolomaAlston,Crabbe &Jermy,Fern Gaz.11(4):262 (1976).

Selaginella lindeniiSpring,Bull.Acad.Roy.Soc.Bruxelles 10:142,no.52 (1843).

Selaginella lineolataMickel &Beitel,Pterid.Fl.Oaxaca (Mem.New York Bot.Gard.46): 343 (1988).

Selaginella longissimaBaker,J.Bot.19: 208 (1881).

Selaginella ludoviciana(A.Braun)A.Braun,Ann.Sci.Nat.,Bot.,sér.4,13: 58 (1860).

Selaginella lychnuchusSpring ex Klotzsch,Linnaea 20: 435(1847),nomen,ex Spring,Monogr.Lyc.2: 247,no.186 (1849).

Selaginella macrostachya(Spring) Spring,Bull.Acad.Roy.Soc.Bruxelles 10: 144 (1843).

Selaginella magnafornensisValdespino &C.López,PeerJ 6(e4708): 11 (2018).

Selaginella marahuacaeA.R.Sm.,Ann.Mo.Bot.Gard.77:268,f.7F-I (1990).

Selaginella martensiiSpring,Mém.Acad.Roy.Sci.Belg.24:129(1849).

Selaginella mazaruniensisJenman,Gard.Chron.22:210(1897).

Selaginella mendoncaeHieron.,Engl.&Prantl,Nat.Pflanzenf.1(4): 693,no.233.(1901 publ.1902).

Selaginella meridensisAlston,Bull.Brit.Mus.(Nat.Hist.) Bot.9(4): 267 (1981).

Selaginella mickeliiValdespino,Brittonia 44(3): 316 (1992).

Selaginella microdendronBaker,J.Bot.23: 116 (1885).

Selaginella microdontaA.C.Sm.,Bull.Torrey Bot.Club 58: 313(1931).

Selaginella microphylla(Kunth) Spring,Bull.Acad.Roy.Soc.Bruxelles 10: 234 (1843).

Selaginella minimaSpring,Bull.Acad.Roy.Soc.Bruxelles 10:139,no.29 (1843).

Selaginella mixtecaMickel &Beitel,Pterid.Fl.Oaxaca (Mem.New York Bot.Gard.46): 345 (1988).

Selaginella mollisA.Braun,Ann.Sci.Nat.,Bot.,sér.V,5,3: 276(1865).

Selaginella monticolaValdespino,Phytotaxa 233(1): 154(2015).

Selaginella moritzianaSpring ex Klotzsch,Linnaea 20: 436(1847).

Selaginella moritzianavar.pearcei(Baker) Valdespino,Taxon 65(6): 1402 (2016).

Selaginella mortonianaCrabbe &Jermy,Am.Fern J.63: 139(1973).

Selaginella mosorongensisHieron.,Hedwigia 43: 4 (1904).

Selaginella mucronataG.Heringer,Salino &Valdespino,PhytoKeys 50: 78 (2015).

Selaginella mucugensisValdespino,PhytoKeys 50: 82 (2015).

Selaginella muscosaSpring,Mart.,Fl.Bras.1(2): 120 (1840).

Selaginella myriostachyaValdespino,C.López &Góes-Neto,Phytotaxa 184(4): 235-244 (2014).

Selaginella nanophyllaValdespino,C.López &Góes-Neto,Phytotaxa 184(4): 235-244 (2014).

Selaginella nanuzaeValdespino,PhytoKeys 57: 98 (2015).

Selaginella neblinaeA.R.Sm.,Ann.Mo.Bot.Gard.77(2): 268(1990).

Selaginella neospringianaValdespino,PhytoKeys 57: 103(2015).

Selaginella nothohybridaValdespino,Brittonia 44(3): 319(1992).

Selaginella novae-hollandiae(Sw.) Spring,Bull.Acad.Roy.Soc.Bruxelles 10: 234,no.161 (1843).

Selaginella oaxacanaSpring,Mém.Acad.Roy.Sci.Belg.24:177(1849).

Selaginella ophiodermaValdespino,Nordic J.Bot.2022(3)-e03411: 2 (2022).

Selaginella orbiculifoliaShelton &Caluff,Willdenowia 33(2):430 (2003).

Selaginella orizabensisHieron.,Hedwigia 41:193,no.16(1902).

Selaginella osaensisA.Rojas,Revista Biol.Trop.49(2): 446(2001).

Selaginella ovifoliaBaker,J.Bot.22: 90 (1884).

Selaginella pallescens(C.Presl)Spring,Mart.,Fl.Bras.1(2):132(1840).

Selaginella palmiformisAlston ex Crabbe &Jermy,Am.Fern J.63: 141 (1973).

Selaginella panurensisBaker,J.Bot.21: 97 (1883).

Selaginella papillosaValdespino,PhytoKeys 159: 81 (2020).

Selaginella pellucidopunctataValdespino,PhytoKeys 57: 107(2015).

Selaginella phiaraValdespino,C.López &L.A.Góes,Phytotaxa 184: 242 (2014).

Selaginella philipsonii(Jermy &J.M.Rankin) Valdespino,PhytoKeys 91: 32 (2017).

Selaginella plagiochilaBaker,J.Bot.21: 212,no.67(1883).

Selaginella plumieriHieron.,Urban,Symb.Ant.7: 488 (1913).

Selaginella polypteraValdespino,Mem.New York Bot.Gard.88(Pterid.Mexico): 588 (2004).

Selaginella popayanensisHieron.,Hedwigia 43: 9 (1904).

Selaginella porelloides(Lam.) Spring,Bull.Acad.Roy.Soc.Bruxelles 10(1): 141 (1843).

Selaginella porphyrosporaA.Braun,Ann.Sci.Nat.,Bot.,sér.V,5,3: 286 (1865).

Selaginella potaroensisJenman,Gard.Chron.3,2: 154 (1887).

Selaginella praestansAlston,Bull.Brit.Mus.(Nat.Hist.) Bot.9:260 (1981).

Selaginella prasinaBaker,J.Bot.22: 113,no.134 (1884).

Selaginella productaBaker,J.Bot.21(8): 243(1883).

Selaginella proliferaValdespino,Mem.New York Bot.Gard.88(Pterid.Mexico): 590 (2004).

Selaginella pruskianaValdespino,Brittonia 44(2): 202 (1992).

Selaginella psittacorrhynchaValdespino,Phytoneuron 63: 2(2017).

Selaginella pubensA.R.Sm.,Ann.Mo.Bot.Gard.77(2): 269(1990).

Selaginella pubimarginataValdespino,PhytoKeys 159: 87,9-12 (2020).

Selaginella pulcherrimaLiebm.ex E.Fourn.,Mexic.Pl.1: 147(1872).

Selaginella quadrifariaAlston,Jermy &Rankin,Bull.Brit.Mus.(Nat.Hist.) Bot.9(4): 261 (1981).

Selaginella rachipterygiaValdespino,Cornejo &C.López,Novon 30(1): 70(2022).

Selaginella radiata(Aubl.)Spring,Bull.Acad.Roy.Soc.Bruxelles 10: 143,no.54 (1843).

Selaginella ramosissimaBaker,J.Bot.23: 295 (1885).

Selaginella raynalianaTardieu,Adansonia 3(3):352-353,t.1,f.1-9 (1963).

Selaginella reflexaUnderw.,Bull.Torrey Bot.Club 21:268(1894).

Selaginella revolutaBaker,J.Bot.21(5): 141 (1883).

Selaginella rhodostachyaBaker,Timehri 5: 221 (1886),also Trans.Linn.Soc.,Bot.2: 296 (1887).

Selaginella roraimensisBaker,Timehri 5:221(1886),also Trans.Linn.Soc.,Bot.2: 295 (1887)

Selaginella roseaAlston,J.Bot.70: 281-282 (1932).

Selaginella rostrataValdespino,PhytoKeys 159: 93,9 and 13(2020).

Selaginella rotundifoliaSpring,Bull.Acad.Roy.Soc.Bruxelles 10(1): 139 (1843).

Selaginella rzedowskiiLorea-Hem.,Bol.Soc.Bot.Mexico 44:24(1983).

Selaginella salazariaeValdespino,Brittonia 45(4): 319 (1993).

Selaginella salinoiL.A.Góes &G.Heringer,Phytotaxa 224(3):292 (2015).

Selaginella saltuicolaValdespino,PhytoKeys 50: 85 (2015).

Selaginella sandwithiiAlston,Jermy &J.M.Rankin,Bull.Brit.Mus.(Nat.Hist.) Bot.9(4): 289 (1981).

Selaginella scalariformisA.C.Sm.,Bull.Torrey Bot.Club 58:314(1931).

Selaginella schultesiiAlston ex Crabbe&Jermy,Am.Fern J.63:141 (1973).

Selaginella seemanniiBaker,J.Bot.21: 244 (1883).

Selaginella sematophyllaValdespino,G.Heringer &Salino,PhytoKeys 50: 89 (2015).

Selaginella serpens(Desv.ex Poir.) Spring,Bull.Acad.Roy.Soc.Bruxelles 10(1): 228 (1843).

Selaginella simplexBaker,J.Bot.23: 293 (1885).

Selaginella smithiorumValdespino,Brittonia 45(4): 322(1993).

Selaginella soboliferaA.R.Sm.,Ann.Mo.Bot.Gard.77(2): 269(1990).

Selaginella solomoniiValdespino,Novon 24(1): 100 (2015).

Selaginella speciosaA.Braun,Ann.Sci.Nat.,Bot.,sér.V,3:274-275,no.7 (1865).

Selaginella squamulosaValdespino,PhytoKeys 91: 24(2017).

Selaginella stenophyllaA.Braun,App.Ind.Sem.Hort.Berol.22(1857).

Selaginella stomatolomaValdespino,PhytoKeys 57:113(2015).

Selaginella striataCaluff &Shelton,Willdenowia 44(3): 313(2014).

Selaginella subrugosaMickel&Beitel,Pterid.Fl.Oaxaca(Mem.New York Bot.Gard.46: 352(1988).

Selaginella substipitataSpring,Bull.Acad.Roy.Soc.Bruxelles 10: 227,no.110 (1843).

Selaginella surucucusensisL.A.Goés &E.L.M.Assis,Kew Bull.72(40): 1 (2017).

Selaginella tanycladaAlston ex Crabbe&Jermy,Am.Fern J.63:143 (1973).

Selaginella tayloriiValdespino,Brittonia 45(4): 320 (1993).

Selaginella terezoanaBautista,Bol.Mus.Paraense Emilio Goeldi,Bot.,45: 1 (1974).

Selaginella thysanophyllaA.R.Sm.,Ann.Mo.Bot.Gard.77(2):270(1990).

Selaginella truncataH.Karst.ex A.Braun,Index Sem.Hort.Berol.1857: 15 (1857).

Selaginella trygonoidesValdespino,PhytoKeys 57: 116 (2015).

Selaginella tuberculataSpruce ex Baker,J.Bot.21(3):83(1883).

Selaginella tuberosaB.McAlpin &Lellinger,Brenesia 24: 409(1986).

Selaginella tyleriA.C.Sm.,Bull.Torrey Bot.Club 58:311(1931).

Selaginella umbrosaLem.ex Hieron.,Engl.&Prantl,Nat.Pflanzenf.1(4): 683,f.404 (1901).

Selaginella undataShelton &Caluff,Willdenowia 33(2): 427(2003).

Selaginella urquiolaeCaluff&Shelton,Willdenowia 39(1):112(2009).

Selaginella valdepilosaBaker,J.Bot.21: 82 (1883).

Selaginella ventricosaValdespino&C.López,PeerJ 6(e4708):15(2018).

Selaginella vernicosaBaker,Timehri 5: 220 (1886),also Trans.Linn.Soc.,Bot.2: 295 (1887).

Selaginella vestiensBaker,J.Bot.21: 97,no.34 (1883).

Selaginella viticulosaKlotzsch,Linnaea 18: 524(1844).

Selaginella wolffiiSodiro,Crypt.Vasc.Quit.620 (1893),also Anales Univ.Centr.Ecuador 12(83): 490 (1895).

Selaginella wurdackiiAlston,Bull.Brit.Mus.(Nat.Hist.)Bot.9:280 (1981).

Selaginella xanthoneuraValdespino,PhytoKeys 159:98,figs.5,14-18 (2020).

Selaginella xiphophyllaBaker,J.Bot.22(10):296 (1884).

Selaginella zartmaniiValdespino,C.López &A.M.Sierra,PeerJ 6(e4708): 21 (2018).

5.9.Incertae sedis

Selaginella birarensisKuhn,Forschungsr.Gazelle,Bot.Alg.19(1889).

Selaginella boschaiHieron.,Hedwigia 51: 243,no.2 (1912).

Selaginella dahliiHieron.,Engl.Bot.Jahrb.50:2,10,no.2(1913).

Selaginella firmuloidesWarb.,Monsunia 1: 105,118,no.77(1900).

Selaginella ketra-ayamAlderw.,Bull.Jard.Bot.Buitenz.II,1:24-25 (1911).

Selaginella ledermanniHieron.,Engl.Bot.Jahrb.56: 224,no.6(1920).

Selaginella pilosulaAlderw.,Bull.Jard.Bot.Buitenzorg,ser.3,5:233(1922).

Selaginella politaRidl.,J.Fed.Mal.States Mus.6: 202,no.348(1915).

Selaginella preslianaSpring,Bull.Acad.Roy.Soc.Bruxelles 10:137,no.16 (1843).

Selaginella propinquaAlderw.,Bull.Jard.Bot.Buitenz.III,5:233(1922).

Selaginella pseudovolkensiiHosok.,Trans.Nat.Hist.Soc.Taiwan 31: 471 (1941).

Selaginella ridleyiBaker,Ann.Bot.8: 131,no.58 (1894).

Selaginella schaffneriHieron.,Nat.Pflanzenf.1: 674 (1902).

Selaginella sonneratiiHieron.,Engl.Bot.Jahrb.50: 2,7,no.1(1913) [soneratii].

Declaration of competing interest

The authors declare no conflict of interest.

Author contributions

Conceptualization: LBZ;Investigation: XMZ;Writing,Reviewing,and Editing: LBZ and XMZ;Visualization: XMZ and LBZ;Supervision: LBZ.

Conflicts of Interest

We declare no conflicts of interest.

Acknowledgments

The research was partially supported by the Natural Science Foundation of China(#31900186,#32260050),Yunnan Fundamental Research Projects (Grant NO.202301BF07001-016),and the Glory Light International Fellowship for Chinese Botanists at Missouri Botanical Garden (MO) to X.M.Zhou.We thank Arthur Haines,Michael Hassler,Petra Korall,Carl Rothfels,Alan Smith,and Alan Weakley for helpful email discussion in Feb.2021,Carl Rothfels,Timothée Le Péchon and other collaborators for sharing materials for our earlier studies,Atsushi Ebihara and Noriaki Murakami for sharing specimen images,and Jing Zhao,Jian-Jun Yang,Shao-Li Fang,and Yu-Xin Li for helping with the figures,Robbin Moran and the owner of plants-of-styria.uni-graz.at/images/selaginella-selaginoides.html for allowing us to use their online images,and Alan Weakley and three anonymous reviewers for helpful comments.

Appendix A.Supplementary data

Supplementary data to this article can be found online at https://doi.org/10.1016/j.pld.2023.07.003.