Dryopteris (Dryopteridaceae) is among the most common and species rich fern genera in temperate forests in the northern hemisphere containing 225–300 species worldwide. The circumscription of Dryopteris has been controversial and various related genera have, over the time, been included in and excluded from Dryopteris. The infrageneric phylogeny has largely remained unclear, and the placement of the majority of the supraspecific taxa of Dryopteris has never been tested using molecular data.The fern genus rbcL gene, rps4-trnS spacer, trnL intron, trnL-F spacer) were used to reconstruct the phylogeny of Dryopteris. A total of 122 accessions are sampled in our analysis and they represent 100 species of the expanded Dryopteris including Acrophorus, Acrorumohra, Diacalpe, Dryopsis, Nothoperanema, and Peranema. All four subgenera and 19 sections currently recognized in Dryopteris s.s. are included. One species each of Arachniodes, Leptorumohra, and Lithostegia of Dryopteridaceae are used as outgroups. Our study confirms the paraphyly of Dryopteris and provides the first strong molecular evidence on the monophyly of Acrophorus, Diacalpe, Dryopsis, Nothoperanema, and Peranema. However, all these monophyletic groups together with the paraphyletic Acrorumohra are suggested to be merged into Dryopteris based on both molecular and morphological evidence. Our analysis identified 13 well-supported monophyletic groups. Each of the 13 clades is additionally supported by morphological synapomophies and is inferred to represent a major evolutionary lineage in Dryopteris. In contrast, monophyly of the four subgenera and 15 out of 19 sections currently recognized in Dryopteris s.s is not supported by plastid data.In this study, DNA sequences of four plastid loci  in Dryopteris strongly suggests that the current taxonomy of this genus is in need of revision. The disagreement between the previous taxonomy and molecular results in Dryopteris may be due partly to interspecific hybridization and polyplodization. More morphological studies and molecular data, especially from the nuclear genome, are needed to thoroughly elucidate the evolutionary history of Dryopteris. The 13 well-supported clades identified based on our data represent 13 major evolutionary lineages in Dryopteris that are also supported by morphological synapomophies.The genera,  Dryopteris Adans. (Dryopteridaceae) is estimated to contain 225 to 300 species worldwide  resolving phylogenetic relationships between Dryopteris and Acrophorus, Acrorumohra, Diacalpe, Dryopsis, Dryopteris, Nothoperanema, and Peranema; [3] assessing the monophylies of supraspecific taxa at sectional and subgeneric ranks recognized in current classifications using relatively large sampling and DNA sequences of multiple loci; and [4] identifying major evolutionary lineages in Dryopteris.The goals of this study include: [1] vigorously testing the monophyly of trnL intron and trnL-F spacer yielded similar tree topologies in both MP and ML analyses (trees not shown). The most parsimonious, parsimony JK, and likelihood JK and BS trees for all analyses are available upon request from the first author. There were no well-supported  clades Dryopteris.The combined data matrix of four plastid regions consisted of 3,638 bases. A simultaneous analysis ,30 of nuUnweighted MP simultaneous analysis generated 1,785,800 most parsimonious trees with a length of 1,831 steps, a consistency index  of 0.83Dryopteris in its current circumscription is paraphyletic in relation to AcrophorusAcrorumohraDiacalpeDryopsisNothoperanema, and Peranema , and AcrorumohraDryopsis, and Nothoperanema, should all be merged into Dryopteris, though each is monophyletic except Acrorumohra. The expanded Dryopteris is supported as monophyletic with strong support , and Peranamea. Such a close relationship among all of them had not previously been suggested in the literature, although the close affinities among Peranemataceae, Dryopsis (previously in Tectariaceae), and Dryopteridaceae have partially long been noticed , and Dryopsis as independent genera, in addition to Dryopteris and another 24 or 25 genera (with one being Incertae Sedis), in his large subfamily Dryopteridoideae, one of the two subfamilies in Dryopteridaceae sensu lato (the other one is Athyrioideae). Recognition of these genera in Dryopteridaceae was largely followed by Smith et al.’s [The family Peranemataceae was established by Ching ; as “Per Peranema-35,37,38 Peranema recognizet al.’s  classifiet al.’s  are highet al.’s ,40).Nothoperanema, normally not viewed as a member of Peranemataceae, is embedded within Peranemataceae. In our analyses, Nothoperanema and Peranemataceae together formed a strongly supported monophyletic group in our analyses , sister to the Aemulae clade within Dryopteris (see below). Our molecular data unambiguously resolved the species of Peranemataceae as members of Dryopteris . Based on our study, Acrophorus belongs to Dryopteris, and represents a specialized group within Dryopteris with round indusia and cordate scales at costa base.The genus s  and Diacalpe and Peranema are paraphyletic in relation to Acrophorus and Nothoperanema, contrasting Nayar & Kaur’s [Diacalpe as a synonym of Peranema while recognizing Acrophorus. Our resolution of Diacalpe is consistent with Liu et al. [Diacalpe were sampled. Morphologically, Diacalpe is characterized by unstalked sori, a few single-celled long and clavate paraphyses on the lower portion of sporangiate stalk, and entire scales at the stipe bases [Sermolli , Wu [33,Sermolli , Wu & ChSermolli , and ChrSermolli , but is r & Kaur  who alsou et al.  and our u et al. , Kramer u et al. , and Smiu et al.  also tre& Kaur’s , Kramer’& Kaur’s , and Smi& Kaur’s  treatmenu et al.  where onpe bases .Diacalpe, D. annamensis Tagawa is resolved as sister to the rest of species, with D. chinensis Ching & S. H. Wu then sister to D. aspidioides + D. christensenae Ching  Fraser-Jenk. Acrorumohra is well defined morphologically. Its pinnules are all anadromous and the terminal pinnules have asymmetrical bases, different from Dryopteris. Ten accessions of three species of Acrorumohra, including the type, A. diffracta (Baker) H. Itô, are sampled in our study. Our analyses demonstrate, for the first time, that Acrorumohra is paraphyletic in relation to Dryopteris polita Rosenst. D. polita is sister to A. hasseltii (Blume) Ching plus A. subreflexipinna (Ogata) H. Itô, and together these three are sister to A. diffracta. Our results clearly show that Acrorumorha is a member of Dryopteris.Originally described as a subgenus, a H. Itô , and lata H. Itô , Acrorumd  Ching. For the first time, Nothoperanema is supported as monophyletic in our analyses  Ching and N. shikokianum (Makino) Ching needs further clarifications.Within Dryopsis was established by Holttum & Edwards [Ctenitis subgen. Dryopsis Ching”. It is now widely recognized [CtenitisDryopsis, and Dryopteris have been controversial. Morphologically, Dryopsis appears to be more distant from Ctenitis than from Dryopteris. Dryopsis has distinct venation on the abaxial surfaces, sori terminal on veinlets, and marginal, entire scales that are clathrate or not, but with long and dull areolae. Ctenitis has venation indistinct on both the adaxial and abaxial surfaces, sori middle on the veinlets, and scales ciliate on their margins, clathrate, and with nearly hexagonal and lustrous areolae [Dryopsis and Dryopteris is that the former has either shallow or deep rachis and costa grooves that are closed near their bases, as well as multi-celled hairs with a thickened base on the margins but not in the grooves of the rachis and costae. Dryopteris, in contrast, always has deep rachis and costa grooves that are connected near their bases, and normally has no hairs on the rachis or costae [The genus  Edwards  based oncognized ,39,53,54 areolae ,13,54. Tr costae ,13,54.Dryopsis contains about 22 species [Dryopsis should be a member of Dryopteridaceae but Liu et al. [DryopsisDryopteris, and Peranemataceae sensu Ching [Dryopsis is not closely related to Ctenitis. species  occurrin species ,55. With species  discoveru et al.  failed tsu Ching ,37, and su Ching . Liu et su Ching  also conDryopsis, including the type, D. apiciflora  Holttum & P. J. Edwards, are sampled in our analysis. Our results demonstrate that Dryopsis is monophyletic , in contrast to the resolution of Li & Lu [Dryopsis formed an unresolved trichotomy with two species of Dryopteris and one species of Acrorumohra. Our results also indicate that Dryopsis is nested within a paraphyletic Dryopteris .Six accessions of five species of  Li & Lu , where ts Figure , stronglDryopsis, the species sampled were resolved into two clades. Morphologically, species of the upper clade  Holttum & P.J. Edwards, and D. sp.) have bullate scales, while those of the lower clade (D. heterolaena (C. Chr.) Holttum & P.J. Edwards, D. mariformis (Rosenst.) Holttum & P.J. Edwards) have flat scales  were represented by two or more species in our study (Appendix I).Our 100-species sampling is still not dense enough to rigorously test the monophylies of all supraspecific taxa (sections or subgenera) recognized in recent classifications by Fraser-Jenkins ,24,25, aween 225 , and in Dryopteris strongly suggests that the current taxonomy of this genus is in need of revision. However, our data do not necessarily falsify the monophyly of these 19 sections. The disagreement between previous taxonomy and molecular results in Dryopteris may be due partly to interspecific hybridization and polyplodization [The non-monophyly of the 19 out of the 21 supraspecific taxa in dization ,58.Dryopteris that have evolved via inter-clade hybridization, based on plastid trnL-F sequences, nuclear PgiC sequences, and/or biochemical evidence. D. guanchica, limited to Spain, Portugal, and the Canary Islands, has been postulated to be of hybrid origin between D. aemula  and possibly D. intermedia (D. sect. Lophodium) [D. shibipedis Sa. Kurata, an obvious member of D. sect. Variae judging from the morphology [D. kinkiensis  and D. pacifica (Nakai) Tagawa  [D. corleyi, an endemic of northern Spain, is of hybrid origin between D. aemula (D. sect. Aemulae) and D. oreades Fomin . Our analyses based on plastid data and the resolution of D. corleyi as sister to D. aemula suggest that D. aemula is the maternal progenitor of D. corleyi. In addition, Sessa et al. [Dryopteris that has involved inter-continental long-distance dispersal as well as inter-clade hybridization. These examples of hybridization not only highlight the importance of reticulate evolution and thus the importance of nuclear data in understanding the evolutionary history of Dryopteris, but also strongly support the inclusion of these 13 lineages, including the small segregates, within Dryopteris, as opposed to breaking Dryopteris into many small genera.There are four well-documented allopolyploids in phodium) . The Japrphology ,42, has e clade) . Using ae clade)  concludea et al.  found evDryopteris : This clade contains species of Peranemataceae sensu Ching , Wu 33,,33,34, aDryopteris aemula and D. corleyi, based on the current sampling. These two species are different enough morphologically to have been placed in different sections by Fraser-Jenkins [D. aemula is consistent with those of Geiger & Ranker [D. corleyi, an endemic of northern Spain, is thought to be of hybrid origin between D. aemula (D. sect. Aemulae) and D. oreades Fomin based on allozyme data  Kuntze, and D. hawaiiensis (Hillebrand) W. Robinson, but D. guanchica is an allotetraploid (see blow) and D. hawaiiensis possibly an allotriploid [2. The Aemulae clade : The Aemulae clade, or the Hawaiian glabra group , contain& Ranker , Juslén & Ranker , and Ses& Ranker . This isme data , and D. polita (D. sect. Politae). The morphological synapomorphy is the flat scales in comparison with the Dryopsis clade, the Erythrovariae clade, and the Variae clade. The gain of bullate scales is considered here as the morphological synapomorphy of the expanded D. subgen. Erythrovariae including the Acrorumohra clade, the Dryopsis clade, the Erythrovariae clade, and the Variae clade.3. The Acrorumohra clade : This clade contains species of Dryopteris sect. Variae Fraser-Jenk. It is characterized by having slightly bullate-based scales and stiffly coriaceous lamina and pinnules with caudate apices and pointed lobes [4. The Variae clade : The Variae clade contains species of ed lobes ,26.Dryopteris sect. Erythrovariae plus D. podophylla. It is characterized by having more bullate scales and herbaceous lamina and pinnules with acute apices and rounded lobes [5. The Erythrovariae clade : This clade contains species of ed lobes ,26.Dryopsis. The relatively moderate branch support may be the result of some missing sequence data for members of this clade. The potential major morphological synapomorphies are the rachis and costa grooves that are closed near their bases and the multi-cellular hairs (see above).6. The Dryopsis clade : This clade contains species of the former genus Dryopteris sect. Cinnamomeae and one species of D. sect. Purpurascentes in our current sampling. This clade is defined by having pinnules angled acroscopically and usually with narrower bases and having linear scales on stipe base [7. The Cinnamomeae clade : The Cinnamomeae clade contains two species of Dryopteris crinalis (Hook. & Arn.) C. Chr., D. mauiensis C. Chr., D. sandwiciensis (Hook. & Arn.) C. Chr., D. tetrapinnata W. H. Wagner & Hobdy, and D. unidentata (Hook. & Arn.) C. Chr. The potential morphological synapomorphy is the absence of indusia [8. The Crinales clade : This clade was named the Hawaiian exindusiate group by Palmer  and it c indusia ,62.Dryopteris sect. Pallidae sensu Fraser-Jenkins [D. aitoniana Pic. Serm., D. odontoloma (Bedd.) C. Chr., D. pallida (Bory) C. Chr. ex Maire & Petitm., and D. mindshelkensis N. Pavl. (synonym: D. submontana (Fraser-Jenk. & Jermy) Fraser-Jenk.), and additional species from other sections, e.g., D. goldiana (Hook.) A. Gray, D. monticola (Makino) (D. sect. Dryopteris), D. oligodonta (Desv.) Pic. Serm. (D. sect. Marginatae), and D. tokyoensis (Matsum. & Makino) C. Chr. (D. sect. Pandae), based our current sampling and Juslén et al. [D. odontoloma in this clade needs further study. This clade is weakly supported as sister to the Crinales clade : This clade contains some species of -Jenkins , e.g., Dn et al. . The incDryopteris sect. Nephrocystis sensu Fraser-Jenkins [10. The Nephrocystis clade : This clade contains those species of -Jenkins .Dryopteris subgen. Dryopteris sensu Fraser-Jenkins [D. reflexosquamataD. rubrobrunnea, etc.) and are mainly distributed in the Sino-Japanese and Sino-Himalayan regions.11. The Dryopteris clade : This clade contains large portion of species of -Jenkins  and is tDryopteris sect. Lophodium Fraser-Jenk. and D. remota. The species of this clade share short-stalked pinnae, long-aristate ultimate segments, and minutely spinulose perispore sculpturing except D. remota[12. The Lophodium clade : This clade contains species of Dryopteris sect. Fragrantes (H. Itô) Seriz., D. fragrans (L.) Schott. Our work shows that D. fragrans is outside of D. subgen. Dryopteris where it was placed by Fraser-Jenkins [D. fragrans as sister to the rest of Dryopteris, though our statistical support values were low .Our data clearly show that the Dryopsis clade is sister to the Erythrovariae clade; these two together are sister to a clade containing the Acrorumohra clade and the Variae clade; these four clades together are sister to a clade containing the Aemulae clade and the Nothoperanema clade; and these six clades are strongly supported as monophyletic . The relationships among the remaining seven clades are resolved in the ML tree : This clade contains one of the two species of -Jenkins , a resol-Jenkins . Most no-Jenkins ,58 in reAcrophorus, Acrorumohra, Diacalpe, Dryopsis, Nothoperanema, and Peranema, should all be merged into Dryopteris. Most species of these genera share a short rhizome and catadromic arrangement of frond segments, unlike the sister genus of Dryopteris s.l., Arachniodes.The genera, Dryopteris strongly suggests that the current taxonomy of this genus is in need of revision. However, our data do not necessarily falsify the monophyly of these 19 sections. The disagreement between previous taxonomy and molecular results in Dryopteris may be due partly to interspecific hybridization and polyplodization.The non-monophyly of the 19 out of the 21 supraspecific taxa in Dryopteris that are supported by morphological synapomophies and may deserve circumscription as supraspecific entities within Dryopteris.The 13 well-supported clades identified with our data represent 13 major evolutionary lineages in Dryopteris recognized by Fraser-Jenkins [D. sect. Caespitosae S. G. Lu, D. sect. Chrysocomae S. G. Lu, D. sect. Indusiatae S. G. Lu) recognized by Lu [D. sect. Purpurascentes Fraser-Jenk. and the monotypic D. sect. Politae Fraser-Jenk. In total, 78 accessions representing 77 species of Dryopteris s.s. were sampled, including all four subgenera and 19 sections in the current classifications of Dryopteris s.s. by Fraser-Jenkins [All four subgenera and 14 out of 16 sections of ed by Lu ,24,25 aned by Lu , were re-Jenkins .Dryopteris and Acrophorus, Acrorumohra, Diacalpe, Dryopsis, Nothoperanema, and Peranema, further included are eight accessions representing five (63%) out of eight species of Acrophorus, 10 accessions representing three (43%) out of seven species of Acrorumohra, seven accessions representing four (40%) out of 10 species of Diacalpe, six accessions representing five (31%) out of 16 species of Dryopsis, 11 accessions representing six (75%) out of eight species of Nothoperanema, and two accessions representing both species of the bitypic Peranema. Type species of all these six genera, Acrophorus, Acrorumohra, Diacalpe, Dryopsis, Nothoperanema, and Peranema, are included. In total, 122 accessions representing 100 species of the expanded Dryopteris  are included in this study.To assess the phylogenetic relationships between Arachniodes Blume, Leptorumohra H. Itô, and Lithostegia Ching of Dryopteridaceae are used as outgroups based on Liu et al. [ArachniodesLeptorumohraLithostegia, and Phanerophlebiopsis Ching together were resolved as sister to a clade consisting of AcrophorusAcrorumohraDiacalpeDryopsisDryopterisNothoperanema, and Peranema. All sequences used in this study together with their GenBank accession numbers and/or voucher information are listed in Appendix II.One species each of u et al.  where ArrbcL gene, rps4-trnS spacer, trnL intron, and trnL-F spacer. The rbcL gene was amplified with primers F1  and BigDye® Terminator v3.1 Cycle Sequencing Kit .Total genomic DNA was extracted from silica-gel dried material or sometimes from herbarium specimens using Plant Genomic DNA Kits  and DNeasy Plant Mini Kits . The PCR protocols followed Zhang et al.  and Ebihy et al. ) and 137y et al. ). The prTCGAATC; ) and Li TCGAATC; ). The trGGRAACC; ) and theGGRAACC; . AmplifirbcLrps4-trnStrnL, and trnL-F, respectively, were included in our analyses. Some 151 DNA sequences are newly generated for this study (GenBank JX535813-JX535961).Additional sequences were obtained from Genbank and had originally been generated by Geiger & Ranker , Li & LurbcL data was manually obtained using Microsoft Wordpad. Preliminary alignments of rps4-trnS and trnL-F (trnL intron + trnL-F spacer) data were obtained using the default alignment parameters in Clustal X [The alignment of the lustal X  followedtrnL intron and trnL-F spacer were also conducted since these two linked regions are sometimes viewed as one locus.Equally weighted maximum parsimony (MP) tree searches were conducted for each data matrix using 1000 tree-bisection-reconnection (TBR) searches in PAUP* ver. 4.0b10  with MAXtrnL-F spacer), GTR + I (trnL intron), GTR + I + G , HKY + G (rps4-trnS spacer), and SYM + I + G (rbcL gene). The selected models and parameters estimated  with 1000 rapid bootstrap analyses followed by a search for the best-scoring tree in a single run.The simultaneous ML analyses of nucleotide characters and ML bootstrapping (BS) were performed using RAxML-HPC2 on TG ver. 7.2.8  Ching, are monophyletic. Sessa et al. [D. subgen Pycnopteris, for which they had insufficient sampling to test monophyly. In the current study, most of the members of D. subgen. Dryopteris are resolved in the Dryopteris clade, while others are placed in other major clades except the Acrorumohra, Dryopsis, Erythrovariae, Nothoperanema, and Variae clades. Members of D. subgen. Erythrovariae sensu Fraser-Jenkins [, Erythrovariae, and Variae clades, but these clades are paraphyletic in relation to the Dryopsis clade as well as D. chinensis (Baker) Koidz. and D. gymnophylla (Baker) C. Chr. (members of D. sect. Aemulae Fraser-Jenk.), and D. podophylla (Hook.) Kuntze (a member of D. subgen. Pycnopteris). Members of D. subgen. Erythrovariae sensu Wu & Lu [D. subgen. Pycnopteris is consistent with Li & Lu’s [rps4-trnS data. Of our three representatives of this subgenus, two fell in the Dryopteris clade (D. bodinieri (Christ) C. Chr. and D. sieboldii (Van Houtte ex Melt.) Kuntze), and one in the Erythrovariae clade (D. podophylla). Our sole sequence of D. podophylla was derived from Li & Lu [D. subgen. Nephrocystis is not monophyletic because Acrorumohra diffracta Baker (= D. diffracta (Baker) C. Chr.), A. hasseltii Blume (=D. hasseltii (Blume) C. Chr.), A. subreflexipinna (Ogata) Ching (= D. subreflexipinna Ogata), and D. futura A. R. Sm., a member of D. sect. Purpurascentes, are resolved in the Acrorumohra clade and the Cinnamomeae clade. Wu & Lu [D. subgen. Nephrocystis.Our results show that none of the four subgenera, a et al.  also rej-Jenkins  are reso Wu & Lu  are resoi & Lu’s ,20 findi Li & Lu . D. subg Wu & Lu  did not D. sect. Cinnamomeae Fraser-Jenk. and D. sect. Variae Fraser-Jenk., are resolved as monophyletic with our current sampling. This is at odds with Sessa et al. [D. sect. Cinnamomeae and D. sect. Variae, and rejected the monophly of all. Although these two sections are found to be monophyletic in the current study, the sampling for both was larger in Sessa et al. [Our results also demonstrated that 15 out of the 17 sections currently recognized ,24-26, fa et al. , which ta et al. , and theDryopteris sect. Aemulae Fraser-Jenk.: Represented by D. aemula (Aiton) Kuntze (Aemulae clade), D. chinensis, and D. gymnophylla (Acrorumohra clade).Dryopteris sect. Caespitosae S. G. Lu: Represented by D. alpestris Tagawa (Dryopteris clade) and D. fragrans (Fragrantes clade).Dryopteris sect. Chrysocomae S. G. Lu: Represented by D. chrysocoma (Christ) C. Chr. and D. himachalensis Fraser-Jenk. (Dryopteris clade).Dryopteris sect. Dryopteris: This section sensu Fraser-Jenkins [D. alpestrisD. filix-mas (L.) Schott, and D. sichotensis V. Komarov (Dryopteris clade).-Jenkins  is repreDryopteris sect. Fibrillosae Ching: Represented by D. affinis (Lowe) Fraser-Jenk. subsp. borreri Fraser-Jenk., D. polylepis  C. Chr., and D. rosthornii (Diels) C. Chr. (Dryopteris clade).Dryopteris sect. Erythrovariae: This section sensu Fraser-Jenkins [D. caudipinna Nakai, D. championii (Benth.) C. Chr., D. cordipinna Ching & Shing, D. decipiens (Hook.) Kuntze, D. erythrosora (D. Eaton) Kuntze, D. gymnosora (Makino) C. Chr., D. indusiata Makino & Yamam.  Makino following Fraser-Jenkins [D. ryo-itoana Kurata, and D. simasakii (H. Itô) Kurata and all are in the Erythrovariae clade. If D. podophylla (Hook.) Kuntze, a member of D. subgen. Pycnopteris, is included, this section sensu Fraser-Jenkins [-Jenkins  is repre-Jenkins ), D. kin-Jenkins ), D. ryo-Jenkins  becomes Dryopteris sect. Hirtipedes Fraser-Jenk.: Represented by D. atrata  Ching, D. commixta Tagawa, D. conjugata Ching, D. cycadina (Franchet & P. A. L. Savat.) C. Chr., D. dickinsii (Franchet & P. A. L. Savat.) C. Chr., D. handeliana C. Chr., D. hangchowensis Ching, D. lunanensis (Christ) C. Chr., D. scottii (Bedd.) Ching, D. stenolepis (Baker) C. Chr. and D. tsutsuiana Kurata, all of which are in the Dryopteris clade. This section becomes monophyletic if D. rosthornii (D. sect. Fibrillosae) and D. uniformis  are included.Dryopteris sect. Indusiatae S. G. Lu: Represented by D. gymnosora (Makino) C. Chr. and D. indusiata Makino & Yamam. (Erythrovariae clade).Dryopteris sect. Lophodium (Newman) C. Chr. ex H. Itô: Represented by D. amurensis Christ, D. expansa (C. Presl) Fraser-Jenk. & Jermy, D. intermedia (Muhlenb. ex Willd.) A. Gray, and D. dilatata (Hoffm.) A. Gray (Lophodium clade). These four species are paraphyletic in relation to D. remota (A. Braun ex Doell) Druce, the type of D. sect. Remotae. The close relationship between D. remota and species of D. sect. Lophodium based on our plastid data shows that D. remota, a triploid, is possibly originated through hybridization with one of the species in D. sect. Lophodium being the maternal donor.Dryopteris sect. Marginatae Fraser-Jenk.: Represented by D. aquilinoides (Desv.) C. Chr. and D. shiroumensis Kurata & Nakaike (Dryopteris clade). They are resolved as paraphyletic in relation to the rest of the Dryopteris clade.Dryopteris sect. Nephrocystis: Represented by Acrorumohra diffractaA. hasseltiiA. subreflexipinnaDryopteris hayatae Tagawa  C. Chr. by Fraser-Jenkins [D. melanocarpa Hayata (subsumed in D. platypus (Kunze) Kuntze by Fraser-Jenkins [D. maximowicziana (Miq.) C. Chr. (not recognized by Fraser-Jenkins [D. sabae (Franchet & P. A. L. Savat.) C. Chr., D. sparsa (Buch.-Ham. ex D. Don) Kuntze, and D. yakusilvicola Sa. Kurata (subsumed in D. cacaiana Tagawa by Fraser-Jenkins [-Jenkins ), D. mel-Jenkins ), D. max-Jenkins ), D. sab-Jenkins ). The fiDryopteris sect. Pallidae Fraser-Jenk.: Represented by D. juxtaposita Christ, D. lacera (Thunb.) Kuntze, D. sublacera Christ, D. uniformis (Makino) Makino (Dryopteris clade), D. odontoloma (Bedd.) C. Chr., and D. pallida (Bory) C. Chr. ex Maire & Petitm. .Dryopteris sect. Pandae Fraser-Jenk.: Represented by D. himachalensis Fraser-Jenk. (Acrorumohra clade) and D. tokyoensis (Matsum. & Makino) C. Chr. .Dryopteris sect. Remotae Fraser-Jenk.: Represented by D. corleyi Fraser-Jenk. (Aemulae clade) and D. remota (Lophodium clade).Dryopteris sect. Splendentes Fraser-Jenk.: Represented by D. reflexosquamata Hayata and D. rubrobrunnea W. M. Chu (Dryopteris clade). These two are paraphyletic in relation to three members of D. sect. Pallidae and two member of D. subgen. Pycnopteris.Acrophorus emeiensis Ching: rbcL zl1474, trnL JX535916, trnL-F JX535867, rps4-trnS JX535815. Acrophorus exstipellatus Ching & S. H. Wu: rbcL JX535857, trnL JX535914, trnL-F JX535865, rps4-trnS JX535813. Acrophorus macrocarpus Ching & S. H. Wu: rbcL DQ054522 (“Acrophorus emeiensis Ching”). Acrophorus nodosus C. Presl: rbcL AB575065, trnL JX535915, trnL-F JX535866, rps4-trnS JX535814. Acrophorus paleolatus Pic. Serm. (“Acrophorus stipellatus T. Moore”): rbcL DQ054510 DQ508756 EF463106, trnL-F DQ514500 EF540696 DQ480130.Acrorumohra diffracta (Baker) H. Itô: rbcL DQ508758 EF463108, trnL-F EU797681 EU797682 EU797683, rps4-trnS EU797685 EU797686 EU797687. Acrorumohra hasseltii (Blume) Ching: rbcL AB575136 DQ054519 DQ508757 EF463107, trnL-F DQ514479 EU797677 EU797679 EU797680, rps4-trnS DQ191888 EU797691 EU797692 EU797693. Acrorumohra subreflexipinna (Ogata) H. Itô: trnL -F EU797675 EU797676 EU797678, rps4-trnS EU797688 EU797689 EU797690.Arachniodes aristata (G. Forst.) Tindale: rbcL AY268851, trnL-F AY268782. Arachniodes assamica (Kuhn) Ohwi: rps4-trnS DQ191891.Diacalpe annamensis Tagawa: rbcL EF463125, trnL-F DQ480132 EF540698. Diacalpe aspidioides Blume: rbcL DQ054523 EF463126, trnL-F DQ514490. Diacalpe chinensis Ching & S. H. Wu: rbcL JX535864, trnL JX535956, trnL-F JX535908, rps4-trnS JX535854. Diacalpe christensenae Ching: rbcL DQ054518 EF540699, trnL-F DQ480131 EF540699, rps4-trnS DQ480131 EF540699.Dryopsis apiciflora  Holttum & P.J. Edwards: rbcL DQ054521, trnL JX535957, trnL-F JX535909. Dryopsis clarkei (Baker) Holttum & P.J. Edwards: trnL JX535958, trnL-F JX535910, rps4-trnS JX535855. Dryopsis heterolaena (C. Chr.) Holttum & P.J. Edwards: rbcL DQ508770, trnL-F DQ514492. Dryopsis mariformis (Rosenst.) Holttum & P.J. Edwards: rbcL DQ054520 EF460683, trnL JX535959, trnL-F JX535911. Dryopsis sp.: rbcL DQ054525.Dryopteris aemula (Aiton) Kuntze: rbcL AY268881, trnL-F AY268816, rps4-trnS JN189189. Dryopteris affinis (Lowe) Fraser-Jenk. subsp. borreri Fraser-Jenk.: rbcL AY268849, trnL-F AY268776, rps4-trnS JN189190. Dryopteris alpestris Tagawa: rbcL JX535858, trnL JX535917, trnL-F JX535868, rps4-trnS JXH11103. Dryopteris amurensis Christ: rbcL AB575112, trnL JX535918, trnL-F JX535869, rps4-trnS JX535816. Dryopteris aquilinoides (Desv.) C. Chr.: rbcL AY268868, trnL-F AY268803, rps4-trnS JN189211. Dryopteris atrata  Ching: rbcL AB575115, trnL JX535919, trnL-F JX535870, rps4-trnS JX535817. Dryopteris bissetiana (Baker) C. Chr.: rbcL AY268862, trnL-F AY268796, rps4-trnS DQ191829. Dryopteris bodinieri (Christ) C. Chr.: rbcL DQ508772, trnL-F DQ514494, rps4-trnS DQ191830. Dryopteris caudipinna Nakai: rbcL AB575117, trnL JX535920, trnL-F JX535871, rps4-trnS JX535818. Dryopteris championii (Benth.) C. Chr.: rbcL AY268863, trnL-F AY268797, rps4-trnS DQ151856. Dryopteris chinensis (Baker) Koidz.: rbcL JX535859, trnL JX535921, trnL-F JX535872, rps4-trnS JX535819. Dryopteris chrysocoma (Christ) C. Chr.: rbcL DQ508773, trnL-F DQ514495, rps4-trnS DQ191832. Dryopteris cinnamomea (Cav.) C. Chr.: rbcL JN189528, trnL-F FR731991, rps4-trnS JN189202. Dryopteris commixta Tagawa: rbcL AB575120, trnL JX535922, trnL-F JX535873, rps4-trnS JX535820. Dryopteris corleyi Fraser-Jenk.: rbcL AY268873, trnL-F AY268808. Dryopteris crinalis (Hook. &Arn.) C. Chr.: AY268835, trnL-F AY268774. Dryopteris cycadina (Franchet & P. A. L. Savat.) C. Chr.: rbcL EF463127, trnL-F AY278400, rps4-trnS DQ191835. Dryopteris decipiens (Hook.) Kuntze var. decipiens: rbcL AB575123, trnL JX535923, trnL-F JX535874, rps4-trnS JX535821. Dryopteris dickinsii (Franchet & P. A. L. Savat.) C. Chr.: rbcL AB575125, trnL JX535924, trnL-F JX535875, rps4-trnS JX535822. Dryopteris dilatata (Hoffm.) A. Gray: rbcL AY268848, trnL-F AY268779, rps4-trnS JN189248. Dryopteris erythrosora (D. Eaton) Kuntze: rbcL DQ508774, trnL-F DQ514496, rps4-trnS JN189255. Dryopteris expansa (C. Presl) Fraser-Jenk. & Jermy: rbcL AY268844, trnL-F AY268775, rps4-trnS JN189180. Dryopteris filix-mas (L.) Schott: rbcL AY268845, trnL-F AY268776, rps4-trnS JN189181. Dryopteris fragrans (L.) Schott: rbcL AB575129, AY268865, trnL-F FR731981 AY268800, rps4-trnS JN189185. Dryopteris futura A. R. Sm.: rbcL JN189534, trnL-F JN189103, rps4-trnS JN189208. Dryopteris gymnophylla (Baker) C. Chr.: rbcL JX535860, trnL JX535925, trnL-F JX535876, rps4-trnS JX535823. Dryopteris gymnosora (Makino) C. Chr.: rbcL AB575132, trnL JX535926, trnL-F JX535877, rps4-trnS JX535824. Dryopteris hadanoi Kurata: rbcL AB575133, trnL JX535927, trnL-F JX535878, rps4-trnS JX535825. Dryopteris handeliana C. Chr.: rbcL AB575134, trnL JX535928, trnL-F JX535879, rps4-trnS JX535826. Dryopteris hangchowensis Ching: rbcL AB575135, trnL JX535929, trnL-F JX535880, rps4-trnS JX535827. Dryopteris hayatae Tagawa: rbcL AB575137, trnL JX535930, trnL-F JX535881, rps4-trnS JX535828. Dryopteris himachalensis Fraser-Jenk.: rps4-trnS DQ191845. Dryopteris indusiata Makino & Yamam.: rbcL AB575140, trnL JX535931, trnL-F JX535882, rps4-trnS JX535829. Dryopteris intermedia (Muhlenb. ex Willd.) A. Gray subsp. maderensis (J. Milde ex Alston) Fraser-Jenkins: rbcL AB575143, trnL-F FR731985. Dryopteris juxtaposita Christ: rbcL AY268875, trnL-F AY268810, rps4-trnS DQ191848. Dryopteris kinkiensis Koidz. ex Tagawa: rbcL AB575144, trnL JX535932, trnL-F JX535883, rps4-trnS JX535830. Dryopteris lacera (Thunb.) Kuntze: rbcL AB575148, trnL JX535933, trnL-F JX535884, rps4-trnS JX535831. Dryopteris laeta (Kom.) C. Chr.: rbcL AB575149, trnL JX535934, trnL-F JX535885, rps4-trnS JX535832. Dryopteris lunanensis (Christ) C. Chr.: rbcL AB575150, trnL JX535935, trnL-F JX535886, rps4-trnS JX535833. Dryopteris mauiensis C. Chr.: rbcL AY268833, trnL-F AY268770. Dryopteris maximowicziana: rbcL AB575151, trnL JX535936, trnL-F JX535887, rps4-trnS JX535834. Dryopteris melanocarpa Hayata: rbcL AB575153, trnL JX535937, trnL-F JX535888, rps4-trnS JX535835. Dryopteris monticola (Makino) C. Chr.: rbcL AB575154, trnL JX535938, trnL-F JX535889, rps4-trnS JX535836. Dryopteris nipponensis Koidz.:AB575156, trnL JX535939, trnL-F JX535890, rps4-trnS JX535837. Dryopteris odontoloma (Bedd.) C. Chr.: rbcL AY268872, trnL-F AY268807, rps4-trnS DQ191859. Dryopteris pacifica (Nakai) Tagawa: rbcL AB575157, trnL JX535940, trnL-F JX535891, rps4-trnS JX535838. Dryopteris pallida (Bory) C. Chr. ex Maire & Petitm.: rbcL AY268874, trnL-F AY268809, rps4-trnS JN189266. Dryopteris patula (Sw.) L. Underw.: rbcL JN189500, trnL-F AY268823, rps4-trnS JN189176. Dryopteris podophylla (Hook.) Kuntze: rps4-trnS DQ191864. Dryopteris polita Rosenst.: rbcL AB575158, trnL-F EU797684, rps4-trnS EU797694. Dryopteris polylepis  C. Chr.: rbcL AY268864, trnL-F AY268798, rps4-trnS JN189263. Dryopteris reflexosquamata Hayata: rbcL JN189604, trnL-F JN189171, rps4-trnS DQ191870. Dryopteris remota (A. Braun ex Doell) Druce: rbcL AY268858, trnL-F AY268792, rps4-trnS JN189204. Dryopteris rosthornii (Diels) C. Chr.: rbcL JX535861, trnL JX535941, trnL-F JX535892, rps4-trnS JX535839. Dryopteris rubrobrunnea W. M. Chu: rbcL JX535862, trnL JX535942, trnL-F JX535893, rps4-trnS JX535840. Dryopteris ryo-itoana Kurata: rbcL AB575161, trnL JX535943, trnL-F JX535894, rps4-trnS JX535841. Dryopteris sabae (Franchet & P. A. L. Savat.) C. Chr.: rbcL AB575162, trnL JX535944, trnL-F JX535895, rps4-trnS JX535842. Dryopteris sacrosancta Koidz.: rbcL AB575163, trnL JX535945, trnL-F JX535896, rps4-trnS JX535843. Dryopteris sandwiciensis (Hook. & Arn.) C. Chr.: rbcL AY268827, trnL-F AY268762. Dryopteris saxifraga H. Itô: rbcL AB575164, trnL JX535946, trnL-F JX535897, rps4-trnS JX535844. Dryopteris scottii (Bedd.) Ching: rbcL JX535863, trnL-F JX535898, rps4-trnS DQ191872. Dryopteris shiroumensis Kurata & Nakaike: rbcL AB575168, trnL JX535947, trnL-F JX535899, rps4-trnS JX535845. Dryopteris sichotensis V. Komarov: rbcL AY268869, trnL-F AY268804. Dryopteris sieboldii (Van Houtte ex Melt.) Kuntze: rbcL AB575169, trnL JX535948, trnL-F JX535900, rps4-trnS JX535846. Dryopteris simasakii (H. Itô) Kurata var. simasakii: rbcL AB575170, trnL JX535949, trnL-F JX535901, rps4-trnS JX535847. Dryopteris sordidipes Tagawa: rbcL AB575172, trnL JX535950, trnL-F JX535902, rps4-trnS JX535848. Dryopteris sparsa (Buch.-Ham. ex D. Don) Kuntze: rbcL AB575173, trnL JX535951, trnL-F JX535903, rps4-trnS JX535849. Dryopteris stenolepis (Baker) C. Chr.: rbcL AY268889, trnL-F AY268824, rps4-trnS DQ191877. Dryopteris sublacera Christ: rbcL DQ508778, trnL-F DQ514501, rps4-trnS DQ191878. Dryopteris tetrapinnata W. H. Wagner & Hobdy: rbcL AY268838, trnL-F AY268772. Dryopteris tokyoensis (Matsum. & Makino) C. Chr.: rbcL AY268861, trnL-F AY268795, rps4-trnS JN189251. Dryopteris tsutsuiana Kurata: rbcL AB575176, trnL JX535952, trnL-F JX535904, rps4-trnS JX535850. Dryopteris unidentata (Hook. & Arn.) C. Chr. var. unidentata: rbcL AY268825, trnL-F AY268766. Dryopteris uniformis (Makino) Makino: rbcL AB575177, trnL JX535953, trnL-F JX535905, rps4-trnS JX535851. Dryopteris varia (L.) Kuntze: rbcL AB575178, trnL JX535954, trnL-F JX535906, rps4-trnS JX535852. Dryopteris xanthomelas (Christ) C. Chr.: rbcL AY587118, trnL-F DQ150394, rps4-trnS DQ151857. Dryopteris yakusilvicola Sa. Kurata: rbcL AB575180, trnL JX535955, trnL-F JX535907, rps4-trnS JX535853.Leptorumohra quadripinnata (Hayata) H. Itô: rbcL DQ508781, trnL-F DQ514505, rps4-trnS EF540707.Lithostegia foeniculacea (Hook.) Ching: rbcL DQ508782, trnL-F DQ514506, rps4-trnS EF540717.Nothoperanema diacalpioides Ching: rbcL DQ054511. Nothoperanema hendersonii (Bedd.) Ching: rbcL AB575138 DQ508783 EF463135 JN189547, trnL-F DQ514507 JN189116, rps4-trnS DQ191885 JN189221. Nothoperanema rubiginosum (Brack.) A. R. Sm. & D. R. Palmer: rbcL AY268836 DQ054511 (“Nothoperanema hendersonii (Bedd.) Ching”) EF463182 (“Nothoperanema squamisetum (Hook.) Ching”), trnL-F AY268771. Nothoperanema shikokianum (Makino) Ching: rbcL AB575167 DQ054509 EF463136, trnL JX535960, trnL-F JX535912, rps4-trnS DQ191886 JX535856. Nothoperanema squamisetum (Hook.) Ching: rbcL DQ054512, trnL JX535961, trnL-F JX535913, rps4-trnS DQ191887.Peranema cyatheoides D. Don: rbcL DQ054513. Peranema luzonicum Copel.: rbcL DQ508784 (“Peranema cyatheoides D. Don”), trnL-F DQ514509 (“Peranema cyatheoides D. Don”).The author(s) declare that they have no competing interests.LBZ desgined the study, conducted data analyses, and wrote the manuscript, LZ, EBS, and AE carried out the lab work, LZ, SYD, EBS, XFG, and AE collected and identified portion of the material. All authors contributed to the manuscript revision. All authors read and approved the final manuscript.


The word "Neuroacanthocytosis" is misspelled in the article title. The correct title is: Alterations of Red Cell Membrane Properties in Neuroacanthocytosis. The correct citation is: Siegl C, Hamminger P, Jank H, Ahting U, Bader B, et al. (2013) Alterations of Red Cell Membrane Properties in Neuroacanthocytosis. PLoS ONE 8(10): e76715. doi:10.1371/journal.pone.0076715


The 4th author's last name is misspelled. The correct author name is: Odile Boespflug-TanguyThe correct citation is:Schiff M, Benoist J-F, Aïssaoui S, Boespflug-Tanguy O, Mouren M-C, et al. (2011) Should Metabolic Diseases Be Systematically Screened in Nonsyndromic Autism Spectrum Disorders? PLoS ONE 6(7): e21932. doi:10.1371/journal.pone.0021932


The authors wish to add Gabriele Campi as the 5th author to the byline. Dr. Campi's affiliation is: Experimental Immunology Unit, Division of Immunology, Transplantation and Infectious Diseases (DIBIT), San Raffaele Scientific Institute, Milano, Italy. The correct byline is: Canderan G, Gruarin P, Montagna D, Fontana R, Campi G, Melloni G, Traversari C, Dellabona P, Casorati G. The correct citation is: Canderan G, Gruarin P, Montagna D, Fontana R, Campi G, et al. (2010) An Efficient Strategy to Induce and Maintain In Vitro Human T Cells Specific for Autologous Non-Small Cell Lung Carcinoma. PLoS ONE 5(8): e12014. doi:10.1371/journal.pone.0012014


The second author's name was spelled incorrectly. The correct name is: Holger Brüggemann.Clostridium botulinum Type A Strain Hall. PLoS ONE 7(7): e41848. doi:10.1371/journal.pone.0041848The correct citation is: Connan C, Brüggemann H, Mazuet C, Raffestin S, Cayet N, et al. (2012) Two-Component Systems Are Involved in the Regulation of Botulinum Neurotoxin Synthesis in


PLoS ONE 7(4): e35493. doi:10.1371/journal.pone.0035493 There was a typographical error in the second author's name. The correct spelling is Markus Damme. The correct citation is: Thelen M, Damme M, Schweizer M, Hagel C, Wong AM, et al. (2012) Disruption of the Autophagy-Lysosome Pathway Is Involved in Neuropathology of the


The name of the fifth author is incorrect. The correct name is: Yun-Wen Zheng. The correct Citation is: Liu NM, Yokota T, Maekawa S, Lü P, Zheng Y-W, et al. (2013) Transcription Profiles of Endothelial Cells in the Rat Ductus Arteriosus during a Perinatal Period. PLoS ONE 8(9): e73685. doi:10.1371/journal.pone.0073685. The correct abbreviation of the fifth author's name in the Author Contributions Statement is: YWZ.


The third author's name was spelled incorrectly. The correct name is: Firas Kobeissy.The correct citation is: Xu G, Stevens SM Jr, Kobeissy F, Brown H, McClung S, et al. (2012) Identification of Proteins Sensitive to Thermal Stress in Human Neuroblastoma and Glioma Cell Lines. PLoS ONE 7(11): e49021. doi:10.1371/journal.pone.0049021


There was an error in the title and citation. The correct title is, "Absence of Colony Stimulating Factor-1 Receptor Results in Loss of Microglia, Disrupted Brain Development and Olfactory Deficits." The correct citation is: Erblich B, Zhu L, Etgen AM, Dobrenis K, Pollard JW (2011) Absence of Colony Stimulating Factor-1 Receptor Results in Loss of Microglia, Disrupted Brain Development and Olfactory Deficits. PLoS ONE 6(10): e26317. doi:10.1371/journal.pone.0026317


FUBP1, CIC, and IDH1. PLoS ONE 8(3): e59773. doi:10.1371/journal.pone.0059773. The name of the fifth author was given incorrectly in the citation. The correct citation is: Klink B, Miletic H, Stieber D, Huszthy PC, Campos Valenzuela JA, et al. (2013) A Novel, Diffusely Infiltrative Xenograft Model of Human Anaplastic Oligodendroglioma with Mutations in


There was a typographical error in the title and citation. The correct title is, "Fragile X Mental Retardation Protein Interacts with the RNA-Binding Protein Caprin1 in Neuronal RiboNucleoProtein Complexes" and the correct citation is:El Fatimy R, Tremblay S, Dury AY, Solomon S, De Koninck P, et al. (2012) Fragile X Mental Retardation Protein Interacts with the RNA-Binding Protein Caprin1 in Neuronal RiboNucleoProtein Complexes. PLoS ONE 7(6): e39338. doi:10.1371/journal.pone.0039338


There was an error in the name of the third author in the citation. The correct citation is:Wavre-Shapton ST, Tolmachova T, Lopes da Silva M, Futter CE, Seabra MC (2013) Conditional Ablation of the Choroideremia Gene Causes Age-Related Changes in Mouse Retinal Pigment Epithelium. PLoS ONE 8(2): e57769. doi:10.1371/journal.pone.0057769


There was an error in the name of the second author.The correct name is: Homayoun Sadeghi-BazarganiThe correct citation is: Arshi S, Sadeghi-Bazargani H, Mohammadi R (2012) Burn Injury-Specific Home Safety Assessment: A Cross-Sectional Study in Iran. PLoS ONE 7(11): e49412. doi:10.1371/journal.pone.0049412


The first author's name was spelled incorrectly. The correct name is: Dharmayati Bambang Utoyo. The correct abbreviation in the Contributions section is DBU. The correct citation is: Utoyo DB, Jaya ES, Arjadi R, Hanum L, Astri K, et al. (2013) Preliminary Study on the Effectiveness of Short Group Cognitive Behavioral Therapy (GCBT) on Indonesian Older Adults. PLoS ONE 8(2): e57198. doi:10.1371/journal.pone.0057198. There were additional errors in the Contributions section. The correct contributions are: Conceived and designed the experiments: DUL ESJ RA LH KA MDDP. Performed the experiments: ESJ RA LH KA MDDP. Analyzed the data: DUL ESJ RA. Contributed reagents/materials/analysis tools: DUL ESJ RA LH KA MDDP. Wrote the paper: DUL ESJ RA.


The name of the first author was incorrectly represented in the Citation. The correct Citation is: de Gouw D, Hermans PWM, Bootsma HJ, Zomer A, Heuvelman K, et al. (2014) Differentially Expressed Genes in Bordetella pertussis Strains Belonging to a Lineage Which Recently Spread Globally. PLoS ONE 9(1): e84523. doi:10.1371/journal.pone.0084523.


There was an error in the same of the fourth author.The correct name is: Stefan HabelitzThe correct citation is: Burwell AK, Thula-Mata T, Gower LB, Habelitz S, Kurylo M, et al. (2012) Functional Remineralization of Dentin Lesions Using Polymer-Induced Liquid-Precursor Process. PLoS ONE 7(6): e38852. doi:10.1371/journal.pone.0038852


The name of the third author is incorrect. The correct name is: Debi Roberson. The correct Citation is: Zhang R, Hu Z, Roberson D, Zhang L, Li H, et al. (2013) Neural Processes Underlying the“Same”-“Different” Judgment of Two Simultaneously Presented Objects- An EEG Study. PLoS ONE 8(12): e81737. doi:10.1371/journal.pone.0081737. The correct abbreviation of the third author's name in the Author Contributions statement is: DR.


Three references were omitted from the PDF during the preparation of this article for publication.These references are:53. Vanderschuren LJMJ, Trezza V, Griffioen-Roose S, Schiepers OJG, Van Leeuwen N, et al. (2008) Methylphenidate disrupts social play behavior in adolescent rats. Neuropsychopharmacology 33: 2946–2956. doi: 10.1038/npp.2008.10.54. Sara SJ, Dyon-Laurent C, Hervé A (1995) Novelty seeking behavior in the rat is dependent upon the integrity of the noradrenergic system. Cogn Brain Res 2: 181–187.55. Tzschentke TM (2007) Measuring reward with the conditioned place preference paradigm: update of the last decade. Addict Biol 12: 227–462.


There was an error in the author byline. Severine Jancek was excluded. The correct byline is: Gwenael Piganeau, Adam Eyre-Walker, Severine Jancek, Nigel Grimsley, Herve Moreau.Severine Jancek's affiliation is 4 Institut de Recherche sur la Biologie de l'Insecte, UMR 6035, CNRS, Université François Rabelais, 37200 Tours, France.The correct citation is: Piganeau G, Eyre-Walker A, Jancek S, Grimsley N, et al. (2011) How and Why DNA Barcodes Underestimate the Diversity of Microbial Eukaryotes. PLoS ONE 6(2): e16342. doi:10.1371/journal.pone.0016342Severine Jancek's author contribution is: Analyzed the data.


Journal of Neuroinflammation 8:47.Correction to Gupta P K, Prabhakar S, Sharma S, Anand A. Vascular endothelial growth factor-A (VEGF-A) and chemokine ligand-2 (CCL2) in amyotrophic lateral sclerosis (ALS) patients.  The authors observe that, in Table two of our study , crude OSmoking: .Alcohol consumption: .Meat consumption: .


The fourth author's name is misspelled. The correct spelling is: Omid Saeed Tehrani. The correct citation is: Pitter KL, Galbán CJ, Galbán S, Tehrani OS, Li F, et al. (2011) Perifosine and CCI 779 Co-Operate to Induce Cell Death and Decrease Proliferation in PTEN-Intact and PTEN-Deficient PDGF-Driven Murine Glioblastoma. PLoS ONE 6(1): e14545. doi:10.1371/journal.pone.0014545


There is an error in reference 33. The correct reference is: Patowary S, Alvarez‑Curto E, Xu TR, Holz JD, Oliver JA, Milligan G, Raicu V. (2013). The muscarinic M3 acetylcholine receptor exists as two differently sized complexes at the plasma membrane. Biochem J 452:303-12.


The second author's name is misspelled. The correct spelling is: Heike MikschofskyThe correct citation is:Nausch H, Mikschofsky H, Koslowski R, Meyer U, Broer I, et al. (2012) Expression and Subcellular Targeting of Human Complement Factor C5a in Nicotiana species. PLoS ONE 7(12): e53023. doi:10.1371/journal.pone.0053023


An initial is missing in the third author's name. The correct name is: Balthasar A Heesters. The correct citation is: Langereis MA, Zeng Q, Heesters BA, Huizinga EG, de Groot RJ , et al. (2012) The Murine Coronavirus Hemagglutinin-esterase Receptor-binding Site: A Major Shift in Ligand Specificity through Modest Changes in Architecture. PLoS Pathog 8(1): e1002492. doi:10.1371/journal.ppat.1002492


Overstreetia cribbi n. sp. from Atherinomorus lacunosus. This species differs from its congeners in the detail of its circum-oral spination and some metrical features. Other new records are of: Diphterostomum plectorhynchi Machida, Kamegai & Kuramochi, 2006 in Diagramma pictum; Parvipyrum acanthuri (Acanthurus dussumieri; Zoogonoides viviparus (Lagocephalus sceleratus; Deretrema ? combesorum (Parupeneus pleurostigma; D? acutum (P. barberinus; and an unidentified immature zoogonid in P. multifasciatus. The newly reported specimens are illustrated and measurements given. The distribution of New Caledonian zoogonids is listed.New and published reports of zoogonid digeneans from New Caledonian waters are recorded, including a description of canthuri  in Acantiviparus  in Lagocmbesorum  early ov? acutum  in P. ba Deretrema species sister to the Zoogoninae + Faustulidae, and Lepidophyllum species sister to that assemblage. As can be seen the sample size is small and, although the support for this arrangement is statistically good, the findings are clearly provisional and preliminary. The known zoogonid fauna of waters around New Caledonia is small, but will probably be found to be larger when explorations of the deep-sea are undertaken. In this paper we list all the known species, including a new species of the genus Overstreetia  will represent a published work according to the International Commission on Zoological Nomenclature (ICZN), and hence the new names contained in the electronic version are effectively published under that Code from the electronic edition alone. This published work and the nomenclatural acts it contains have been registered in ZooBank, the online registration system for the ICZN. The ZooBank LSIDs (Life Science Identifiers) can be resolved and the associated information viewed through any standard web browser by appending the LSID to the prefix “Family Zoogonidae Odhner, 1902Subfamily Zoogoninae Odhner, 1902GenusDiphterostomumStossich, 1903urn:lsid:zoobank.org:act:D56035F2-141C-4607-8AFE-7AEFED0AE0B3.ZooBank: Diphterostomum tropicumDurio & Manter, 1968urn:lsid:zoobank.org:act:71827EC5-F9EA-44A4-888D-12BF3E1FE770.ZooBank: Diphtherostomum tropicum Durio & Manter, 1968Syn: Record from off New Caledonia: Durio & Manter (1968a)Lethrinus sp., ‘bec de cane’; ‘can be safely identified as’ Lethrinus nebulosus   .Lethrinus lentjan   from Green Island, on the Great Barrier Reef. The only other report is from the sparid Chrysophrys auratus  from off New Zealand by Discussion: Diphterostomum plectorhynchiMachida, Kamegai & Kuramochi, 2006 , Perciformes, Haemulidae, painted sweetlips.Host: Site: digestive tract.Locality: Between Larégnière and Récif Crouy ; Interior Lagoon near Récif Toombo .Specimens: MNHN JNC2511, JNC2512, JNC3023, BMNH 2014.1.31.2-3.Previous New Caledonian records: none.Plectorhinchus spp. from off Japan   (as P. hasta), the banded grunter P. furcatus  (as Rhonciscus furcatus) and the saddle grunt P. maculatus  from the Bay of Bengal  . D. indif Bengal , is saidDiphterostomum spp.  with, according to our records, over 200 reports of various life-cycle stages. More than 130 of these (66%) refer to the species D. brusinae. About 5% of these records are from the Indo-West Pacific Region, but no reports are from haemulids. Members of this genus are all similar with few distinguishing features.There are about 13 recognised GenusParvipyrumPritchard, 1963urn:lsid:zoobank.org:act:7C176519-62E1-401E-BDDE-F19F8B2CBA7D.ZooBank: Parvipyrum acanthuriPritchard, 1963 ; ex A. dussumieri, Nouméa Fish Market (08/04/2011), Récif Snark .Localities: ex A. blochii, MNHN JNC2213, BMNH 2007.11.14.51; ex A. dussumieri, MNHN JNC2545, JNC3374, BMNH 2014.1.31.6.Specimens: ex Previous New Caledonian record: Acanthurus blochii.Previously reported New Caledonian host: Acanthurus, and has been reported only from Hawaii  Odhner, 1902 , Tetraodontiformes, Tetraodontidae, silver-cheeked toadfish.Host: Site: digestive tract.Locality: near Îlot Pandanus .Specimens: MNHN JNC2982.Previous New Caledonian records: none.Zoogonoides have been described from the Indo-Pacific region. Three of these can be distinguished from our specimen by the sucker ratio, with the ventral sucker smaller than, or of similar size to, the oral sucker , and Z. yamagutii Kamegai, 1973. Z. viviparus is reported mainly in the North Atlantic Ocean, having been originally reported off Norway  [as P. anguillaris] (Siluriformes: Plotosidae) from Nishidomari Bay, Tsushima Island, Japan , the second in intermediate host may be an ophiuroid or holothurian echinoderm, a polychaete, bivalve or gastropod, or possibly a mysid crustacean , Perciformes, Lethrinidae, Japanese large-eye bream; Lethrinus atkinsoni Seale, 1910, Perciformes, Lethrinidae, Pacific yellowtail emperor; L. genivittatus Valenciennes, 1830, Perciformes, Lethrinidae, longspine emperor:Hosts: Site: intestine, digestive tract.G. euanus, Inside Lagoon, facing Récif Toombo , Off Récif Kué ; ex L. atkinsoni, Off Ever Prosperity ; ex L. genivittatus, Off Baie des Citrons, Nouméa , Baie Maa .Localities: ex G. euanus, MNHN JNC2388, BMNH 2014.1.31.4; ex L. atkinsoni, MNHN JNC1789, BMNH 2007.11.14.52; ex L. genivittatus, MNHN JNC2293, BMNH 2007.11.14.52.Specimens: ex Previous New Caledonian records: 1. G. euanus , L. atkinsoni , L. genivittatus .Previously reported New Caledonian host: Chrysophrys auratus Forster 1801 (as Pagrosomus unicolor) from the Inland Sea of Japan  from off Callao, Peru , Perciformes, Mullidae, manybar goatfish.New Caledonian host: Deretrema combesorumBray & Justine, 2008urn:lsid:zoobank.org:act:CE93F29C-4B9E-4E39-B986-BBB772DA8016.ZooBank: Record from off New Caledonia: Parupeneus multifasciatus (Quoy & Gaimard 1825), Perciformes, Mullidae, manybar goatfish.New Caledonian host: Deretrema ? combesorumBray & Justine, 2008, early ovigerous forms , Perciformes, Mullidae, sidespot goatfish.Host: Site: digestive tractLocality: West of Passe de Dumbéa .Specimens: MNHN JNC2416, BMNH 2014.1.31.5.C. combesorum. They show all the diagnostic characters listed in D. combesae and C. combesorum. Nevertheless, several metric and ratio characters differ, e.g., size, width ratio, and the ratios of the suckers and the sizes of most gonads relative to body-length , Perciformes, Mullidae, dash-and-dot goatfish.Host: Site: digestive tract.Locality: West of Îlot Goëland .Specimens: MNHN JNC2346.Previous New Caledonian records: none.D. acutum as described by Naso, from off Hawaii  synonymous with D. acutum, following the discussion of Discussion: The single specimen available appears indistinguishable  from D. f Hawaii . Bray , Perciformes, Priacanthidae, moontail bullseye.New Caledonian host: GenusLecithostaphylusOdhner, 1911urn:lsid:zoobank.org:act:431229B4-A8F9-4107-B3A4-779055CD30F4.ZooBank: Lecithostaphylus nitens  Linton, 1940urn:lsid:zoobank.org:act:EDDA9CF7-19AA-4386-AAE2-BEBF6ABC566D.ZooBank: Record from off New Caledonia: Tylosurus crocodilus , Beloniformes, Belonidae, hound needlefish.New Caledonian host: Tylosurus, Platybelone and Ablennes, mostly from the former. It was originally reported in ‘Tylosurus caribbaeus’ [? T. acus] from off Woods Hole, Massachusetts in the Northwestern Atlantic Ocean. Most reports are from the northwestern Atlantic or Gulf of Mexico, including one from T. crocodilus , Atheriniformes, Atherinidae, hardyhead silverside.Host: Site: digestive tract.Locality: Anse Vata, Nouméa .Specimens: Holotype MNHN JNC2656, Paratype BMNH 2014.1.31.1.Etymology: This species is named after our colleague Dr. Tom Cribb, of the University of Queensland, who has contributed immeasurably to our understanding of the taxonomy and biology of digeneans.Based on 2 ovigerous specimens. Measurements on Testes 2, oval, entire to slightly irregular, tandem, contiguous, in anterior half of hindbody. Cirrus-sac broadly claviform, posterior end overlaps anterior edge of ventral sucker. Seminal vesicle allantoid, undivided, surrounded by gland-cells, in proximal region of cirrus-sac. Pars prostatica vesicular. Ejaculatory duct short, thick-walled. Genital atrium small. Genital pore submarginal, sinistral, bifurcal.Ovary oval, entire, contiguous with ventral sucker and close to anterior testis. Proximal female system obscured by eggs. Uterus runs posteriorly from ovary passes ventrally over testes, fills bulk of body posterior to anterior testis, presumably reaches extracaecally, but caeca obscured by eggs. Metraterm short, muscular, with narrow sheath of gland-cells. Eggs numerous, operculate, tanned. Vitellarium forms 2 lateral fields of few  irregularly oval follicles between levels of ventral sucker and posterior testis.Excretory pore terminal. Excretory vesicle anterior extent and shape obscured by eggs.Overstreetia and Pseudopalaeorchis Kamegai, 1970. Since the key was produced two further zoogonid genera have been described with tandem testes, Whitegonimus Jeżewski, Zdzitowiecki & Laskowski, 2009 and Oesophagotrema Chaari, Derbel & Neifar, 2011  off Sodwana, Natal, South Africa  off Heron Island, Queensland, Australia  and the paratype of O. olsoni (BMNH 2005.3.11.6) and included the data in Only six ovigerous specimens of O. cribbi the enlarged spines form an arc beside the aperture of the oral sucker and pass dorsally around the oral sucker region. This contrasts with the condition in O. sodwanaensis where the oral spine rows are limited to the anterior part of the oral sucker region , Perciformes, Mullidae, manybar goatfish.Host: Site: digestive tract.Locality: Off Récif Kué, Middle of Reef .Specimens: JNC2827.The zoogonid fauna of New Caledonian waters as described here is small, and probably represent a low proportion of the complete fauna as the Zoogonidae (and particularly the Lepidophyllinae) is one of the relatively few digenean families with a good representation in the deep-sea, i.e., off the continental shelf . NeverthDeretrema combesae, Deretrema combesorum, Dupliciporia lanterna and Overstreetia cribbi n. sp.Four species (31% of the fauna) are endemic: Diphterostomum tropicum.One species (8%) is restricted to South Western Pacific close to New Caledonia (FAO Major Fishing Area 71): Diphterostomum plectorhynchi, Parvipyrum acanthuri, Deretrema triodontis and Sacculoacetabulum ohjibah.Four species (31%) are found in the northern and southern Western Pacific (FAO 61 & 71): Deretrema acutum.One species (8%) is reported in the Western and Central Pacific : Zoogonus pagrosomi.One species is reported from sites across the Pacific Ocean : Zoogonoides viviparus, Lecithostaphylus nitens. These are probably cryptic complexes.Two species (15%) are cosmopolitan :


Dr. Sara Ares-Carrasco was not correctly included in the author byline. Dr. Ares-Carrasco should be listed as the fifth author and affiliated with Instituto de Investigaciones Sanitarias-Fundación Jiménez Díaz, Madrid, Spain. The contributions of this author are as follows: Performed the experiments. Dr. Ares-Carrasco's name should be abbreviated in the Author Contributions Statement as follows: SAC.The correct Citation is: Picatoste B, Ramírez E, Caro-Vadillo A, Iborra C, Ares-Carrasco S, et al. (2013) Sitagliptin Reduces Cardiac Apoptosis, Hypertrophy and Fibrosis Primarily by Insulin-Dependent Mechanisms in Experimental type-II Diabetes. Potential Roles of GLP-1 Isoforms. PLoS ONE 8(10): e78330. doi:10.1371/journal.pone.0078330.


The fourth author's name was misspelled in the author list and citation. The correct name is David J. Diemert.The citation should read: Saichua P, Sithithaworn P, Jariwala AR, Diemert DJ, Sithithaworn J, et al. (2013) Microproteinuria during Opisthorchis viverrini Infection: A Biomarker for Advanced Renal and Hepatobiliary Pathologies from Chronic Opisthorchiasis. PLoS Negl Trop Dis 7(5): e2228. doi:10.1371/journal.pntd.0002228


The first author's name was spelled incorrectly. The correct name is: Tamara Ben-Ari. The correct citation is: Ben-Ari T, Neerinckx S, Gage KL, Kreppel K, Laudisoit A, et al. (2011) Plague and Climate: Scales Matter. PLoS Pathog 7(9): e1002160. doi:10.1371/journal.ppat.1002160


The name of the second author is incorrect. The correct name is: Ricardo Sant'Anna. The correct Citation is: Ferreira P, Sant’Anna R, Varejão N, Lima C, Novis S, et al. (2013) Structure-Based Analysis of A19D, a Variant of Transthyretin Involved in Familial Amyloid Cardiomyopathy. PLoS ONE 8(12): e82484. doi:10.1371/journal.pone.0082484.The correct abbreviation of the second author's name in the Author Contributions Statement is: RS.