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Bayesian phylogram illustrating the relationships between the three species of the Didymodon tophaceus complex. Didymodon spadiceus, Didymodon maximus and Didymodon luridus were used as outgroup species. Posterior probabilities as well as MP and NJ bootstrap values are indicated.
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![Figure 3. Didymodon tophaceus (from Ku a n ] ˇ era 5861 ): (A) leaf;...](profile/Rosa-Ros-Espin/publication/233441764/figure/fig1/AS:299942179557382@1448523196300/Didymodon-tophaceus-from-Ku-a-n-era-5861-A-leaf-D-leaf-apex-and-G_Q320.jpg)
Recently two Mediterranean species closely related to Didymodon tophaceus (Brid.) Lisa have been described in the genus Didymodon Hedw.: Didymodon sicculus M. J. Cano et al. and Didymodon erosus J. A. Jiménez & J. Guerra. The former has been proved to be widespread around the Mediterranean basin in recent years, and this study points to a considera...
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... At the molecular level, only a few phylogenetic studies on Didymodon have been conducted to date, all with a very limited number of species (Werner et al., 2004(Werner et al., , 2005(Werner et al., , 2009Kučera & Ignatov, 2015, Kučera et al., 2018Ronikier et al., 2018;Inoue et al., 2020). Werner et al. (2004), based on ITS sequences and including 12 species of Didymodon, placed Kingiobryum H. Rob. ...
... (Dicranaceae), an Andean monotypic genus, into Didymodon. Werner et al. (2009) and Kučera et al. (2018) studied the systematic affinities between Didymodon tophaceus (Brid.) Lisa and its relatives Didymodon sicculus M.J. Cano, Ros, García-Zam. ...
... The systematic affinities of Didymodon sicculus and D. erosus, two species closely related to Didymodon tophaceus, were studied by Werner et al. (2009) and Kučera et al. (2018) based almost exclusively on European specimens. Both works showed that these species were closely related, and that they formed three clearly differentiated genetic lineages. ...
Didymodon s.l. is one of the largest genera in the moss family Pottiaceae, with about 122 species distributed in all continents. This, together with its high degree of morphological variation, has made it one of the taxonomically most challenging genera of Pottiaceae. Circumscription of Didymodon s.l. has been, and still is, controversial. To date, the only molecular study that has investigated the delimitation of the genus has been far from comprehensive (35 samples from 27 species), limited in geographical scope (mainly restricted to Europe), and based exclusively on ITS sequences. To evaluate the circumscription of Didymodon s.l. and its relationships with the allied genera Andinella, Gertrudiella, and Tridontium, we conducted phylogenetic analyses of DNA sequences for three plastid markers (atpB-rbcL, trnG, and trnL-F) and one nuclear locus (ITS) for 335 samples representing 86 species of Didymodon s.l. (ca. 70%), and all taxa of the genera Andinella, Gertrudiella, and Tridontium. Individual markers and concatenated matrices were analyzed using maximum likelihood and Bayesian approaches. Our results indicate that Didymodon s.l. is not monophyletic, because Andinella, Gertrudiella, and Tridontium species are nested within it. Species of these four genera can be divided into eight well-supported and morphologically distinct genera: Didymodon s.s., Geheebia, Gertrudiella, Husnotiella, Trichostomopsis, Tridontium, Vinealobryum, and Zanderella. In correspondence with the results presented, 38 new combinations, 10 new synonyms, and a new name are provided for those taxa where required, and lectotypes are designated for 13 names. A diagnostic key to the eight recognized genera is provided.
... We employed one nuclear (ITS) and two chloroplast markers (rps4 and trnM-trnV), which had been used successfully in previous phylogenetic studies in Didymodon s. lat. and enabled the re-use of earlier results and easier interpretation of new data (Werner et al. 2004(Werner et al. , 2005(Werner et al. , 2009Kučera and Ignatov 2015;Kučera et al. 2018;Ronikier et al. 2018;Jiménez et al. 2022;Zhang et al. in press). Phylogenetic trees are created and shown separately. ...
Inner Mongolia steppe is one of the suitable habitats for Didymodon species and a new species, Didymodon manhanensis C. Feng & J. Kou from Manhan Mountain in semi-arid region in Inner Mongolia, China is described and illustrated. It is characterised by leaves incurved and slightly twisted when dry, spreading when moist, narrowly lanceolate from an ovate base; subulate and fragile leaf apices; distally bistratose leaf margins that are recurved in proximal 2/3–3/4; excurrent costa with guide cells in 2–3 layers and without ventral stereids; smooth laminal cells and red KOH laminal colour reaction. Our morphological analyses and molecular results, based on DNA sequences of ITS, rps 4 and trn M- trn V, confirm that D. manhanensis belongs to a group that includes D. obtusus J. Kou, X.-M. Shao & C. Feng and D. daqingii J. Kou, R.H. Zander & C. Feng. This new species is compared with similar species and its phylogenetic position and ecology are discussed.
... However, a few studies have clarified much of the confusion, such as the separating Didymodon from Barbula Hedw. (Saito 1975, Zander 1978, 1993, the key characters useful in species recognition (Zander 1978(Zander , 1993(Zander , 2007 and molecular studies (Werner et al. 2005b(Werner et al. , 2009. A recent significant approach related to Didymodon s. lat. ...
A new species, Didymodon sinicus C. Feng & J. Kou, is described and illustrated based on materials collected from Inner Mongolia, Yunnan and Tibet of China. It is characterized by ovate-lanceolate to ovate-triangular leaves that are erect to patent when wet, auriculate leaf base, acute leaves apices, distally bistratose leaf margins, the yellowish color in KOH, short-excurrent costa with guide cells in 2–3 layers and without ventral stereids, and smooth laminal cells. A phylogenetic analysis based on DNA sequences of ITS, rps4, trnM-trnV confirms D. sinicus as a sister to the clade of D. obtusus J. Kou, X.-M. Shao & C. Feng and the new species belongs to the genus Didymodon s.str.
... based on ITS sequences. Werner et al. (2009) and Kučera et al. (2018) confirmed D. tophaceus, D. erosus and D. sicculus into a highly supported lineage of closely related taxa by ITS and the chloroplast genome. Ronikier et al. (2018) used nuclear (ITS) and plastid (atpIH, trnLF, trnG, rps4) sequences to assess the genetic differentiation of D. gelidus in the Southern Hemisphere and D. brachyphyllus in the Northern Hemisphere. ...
... Our study provides the most comprehensive phylogenetic relationship of species within the genus Didymodon s.lat. based on five plastid markers (trnL-trnF, trnG, atpB-rbcL, rps4 and trnM-trnV) and one nuclear locus (ITS), including a total of 107 species, some of which have not been documented by previous studies (Werner et al., 2004(Werner et al., , 2005(Werner et al., , 2009Kučera & Ignatov, 2015;Kučera et al., 2018;Ronikier et al., 2018;Inoue et al., 2020;Jiménez et al., 2022). ...
Didymodon Hedw., with approximately 140 species in the family Pottiaceae, is distributed nearly throughout the world, with the greatest diversity and important ecological functions in drought lands and alpine ecosystems. Several studies involving morphology, molecular systematics and macro-systematic analysis have addressed the infrageneric classification of Didymodon, but controversy over the position of the infrageneric and species classification remains due to its high degree of morphological variation in micro-habitats and strong sensitivity to climate change at regional and global scale. To date, only a few phylogenetic studies have been conducted with an incomplete number of Didymodon species; further, there is no study published regarding the divergence time of Didymodon. Consequently, we constructed a comprehensive phylogenetic relationship of Didymodon species, a total of 107 species, based on one nuclear (ITS) and five chloroplast DNA. Moreover, divergence time analysis was conducted to infer the age of origin and divergence of Didymodon species. Our results presented the largest-scale phylogenetic relationship of Didymodon to date and resolved the phylogenetic status of some controversial taxa and the new species. The divergence time estimation showed that Didymodon species originated to be around the early Cretaceous, and the diversification was concentrated in the Cretaceous and Eocene. Paleoclimate and environmental change have a direct impact on the origin and divergence of Didymodon species by shaping their morphology, resource availability and ecological niche. Our study will help understand species origin and speciation as well as reflecting species adaptability and experience to historical events.
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... At the molecular level, only a few phylogenetic studies on Didymodon have been conducted to date, all with a very limited number of species (Werner et al., 2004(Werner et al., , 2005(Werner et al., , 2009Kučera & Ignatov, 2015, Kučera et al., 2018Ronikier et al., 2018;Inoue et al., 2020). Werner et al. (2004), based on ITS sequences and including 12 species of Didymodon, placed Kingiobryum H. Rob. ...
... (Dicranaceae), an Andean monotypic genus, into Didymodon. Werner et al. (2009) and Kučera et al. (2018) studied the systematic affinities between Didymodon tophaceus (Brid.) Lisa and its relatives Didymodon sicculus M.J. Cano, Ros, García-Zam. ...
... The systematic affinities of Didymodon sicculus and D. erosus, two species closely related to Didymodon tophaceus, were studied by Werner et al. (2009) and Kučera et al. (2018) based almost exclusively on European specimens. Both works showed that these species were closely related, and that they formed three clearly differentiated genetic lineages. ...
Didymodon s.l. is one of the largest genera in the moss family Pottiaceae, with about 122 species distributed on all continents. This, together with its high degree of morphological variation has made it one of the taxonomically most challenging genera of Pottiaceae. Circumscription of Didymodon s.l. has been, and still is, controversial. To date, the only molecular study that has investigated the delimitation of the genus has been far from comprehensive (35 samples from 27 species), limited in geographical scope (mainly restricted to Europe), and based exclusively on ITS sequences. In order to evaluate the circumscription of Didymodon s.l. and its relationships with the allied genera Andinella, Gertrudiella and Tridontium, we conducted phylogenetic analyses of DNA sequences for three plastid markers (atpB‐rbcL, trnG and trnL‐F) and one nuclear locus (ITS) for 335 samples representing 86 species of Didymodon s.l. (ca. 70%), and all taxa of the genera Andinella, Gertrudiella and Tridontium. Individual markers and concatenated matrices were analyzed using maximum likelihood and Bayesian approaches. Our results indicate that Didymodon s.l. is not monophyletic because Andinella, Gertrudiella and Tridontium species are nested within it. Species of these four genera can be divided into eight well‐supported and morphologically distinct genera: Didymodon s.s., Geheebia, Gertrudiella, Husnotiella, Trichostomopsis, Tridontium, Zanderella, and Vinealobryum. In correspondence with the results presented, 38 new combinations, 10 new synonyms, and a new name are provided for those taxa where required, and lectotypes are designated for 12 names. A diagnostic key to the eight recognized genera is provided.
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... A metadata study has been conducted by Zander (2019b) of the number of molecular strains per species in the Pottiaceae (these all termed "races" in that study). The data for this study was based on several recent publications (Alonso et al., 2016;Cano et al., 2009;Grundmann et al., 2006;Köckinger et al., 2010;Kučera, Ignatov, 2015;Kučera et al., 2013Kučera et al., , 2018Werner et al., 2005Werner et al., , 2009Werner et al., , 2014. Molecular strains were inferred when two exemplars of the same species share at minimum one trait not shared by a third exemplar of the same species, i.e., an internal node with two exemplars terminal and one more basal. ...
The analytic orientation of this paper is intended as a replacement for the antiquated but still prevalent phylogenetic inferential models and techniques of the late 20th century that are focused entirely on shared descent. Serial descent, that is, progenitor to descendant, may occur at the species or infraspecies level. In molecular systematics, species level paraphyly occurs when two examples of the same species are separated on a cladogram by a second species. This implies linear macroevolution of the second species from the first. Molecular cladograms often show cladistic structure (branching) among examples of the same species. If well-supported, such indicates a potential for evolution. A range of infraspecific and intraspecific cladistic structure in species of Pottiaceae (Bryophyta) was demonstrated in previously published molecular cladograms and data sets of other authors. This includes well-supported cladistic structure of molecular strains, and well-supported paraphyly involving other species. Large numbers of base pair changes among strains are considered here evidence of evolvability and increasing age of a species. Infraspecific strains are apparently lost in older species through speciation and extinction. Cluster analysis using DNA metadata of Oxystegus species matched published molecular cladograms to a large extent. The fact that apparent molecular strains are present in both nonparaphyletic and paraphyletic species, about half the species studied, shore up the theory that internal racial differentiation at the molecular level leads to or signals serial descent of multiple extant morphotaxa. It is because much infraspecific molecular cladistic structure exists that newly speciated taxa are already strongly cladistically dichotomized. Thus, the ultimate source of molecular paraphyly is internal to each species, and does not imply polyphyly by convergent species or cryptic taxa. Molecular systematics cannot effectively model progenitor-descendant radiation. Species with many strains are potential sources of future biological diversity. Recognition of differential evolvability may allow facilitation of complex, interactive, diverse ecosystems successfully tracking climate change.
... A metadata study has been conducted by Zander (2019b) of the number of molecular strains per species in the Pottiaceae (these all termed "races" in that study). The data for this study was based on several recent publications (Alonso et al., 2016;Cano et al., 2009;Grundmann et al., 2006;Köckinger et al., 2010;Kučera, Ignatov, 2015;Kučera et al., 2013Kučera et al., , 2018Werner et al., 2005Werner et al., , 2009Werner et al., , 2014). Molecular strains were inferred when two exemplars of the same species share at minimum one trait not shared by a third exemplar of the same species, i.e., an internal node with two exemplars terminal and one more basal. ...
General recommendations regarding proper type designation of names of taxa described by Turczaninow in his Animadversiones series of articles (as well as in some other publications) are provided. It is concluded that, as clearly indicated in the protologues, all (or almost all) taxa described in these publications are based on specimens from the private herbarium of Turczaninow which was donated in the 1840s to the Kharkiv University (CWU) and in the 1940s was transferred to the Institute of Botany in Kyiv (KW). Consequently, holotypes and syntypes of these taxa are now almost exclusively in KW. Several cases of correct and incorrect type designations are discussed, specifically of some South American Brassicaceae, Geraniaceae and Hypericaceae, Central American Malvaceae, and southern African Polygalaceae. Information on the re-discovered holotype (KW) of Abelmoschus achanioides Turcz. (now accepted as Malvaviscus achanioides (Turcz.) Fryxell, Malvaceae) is provided, and an earlier lectotypification of that name with a specimen from G is considered ineffective. The holotype of Stenocalyx involutus Turcz. (now considered a synonym of Mezia includens (Benth.) Cuatrec., Malpighiaceae) was originally in the Turczaninow herbarium, but
the whole folder with that specimen is now missing in KW (considered lost or destroyed), and it was already missing in the mid-1920s, when
the collection was still in CWU. Because of that the lectotype of S. involutus is designated here, the specimen from MPU, to replace the lost or destroyed holotype. The need for thorough analysis of protologues, available original material, and associated information for correct type designation/indication is emphasized.
... A study has been conducted by Zander (2019b) of the number of molecular races per species in the Pottiaceae. The data for this study were extracted from several recent publications (Alonso et al. 2016;Cano et al. 2009;Grundmann et al. 2006;Köckinger et al. 2010;Kučera & Ignatov 2015;Kučera et al. , 2018Werner et al. 2005Werner et al. , 2009Werner et al. , 2014. Molecular races were inferred when two exemplars of the same species share on a cladogram a trait not shared by a third exemplar of the same species, i.e., an internal node with two exemplars terminal and one more basal. ...
Three species of Pottiaceae (Bryophyta) of limited distribution worldwide are reported for hyperoceanic northwestern North America. The Asian Pseudosymblepharis angustata is a genus and species new to North America from Alaska. Oxystegus daldinianus, previously known from Europe and the Southern Appalachians in the U.S.A., is reported as new to Canada from British Columbia. A new species is described in Oxystegus from British Columbia. These genera continue to be recognized here instead of the broad molecular systematics concept of Chionoloma in part because of lack of resolution of molecular cladograms reported by a recent study on molecular races in the Pottiaceae.
... Four major molecular studies are involved in re-classification involving Didymodon s.lat., those of Kučera & Ignatov (2015) focusing on Didymodon sect. Rufiduli (P.C.Chen) R.H. Zander, Kučera et al. (2018) on Didymodon tophaceus and allies, and of Werner et al. (2005Werner et al. ( , 2009, which also deals with Didymodon tophaceus and near relatives. The three studies include exemplars of other species in Didymodon s.lat. ...
... Of course generic diversity is common in mosses (e.g., Shaw & Schneider 1995;Shaw 2000Shaw , 2001 but we are concerned here with DNA differences at a level important to taxonomy and classification. Table 3 is the summary result of tabulation of the number of molecular races per species in the Pottiaceae as given in several recent publications (Werner et al. 2005(Werner et al. , 2009(Werner et al. , 2014Grundmann et al. 2006;Cano et al. 2009;Köckinger et al. 2010;Kučera & Ignatov 2015;Kučera et al. 2013;Alonso et al. 2016). Two molecular races are here inferred when two exemplars of the same species share a molecular trait not shared by a third exemplar of the same species, i.e., there is an internal node with two exemplars terminal and one more basal. ...
Molecular analysis has several problems including (1) small samples, (2) radiation of molecular strains or races that separately generate their own species-level lineages, (3) all problems associated with cladistics analysis done using morphological data, (4) conflicting studies, (5) reuse of data contributing to small samples and absence of experimental replication, (6) no native taxon concept, and (7) use of combined data sets to achieve an invented gene history. A macroevolutionary re-interpretation of a recent molecular revision of species of Didymodon s.lat. (Bryophyta) introduces a monophyletic evolutionary model fully compatible with both molecular and morphological data. Multiple molecular races and their contribution to paraphyly and apparent polyphyly are quantized through analysis of recent published studies to demonstrate uncertainty of monophyly on the order of a distance of 4.5 contiguous nodes per species in molecular cladograms. The number of trait changes per speciation event averaged 3.57, on a par with previous studies. This work is the first successful prediction of the actual existence of a missing link, Exobryum rufidulum, hypothetically described in a previous macroevolutionary analysis. For the first time a taxon higher than genus was established based on an extension of the empirical dissilient genus concept. Six new combinations are made in Exobryum. A new genus, Aithobryum, is established with three species transferred from Didymodon. Didymodon sinuosus and D. californicus are transferred to Vinealobryum.
... Didymodon sicculus (obr. v příloze č. 1, 2) je nejblíže příbuzný druhu D. tophaceus a společně s druhem D. erosus J. A. Jiménez & J. Guerra tvoří komplex blízce příbuzných a někdy obtížně morfologicky rozlišitelných druhů (Werner et al. 2009). Na první pohled však může být D. sicculus mezi našimi druhy zaměněn spíše za D. luridus, od kterého se liší (spolu s ostatními dvěma druhy komplexu) zejména přítomností papil na obou stranách listové čepele. ...
Altogether 139 taxa (9 liverworts and 130 mosses) were recorded during the course of four excursions to the region of the Pavlov Hills in South Moravia, including one species new to the Czech Republic (Didymodon sicculus; the identity of another species not recorded in the Czech Republic, Pterygoneurum kozlovii, needs to be confirmed), one species considered Vanished (Hennediella heimii), three Critically Endangered species (Bryum intermedium, Hilpertia velenovskyi and Tortula inermis), five Endangered taxa (Aloina aloides var. ambigua, Conardia compacta, Grimmia anodon, G. crinita and Pseudocrossidium revolutum) and five Vulnerable species (Acaulon triquetrum, Didymodon cordatus, D. sinuosus, Pterygoneurum subsessile and Rhynchostegium megapolitanum). Important records are briefly commented on.
