Cryptogamie, Bryologie, 2016, 37 (4): 361-382 © 2016 Adac. Tous droits réservés Lewinskya, a new genus to accommodate the phaneroporous and monoicous taxa of Orthotrichum (Bryophyta, Orthotrichaceae) Francisco LARA a*, Ricardo GARILLETI b, Bernard GOFFINET c, Isabel DRAPER a, Rafael MEDINA c, Beatriz VIGALONDO a & Vicente MAZIMPAKA a aDepartamento de Biología (Botánica), c/ Darwin 2, Facultad de Ciencias, Universidad Autónoma de Madrid, E-28049 Madrid, Spain. bDepartamento de Botánica y Geología, Facultad de Farmacia, Universidad de Valencia, Av. Vicente Andrés Estellés s/n, E-46100 Burjassot, Spain. cEcology and Evolutionary Biology, University of Connecticut, 75 North Eagleville Rd., Storrs, Connecticut 06269-3043, U.S.A. Abstract – Molecular analyses have consistently evidenced the phylogenetic heterogeneity of Orthotrichum Hedw., and suggested the need to segregate the species with superficial stomata in a separate genus. A recent proposal has been made to accommodate the monoicous species with such stomata in the genus Dorcadion Adans. ex Lindb., which is, however, an illegitimate name according to the current Code of nomenclature of algae, fungi and plants. Consequently a new name is required, and the generic name Lewinskya F.Lara, Garilleti & Goffinet is proposed. New combinations are made for all the species included in the new genus. Given the long history of the genus Orthotrichum and the similarities between this genus and Lewinskya, the morphological and geographic circumscriptions of both genera are provided to define them accurately. The taxa remaining in Orthotrichum s.str. are also listed. Biogeography / Diversity / Musci / Nomenclature / Systematics INTRODUCTION Acrocarpic mosses growing on trees or rocks, with orthotropic stems, leaves with short isodiametric cells above, elongate basal cells, undifferentiated marginal cells, a mitrate calyptra and with sporangia, which upon dehiscence reveal a diplolepideous (though often variously reduced) peristome, are accommodated in the genus Orthotrichum Hedw. The genus is part of the tribe Orthotricheae Goffinet & Vitt, within the Orthotrichaceae Arnott, a highly speciose and cosmopolitan family. At the end of the last century, Goffinet & Vitt (1998) proposed a revised classification whereby the Orthotricheae included five genera (i.e., Muelleriella Dusén, Orthomitrium Lewinsky & Crosby, Orthotrichum, Stoneobryum Norris & H.Rob. and Ulota D.Mohr). Lewinsky-Haapasaari & Hedenäs (1998) subsequently * Corresponding author: francisco.lara@uam.es doi/10.7872/cryb/v37.iss4.2016.361 362 F. Lara et al. segregated O. paraguense Besch. in the new genus Sehnemobryum Lewinsky & Hedenäs. Most species of the Orthotricheae belong to either Orthotrichum or Ulota, with the remaining genera holding only one to five species (Crosby et al., 1999). Generic concepts within the tribe reflect patterns in the ornamentation of laminal cells, the precocious germination of spores (endospory), differentiation of the laminal margin, costal anatomy and architecture and distribution of the stomata (Gofffinet & Vitt, 1998; Lewinsky-Haapasaari & Hedenäs, 1998). However, both speciose genera harbor variation in these traits, most notably with Orthotrichum including species with either superficial (phaneroporous) or immersed (cryptoporous) stomata. Lewinsky (1993a) critically revised the taxonomic diversity of Orthotrichum. The circumscription of the genus continues to grow, with 45 taxa added since 1993. The species have been historically organized into subgenera and sections (e.g., Vitt, 1971). Phylogenetic inferences from morphological traits providing the basis for these infrageneric taxa suggested, however, significant homoplasy in these traits (Lewinsky-Haapasaari & Hedenäs, 1998). Furthermore, the phylogenetic significance of transformations in the generic traits appeared ambiguous, with Goffinet et al. (1998) providing the first evidence from analyzing rbcL sequences that Orthotrichum as circumscribed and diagnosed by Lewinsky (1993a) was an artificial lineage. Goffinet et al. (2004) subsequently corroborated this hypothesis based on inferences from four loci, demonstrating that a) species with superficial stomata shared a unique ancestor with Ulota, b) Muelleriella and Orthomitrium were nested among species with immersed stomata, c) Orthotrichum exiguum belonged to the tribe Zygodonteae (Schimp.) Goffinet, but that d) Stoneobryum and Sehnemobryum were indeed distinct. However, Goffinet et al. (2004) preferred to retain Orthotrichum sensu Lewinsky until the inferences could be strengthened by additional data from a more extensive sampling of taxa. Thus by 2004, the Orthotricheae included four genera: Orthotrichum, Sehnemobryum, Stoneobryum and Ulota (Goffinet & Buck, 2004). The polyphyly of Orthotrichum was repeatedly suggested by inferences from molecular data (Goffinet et al., 2007; Plášek et al., 2009, 2011; Sawicki et al., 2009, 2010, 2012; Vigalondo et al., 2016a). Sawicki et al. (2010) proposed the first systematic consequences since 2004 to reflect the phylogenetic reconstruction, and reinstated Nyholmiella Holmen & E.Warncke for O. gymnostomum Bruch ex Brid. and O. obtusifolium Brid., traditionally composing subgenus Orthophyllum Delogne (Vitt, 1971). Plášek et al. (2015) further divided Orthotrichum and segregated O. lyellii Hook. & Taylor in the new genus Pulvigera Plášek, Sawicki & Ochyra, and resurrected Dorcadion Adans. ex Lindb. to accommodate the monoicous and phaneroporous taxa of Orthotrichum. The genus Orthotrichum s.str. would thus be restricted to the cryptoporous species. Although this revised classification of the Orthotricheae settles the incongruence between phylogenetic inferences and systematic concepts, assuming that a criterion of monophyly is considered necessary (see Zander, 2013), nevertheless it invokes an illegitimate name. Consequently we propose a new name, Lewinskya gen. nov. for Dorcadion Adans. ex Lindb. sensu Plášek et al. (2015). Furthermore we provide an expanded morphological description of the genus, an assessment of its geographic distribution and a complete taxonomic circumscription with all necessary new combinations. With the proposed segregation of a significant number of species from Orthotrichum s.str., the morphological and geographic distribution of this genus needs to be revised, too. Hence we present along with the current circumscription of Orthotrichum a description of its morphological traits and distribution. Lewinskya gen. nov. 363 METHODS The characterization of morphological traits and geographic distributions is based on descriptions in Lewinsky (1993a) complemented with those from all subsequently discovered and recognized taxa (Lewinsky, 1993b, 1994; Ignatov & Lewinsky-Haapasaari, 1994; Lara et al., 1994, 1996, 1999a, 1999b, 1999c, 2000, 2006, 2007, 2009a, 2009b, 2009c, 2010; Lewinsky-Haapasaari, 1994, 1995, 1999a, 1999b; Vitt, 1994, 2014; Lewinsky-Haapasaari & Tan, 1995; Tan & Yu, 1997; Vitt & Darigo, 1997; Lewinsky-Haapasaari & Norris, 1998a, 1998b; Magill & van Rooy, 1998; Mazimpaka et al., 1999, 2000a, 2000b; Matteri, 2000; Garilleti et al., 2001, 2006a, 2006b, 2009, 2011; Ignatov et al., 2001, 2006; Allen, 2002; Goffinet, 2002; Draper et al., 2003, 2006; Akatova et al., 2004; Norris et al., 2004; Kürschner & Erdağ, 2005; Goffinet et al., 2007; Albertos et al., 2008; Medina et al., 2008, 2009, 2011, 2012, 2013; Atwood & Allen, 2009; Frahm et al., 2009; Plášek et al., 2009, 2014; Fedosov & Ignatova, 2010, 2011; Jia et al., 2011; Bosanquet & Lara 2012; Lara & Garilleti, 2014; Wang & Jia, 2014; Ellis et al., 2015a; Vigalondo et al., 2016b). Distribution areas used for listed taxa and maps follow Wijk et al. (1964). RESULTS AND DISCUSSION Nomenclature and taxonomy Lindberg (1878) proposed to restore the pre-Hedwigian name Dorcadion (Adanson, 1763) for the genus Orthotrichum (Hedwig, 1801), thus creating a superfluous name. The author (Lindberg, 1878: 6) pointed this out neatly and indisputably, when he refers to the names of Adanson (1763): “All of the new ones are composed of rather heterogeneous links and therefore unnatural, with the exception of two, Sekra and Dorcadion, the former the same as Cinclidotus P.-B., the latter completely the same as Orthotrichum Hedw.” (“Af de nya äro alla sammansatta af ganska heterogena länkar och derför onaturliga, med undantag af två, Sekra och Dorcadion, det förra alldeles detsamma som Cinclidotus P.-B., det sednare fullkomligt liktydigt med Orthotrichum Hedw.”). A few pages later, Lindberg (1878: 35) created some confusion when he listed the genera that would constitute the Grimmiaceae, since using a broad concept he cited “Weissia (Ulota), Dorcadion (Orthotrichum Hedw. p. p.), ...”, which could suggest that Dorcadion is a new taxon incorporating only part of the members of Orthotrichum. However, what he did here was to indicate that under Dorcadion he included all the species of Orthotrichum as defined by Hedwig, except for those included in Weissia Ehrh. ex G. Gaertn., B.Mey. & Scherb. (Ulota D.Mohr). At that time, however, Orthotrichum already excluded the species of Ulota, given the combination U. crispa (Hedw.) Brid. established by Bridel (1818). Furthermore, Lindberg (1879), in his Musci Scandinavici, proposed two subgenera within Dorcadion, Calyptoporus and Gymnoporus (nomina nuda) that included the species with immersed and superficial stomata, respectively, that had until then been included in Orthotrichum (excluding Ulota). Therefore, we argue that Lindberg had the same concept for Dorcadion and Orthotrichum, once Ulota species were excluded. Consequently, Dorcadion is superfluous and thus illegitimate according to the Code of nomenclature of algae, fungi and plants 364 F. Lara et al. (McNeill et al., 2012), as previously indicated by Wijk et al. (1964) and Lewinsky (1993a). Since Dorcadion is an illegitimate name, it must be replaced. We propose the name Lewinskya F.Lara, Garilleti & Goffinet, in honour of the late Jette LewinskyHaapasaari, whose taxonomic work underlies much of our current knowledge and understanding of the diversity of the Orthotrichaceae. The establishment of a new genus requires the designation of a type, which ideally reflects the typical morphological features of the genus. With the proposed selection of Orthotrichum striatum Hedw., each of the three species originally included in Orthotrichum by Fig. 1. Regional diversity and endemicity of Lewinskya F.Lara, Garilleti & Goffinet considering the world geographical areas used in Index Muscorum (Wijk et al., 1964). The first numeral indicates the Lewinskya gen. nov. 365 Hedwig (1801) are now the type species for the three speciose genera of the Orthotricheae: Orthotrichum s.str., Ulota, and Lewinskya. Lewinskya F.Lara, Garilleti & Goffinet gen. nov. Type: Lewinskya striata (Hedw.) F.Lara, Garilleti & Goffinet, comb. nov. ≡ Orthotrichum striatum Hedw. Dorcadion Adans. ex Lindb. nom. illeg. sensu Plášek et al. (2015). Diagnosis: Orthotricho similis, a quo plerumque differt foliorum basalibus cellulis elongatis, parietibus incrassatis nodulosisque, sexuali conditione gonioautoica, total number of taxa recognized for the respective areas. The second numeral (in bold) refers to the number of taxa exclusive of each area, indicating between brackets the endemicity rate for each area. 366 F. Lara et al. capsulis saepe laevibus, stomatibus phaneroporis, endostoma sine membrana connectivali, sporisque diametro generatim 20 μm maiore. Circumscription of Lewinskya and Orthotrichum s.str. Lewinskya Plants small or large, to 5 cm high, rarely smaller than 1 cm, in more or less loose cushions. Leaves erect or erect-patent when dry, sometimes flexuose, Fig. 2. Regional diversity and endemicity of Orthotrichum Hedw. s.str. considering the world geographical areas used in Index Muscorum (Wijk et al., 1964). The first numeral indicates the total number of taxa Lewinskya gen. nov. 367 rarely crisped, with plane or recurved margins, sometimes undulate, exceptionally incurved. Leaf apex acute to long acuminate, sometimes aristate, rarely obtuse, rounded or retuse. Leaf lamina unistratose, exceptionally predominantly bistratose in the upper half. Upper leaf cells isodiametric or oblong, sometimes elongate, variably papillose or smooth. Basal leaf cells mostly elongate, with incrassate walls, often dimorphic, with rows of cells with nodulose and coloured walls between cells with hyaline and more or less sinuose walls; alar cells occasionally differentiated. Propagules absent, except for one report of claviform propagules, with longitudinal and transverse walls, similar to those of Zygodon. Goniautoicous. Calyptra more or less hairy, frequently densely so, exceptionally naked. Vaginula naked or hairy, recognized for the respective areas. The second numeral (in bold) refers to the number of taxa exclusive of each area, indicating between brackets the endemicity rate for each area. 368 F. Lara et al. commonly with some long and papillose hairs. Sporophytes usually 1 per perichaetium, occasionally in pairs in several species, consistently many in some species. Seta short, generally as long as capsule, or more or less longer, exceeding the perichaetial leaves. Capsule immersed, emergent or exserted, then often longly so. Urn frequently cylindrical, sometimes ovoid, rarely globose or cup-shaped, often smooth or with 8 ribs scarcely and variably developed, less frequently with prominent ribs. Exothecial bands not or scarcely differentiated in most species, well differentiated in others. Stomata superficial, generally in the middle of the urn, less frequently in its upper or lower portions. Peristome generally double, rarely simple, exceptionally vestigial. Prostome frequent, variably developed. Exostome of 8 pairs of fused teeth, less frequently of 16 free teeth, occasionally very reduced or almost absent. Endostome of 8 or 16 segments, rarely vestigial or absent, frequently very wide but sometimes more or less narrow, almost always internally biseriate, exceptionally uniseriate, almost always roughly ornamented, with irregular or regular margins, without connecting membrane. Spores frequently over 20 μm in diameter, usually unicellular, sometimes with a proportion two-celled, exceptionally multicellular. The genus comprises 70 taxa (66 species and 4 varieties), which are mostly epiphytes, with some being saxicolous. The diversity of species is unevenly distributed (Fig. 1). The Northern Hemisphere harbors 46 taxa (66% of the total) and 38 of these (83%) are not known from the austral territories. Thirty-two taxa (46% of the total) occur in the Southern Hemisphere, and 24 of these (75%) are absent from septentrional territories. The percentage of endemics in each region (based on the divisions in Wijk et al., 1964) is notably higher in the Southern Hemisphere. Orthotrichum s.str. Type: Orthotrichum anomalum Hedw. Plants small or medium sized, frequently less than 1 cm high, forming more or less compact cushions or, more rarely, patches. Leaves erect when dry, rarely contorted, with recurved margins, rarely plane. Leaf apex commonly acute or acuminate but frequently specialized: obtuse, rounded, denticulate, channeled, mucronate or piliferous. Leaf lamina generally unistratose, frequently with bistratose bands, in several species predominantly bistratose. Upper leaf cells isodiametric, rarely oblong, generally papillose, with simple or branched papillae, sometimes smooth or almost so. Basal leaf cells rectangular, with straight or somewhat sinuose walls, thin or slightly incrassate, only occasionally more or less nodulose, monomorphic; alar cells not differentiated. Propagules frequently present on leaves, more rarely also on rhizoids, cylindrical to vermiform, only with transverse walls, sometimes branched; exceptionally with rhizoidal tubers. Cladautoicous, always with a terminal perigonium in the male branches, frequently also with lateral perigonia. Calyptra variably hairy to naked. Vaginula naked or, less frequently, hairy. One sporophyte per perichaetium, polysety exceptional. Seta short, often extremely so, rarely longer than the perichaetial leaves. Capsule immersed to long-emergent, only occasionally exserted, rarely long-exserted. Urn cylindrical to urceolate, often shortly so, with 8 ribs, rarely 16, more or less prominent and wide, occasionally limited to the distal area, never totally smooth. Exothecial bands generally well differentiated. Stomata immersed, more or less covered by the subsidiary cells, exceptionally almost uncovered and apparently superficial, often limited to the urn base, occasionally spreading across the neck. Peristome generally double, rarely Lewinskya gen. nov. 369 simple or absent. Prostome occasional, variably developed. Exostome of 8 pairs of teeth, rarely 16 completely independent, occasionally vestigial or absent. Endostome of 8 or 16 segments, rarely completely absent, generally filiform, rarely wide, exceptionally fused, smooth or ornamented, with differentiated connecting membrane, sometimes incomplete. Spores usually unicellular, in most cases with diameters of less than 20 μm, pluricellular in some species. The genus comprises 103 taxa (94 species, 1 subspecies and 8 varieties) that are mostly epiphytes, although some species are exclusively or preferentially saxicolous. The distribution of species diversity and endemics is also very uneven (Fig. 2). The Northern Hemisphere hosts 83 taxa (81% of the total), with 76 of these (92%) being exclusive. In the Southern Hemisphere 27 taxa are known (26% of the total), and 20 of these (74%) do not reach any of the septentrional territories. Endemism is high across the Southern Hemisphere, but also in the Northern Hemisphere, and particularly in North America (Am 1) and Eastern Asia (the ChinaJapan area, AS 2; Fig. 2). The Mediterranean basin is a center of diversity (Lara et al., 2009b) not evidenced in the chosen geographic divisions. KEY TO GENERA Stomata superficial. Basal leaf cells mostly elongate, with incrasate and sinuose or nodulose walls. Goniautoicous. Endostome without connecting membrane. Gametophores usually taller than 1 cm. Capsules often smooth or faintly ribbed when dry, frequently exserted. Propagules absent. Spores frequently over 20 μm in diameter ........................................................................................... Lewinskya Stomata immersed, more or less covered by the subsidiary cells. Basal leaf cells rectangular, usually with thin, not nodulose walls. Cladautoicous. Endostome with connecting membrane, sometimes incomplete. Gametophores usually smaller than 1 cm. Capsules mostly strongly ribbed when dry, rarely exserted. Propagules frequently present. Spores usually less than 20 μm in diameter ......... ................................................................................................................................ Orthotrichum Acknowledgments. We would like to address our warmest thanks to Bill Buck for his critical advice in the nomenclatural part of this work. Bill, Tom Blockeel and Rosa María Ros made constructive comments and suggestions on an earlier version, which helped to improve the manuscript and are gratefully acknowledged. This research was supported by the Spanish Ministries of Education and Science (CGL2007-61389/BOS), Science and Innovation (CGL2011–28857/BOS), and Economy and Competitiveness (CGL2013–43246–P). Support to BG was provided by NSF grants DEB-1146295 & 1240045. REFERENCES ADANSON M., 1763 — Familles des plantes. Paris, Vincent. AKATOVA T.V., KHARZINOV Z.KH., IGNATOVA E.A. & IGNATOV M.S., 2004 — On three rare species of Orthotrichum (Orthotrichaceae, Musci) in the Caucasus. Arctoa 13: 41-49. ALBERTOS B., GARILLETI R., MAZIMPAKA V. & LARA F., 2008 — Re-evaluation of Orthotrichum rhytiore (Orthotrichaceae). The bryologist 111: 323-327. ALLEN B.H., 2002 — Moss flora of Central America. Part 2. 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Utrecht, International Bureau for Plant Taxonomy and nomenclature of the International Association for Plant Taxonomy. ZANDER R.H. 2013. Framework for Post-Phylogenetic Systematics. St. Louis, Zetetic, Publications. 374 F. Lara et al. APPENDIX 1 Taxa included in the genus Lewinskya F.Lara, garilleti & goffinet, with indication of their known distribution as shown in Fig. 1. Lewinskya acuminata (H.Philib.) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum acuminatum H.Philib., Rev. Bryol. 8: 28. 1881 Distribution: Am 1, Eur, AFR 1, AFR 2, AFR 5, AS 5. Lewinskya affinis (Schrad. ex Brid.) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum affine Schrad. ex Brid., Muscol. Recent. 2: 22-23. 1801 Distribution: Am 1, Eur, AFR 1, AFR 5, AS 1, AS 5. Lewinskya anaglyptodon (Cardot & Broth.) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum anaglyptodon Cardot & Broth., Kongl. Svenska Vetensk. Acad. Handl. n.s. 63: 31, 2 f. 10. 1923 Distribution: Am 6. Lewinskya araucarieti (Müll.Hal. in Brotherus) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum araucarieti Müll. Hal. in Brotherus, Ergebn. Bot. Exped. Südbras. Musci 287. 1924 Distribution: Am 5, Am 6. Lewinskya arborescens (Thér. & Naveau) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum arborescens Thér. & Naveau, Bull. Soc. Roy. Bot. Belgique 60: 48. 1927 Distribution: AFR 2. Lewinskya armata (Lewinsky & van Rooy) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum armatum Lewinsky & van Rooy, J. Bryol. 16: 69, f. 2–5. 1990 Distribution: AFR 4. Lewinskya bolanderi (Sull.) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum bolanderi Sull., Icon. Musc. Suppl. 64, pl. 46. 1874 Distribution: Am 1, Am 2. Lewinskya brassii (E.B.Bartram) F Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum brassii E.B.Bartram, Lloydia 5: 268, 25. 1942 Distribution: AS 4. Lewinskya breviseta (F.Lara, Garilleti & Mazimpaka) F.Lara, Garilleti & Goffinet, comb. et stat. nov. ≡Orthotrichum speciosum var. brevisetum F.Lara, Garilleti & Mazimpaka, J. Bryol. 25: 276, f. 2. 2003 Distribution: Eur, AFR 1, AS 5. Within the context of the new genus, L. breviseta differs from Lewinskya speciosa in significative morphological characters, such as the short seta and the ribbed capsules. Additionally, both mosses show very different ecological preferences and distribution ranges, being L. breviseta restricted to Mediterranean environments whereas L. speciosa is a Boreal widespread species that seems to avoid climates with strong summer drought (Ellis et al., 2015b). All these divergences lead us to propose the recognition of L. breviseta at the species level. Lewinskya brotheri (Dusén ex Lewinsky) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum brotheri Dusén ex Lewinsky, Mem. New York Bot. Gard. 45: 348, f. 133–143. 1987 Distribution: Am 6. Lewinskya cyathiformis (R.Br.bis) F.Lara, Garilleti & Goffinet, Lewinskya gen. nov. comb. nov. ≡Orthotrichum cyathiforme R.Br.bis, Trans. & Proc. New Zealand Inst. 27: 436, 39 f. 24. 1895 Distribution: AUSTR 2. Lewinskya dasymitria (Lewinsky in T.Koponen & J.X.Luo) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum dasymitrium Lewinsky in T.Koponen & J.X.Luo, Bryobrothera 1: 169, f. 1–2. 1992 Distribution: AS 2, AS 3. Lewinskya densa (Lewinsky) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum densum Lewinsky, Mem. New York Bot. Gard. 45: 336, f. 51–62. 1987 Distribution: Am 6. Lewinskya elegans (Schwägr. ex Hook. & Grev.) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum elegans Schwägr. ex Hook. & Grev., Edinburgh J. Sci. 1: 122, pl. 6 [upper right]. 1824 Distribution: Am 1. Lewinskya elegantula (Schimp. ex Mitt.) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum elegantulum Schimp. ex Mitt., J. Linn. Soc. Bot. 12: 187. 1869 Distribution: Am 6. Lewinskya elongata (Taylor) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum elongatum Taylor, London J. Bot. 5: 45. 1846 Distribution: Am 4. Lewinskya erosa (Lewinsky) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum erosum Lewinsky, J. Hattori Bot. Lab. 72: 32, f. 17. 1992 Distribution: AS 2. Lewinskya fenestrata (Cardot & Thér.) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum fenestratum Cardot & Thér., Proc. Wash. Acad. Sci. 4: 375 310-311, pl. 16, f. 2. 1902 Distribution: Am 1. Lewinskya firma (Venturi) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum firmum Venturi, Nuovo Giorn. Bot. Ital. 4: 15. 1872 Distribution: AFR 2, AFR 4, AS 3. Lewinskya galiciae (F.Lara, Garilleti & Mazimpaka) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrihum galiciae F.Lara, Garilleti & Mazimpaka, Nova Hedwigia 83: 200. 2006 Distribution: AFR 2. Lewinskya graphiomitria (Müll.Hal. ex Beckett) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum graphiomitrium Müll.Hal. ex Beckett, Trans. & Proc. New Zealand Inst. 25: 291. 1893 Distribution: AUSTR 2. Lewinskya hawaiica (Müll.Hal.) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum hawaiicum Müll. Hal., Flora 82: 451. 1896 Distribution: OC. Lewinskya holzingeri (Renauld & Cardot in Holzinger) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum holzingeri Renauld & Cardot in Holzinger, Contr. U.S. Natl. Herb. 3: 270. 1895 Distribution: Am 1. Lewinskya hookeri (Wilson ex Mitt.) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum hookeri Wilson ex Mitt., J. Proc. Linn. Soc. Bot. Suppl. 1: 48. 1859 Distribution: AS 2, AS 3. Lewinskya hookeri var. granulata (Lewinsky) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum hookeri var. granulatum Lewinsky, J. Hattori Bot. Lab. 72: 20. 1992 Distribution: AS 2, AS 3. 376 F. Lara et al. Lewinskya hortensis (Bosw.) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum hortense Bosw., J. Bot. 30: 97. 1892 Distribution: Am 6, AUSTR 1, AUSTR 2. Lewinskya hortoniae (Vitt) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum hortoniae Vitt, Bryologist 82: 2, f. 1-9, 62, 64. 1979 Distribution: Am 2. Lewinskya iberica (F.Lara & Mazimpaka) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum ibericum F.Lara & Mazimpaka, Nova Hedwigia 56: 263, f. 1-3. 1993 Distribution: Eur, AFR 1, AS 5. Lewinskya incana (Müll.Hal.) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum incanum Müll.Hal., Flora 68: 419. 1885 Distribution: Am 6. Lewinskya incrassata (Lewinsky) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum incrassatum Lewinsky, Bryobrothera 2: 67, f. 4-5. 1993 Distribution: OC. Lewinskya incurvomarginata (Lewinsky & van Rooy) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum incurvomarginatum Lewinsky & van Rooy, J. Bryol. 16: 67, f. 1, 4. 1990 Distribution: AFR 4. Lewinskya iwatsukii (Ignatov) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum iwatsukii Ignatov, Arctoa 10: 172. 2001 Distribution: Eur, AS 1, AS 2, AS 3. Lewinskya johnstonii (E.B.Bartram) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum johnstonii E.B.Bartram, Trav. Bryol. 1: 132. 1942 Distribution: Am 6. Lewinskya keeverae (H.A.Crum & L.E.Anderson) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum keeverae H.A.Crum & L.E.Anderson, J. Elisha Mitchell Sci. Soc. 72: 281, f. 3–12. 1956 Distribution: Am 1. Lewinskya laevigata (J.E.Zetterst.) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum laevigatum J.E.Zetterst., Öfvers. Förh. Kongl. Svenska Vetensk.-Akad. 19: 363. 1862 Distribution: Am 1, Eur, AS 1. Lewinskya latimarginata (Lewinsky) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum latimarginatum Lewinsky, Mem. New York Bot. Gard. 45: 331, f. 14-24. 1987 Distribution: Am 4. Lewinskya laxifolia (Wilson ex Mitt.) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum laxifolium Wilson ex Mitt., Hooker’s J. Bot. Kew Gard. Misc. 3: 57. 1851 Distribution: Am 4. Lewinskya leiolecythis (Müll.Hal.) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum leiolecythis Müll. Hal., Nuovo Giorn. Bot. Ital. n.s. 3: 107. 1896 Distribution: AS 2. Lewinskya ludificans (Lewinsky) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum ludificans Lewinsky, Mem. New York Bot. Gard. 45: 347, f. 122-132. 1987 Distribution: Am 6. Lewinskya mandonii (Schimp. ex Hampe) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum mandonii Schimp. ex Hampe, Ann. Sci. Nat. Bot. sér. 5, 4: 332. 1865 Distribution: Am 4. Lewinskya pariata (Mitt.) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum pariatum Mitt., Lewinskya gen. nov. J. Linn. Soc. Bot. 12: 186. 1869 Distribution: Am 4, Am 6. Lewinskya praemorsa (Venturi) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum praemorsum Venturi, Bot. Centralbl. 44: 418. 1890 Distribution: Am 1. Lewinskya psychrophila (Mont.) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum psychrophilum Mont., Ann. Sci. Nat. Bot. sér. 2, 9: 52. 1838 Distribution: Am 6. Lewinskya pulchra (Lewinsky) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum pulchrum Lewinsky, J. Hattori Bot. Lab. 72: 20, f. 1-10. 1992 Distribution: AS 2. Lewinskya pycnophylla (Schimp. in C.MüllerHal.) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum pycnophyllum Schimp. in C.MüllerHal., Syn. Musc. Frond. 1: 709. 1849 Distribution: Am 1, Am 2, Am 4, Am 6. Lewinskya pycnophylla var. verrucosa (Müll.Hal.) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum verrucosum Müll. Hal., Linnaea 42: 359. 1879 Distribution: Am 4, Am 6. Lewinskya pylaisii (Brid.) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum pylaisii Brid., Bryol. Univ. 1(1): 722. 1826 Distribution: Am 1, Eur, AS 1. Lewinskya rupestris (Schleich. ex Schwägr.) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum rupestre Schleich. ex Schwägr., Sp. Musc. Frond. Suppl. 1: 27, pl. 53 [top]. 1816 Distribution: Am 1, Am 2, Am 4, Am 5, Am 6, Eur, AFR 1, AFR 2, AFR 4, AFR 5, AS 1, AS 2, AS 3, AS 5, AUSTR 1, AUSTR 2, OC, ANT. 377 Lewinskya rupestris var. papillosa (Lewinsky) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum rupestre var. papillosum Lewinsky, J. Hattori Bot. Lab. 56: 402. 1984 Distribution: Am 6, AUSTR 2. Lewinskya sainsburyi (Allison) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum sainsburyi Allison, Trans. & Proc. Roy. Soc. New Zealand 77: 280. 1947 Distribution: AUSTR 2. Lewinskya schoddei (Lewinsky) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum schoddei Lewinsky, J. Hattori Bot. Lab. 72: 34, f. 20. 1992 Distribution: AS 4. Lewinskya shawii (Wilson) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum shawii Wilson, Musci Eur. Nov. fasc. 1–2, Monogr. 1. 1. 1864 Distribution: Am 1, Eur, AFR 1, AS 5. Lewinskya sordida (Sull. & Lesq. in Austin) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum sordidum Sull. & Lesq. in Austin, Musci Appalach. 30. 1870 Distribution: Am 1, Eur, AS 1, AS 2, AS 5. Lewinskya spanotricha (Lewinsky) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum spanotrichum Lewinsky, Lindbergia 18: 116, f. 1–2. 1993 [1994] Distribution: Am 4. Lewinskya speciosa (Nees) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum speciosum Nees, Deutschl. Fl. Abt. II Cryptog. 5: 5. 1819 Distribution: Am 1, Eur, AFR 1, AS 1, AS 2, AS 5. 378 F. Lara et al. Lewinskya spjutii (D.H.Norris & Vitt) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum spjutii D.H.Norris & Vitt, Nova Hedwigia 56: 260, f. 1. 1993 Distribution: Am 1. Lewinskya steerei (Lewinsky) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum steerei Lewinsky, Mem. New York Bot. Gard. 45: 357, f. 188-197. 1987 Distribution: Am 4. Lewinskya striata (Hedw.) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum striatum Hedw., Sp. Musc. Frond. 163. 1801 Distribution: Am 1, Eur, AFR 1, AFR 5, AS 1, AS 2, AS 3, AS 5. Lewinskya subulata (Mitt.) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum subulatum Mitt., J. Linn. Soc. Bot. 12: 184. 1869 Distribution: Am 4. Lewinskya taiwanensis (Lewinsky) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum taiwanense Lewinsky, J. Hattori Bot. Lab. 72: 25, f. 12. 1992 Distribution: AS 2. Lewinskya tanganyikae (P. de la Varde) F.Lara, Garilleti & Goffinet, comb. nov. ≡Ulota tanganyikae P. de la Varde, Ark. Bot., n.s. 3: 174, f. 25. 1955 Distribution: AFR 2, AFR 4. Lewinskya tasmanica (Hook.f. & Wilson) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum tasmanicum Hook.f. & Wilson, London J. Bot. 7: 27, pl. 1 c. 1848 Distribution: AUSTR 1, AUSTR 2. Lewinskya tasmanica var. parvitheca (R.Br.bis) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum parvithecum R.Br. bis, Trans. & Proc. New Zealand Inst. 27: 440, pl. 41 f. 1894[1895] Distribution: AUSTR 2. Lewinskya tenuicaulis (Lewinsky) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum tenuicaule Lewinsky, J. Hattori Bot. Lab. 75: 45, f. 1. 1994 Distribution: Am 4. Lewinskya tortidontia (F.Lara, Garilleti & Mazimpaka) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum tortidontium F.Lara, Garilleti & Mazimpaka, Nova Hedwigia 63: 517, f. 1-17. 1996 Distribution: Eur, AFR 1, AS 5. Lewinskya tortifolia (Lewinsky) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum tortifolium Lewinsky, J. Hattori Bot. Lab. 72: 23, f. 11. 1992 Distribution: AS 3. Lewinskya truncatodentata (Müll.Hal.) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum truncatodentatum Müll.Hal., Linnaea 43: 442. 1882 Distribution: Am 6. Lewinskya truncata (Lewinsky & Deguchi) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum truncatum Lewinsky & Deguchi, Bryologist 89: 216, f. 1-17. 1986 [1987] Distribution: Am 6. Lewinskya vicaria (Laz.) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum vicarium Laz., Žurn. Inst. Bot. Vseukrajins›k. Akad. Nauk 26-27: 210. 1938. Distribution: AS 1. Lewinskya vladikavkana (Venturi in Husnot) F.Lara, Garilleti & Goffinet, comb. nov. ≡Orthotrichum vladikavkanum Venturi in Husnot, Muscol. Gall. 167. 1887 Distribution: Eur, AS 1, AS 5. Lewinskya gen. nov. 379 APPENDIX 2 Taxa included in the genus Orthotrichum Hedw., with indication of their known distribution as shown in Fig. 2. Orthotrichum aequatoreum Mitt., J. Linn. Soc. Bot. 12: 184. 1869. Distribution: Am 2, Am 4, AFR 2. Orthotrichum alpestre Hornsch. ex Bruch & Schimp., Bryol. Eur. 3: 75, pl. 213 (fasc. 42. Monogr. Suppl. 1: 1. pl. 1). 1849. Distribution: Am 1, Eur, AS 1, AS 3. Orthotrichum alpestre var. berggrenii (E.B.Bartram) Lewinsky, Bryobrothera 2: 63. 1993. Distribution: OC. Orthotrichum angustifolium Hook.f. & Wilson, London J. Bot. 3: 547. 1844. Distribution: AUSTR 2. Orthotrichum anodon F.Lara, Garilleti & Mazimpaka, Bryologist 109: 188, f. 1–17, 28. 2006. Distribution: Am 1. Orthotrichum anomalum Hedw., Sp. Musc. Frond. 162. 1801. Distribution: Am 1, Am 2, Am3, Eur, AFR 1, AS 1, AS 2, AS 3, AS 5. Orthotrichum assimile Müll.Hal., Syn. Musc. Frond. 1: 704. 1849. Distribution: Am 6, AUSTR 1, AUSTR 2. Orthotrichum atratum Mitt., J. Linn. Soc. Bot. 15: 66. 1876 [1877]. Distribution: AFR 4. Orthotrichum aucklandicum (Vitt) B.Goffinet, Monogr. Syst. Bot. Missouri Bot. Gard. 98: 287. 2004. Distribution: AUSTR 2. Orthotrichum bartramii R.S.Williams, Bryologist 28: 76, pl. 9. 1925. Distribution: Am 1, Am 2. Orthotrichum bistratosum (Schiffn.) J.Guerra, Anales Jard. Bot. Madrid 42: 248. 1985. Distribution: Eur, AS 5. Orthotrichum callistomum Fisch.-Oost. ex Bruch & Schimp., Bryol. Eur. 3: 77, pl. 224 (fasc. 43. Mon. suppl. 2: 1. pl. 3). 1850. Distribution: Eur, AS 2, AS 3, AS 5. Orthotrichum calvum Hook.f. & Wilson, Fl. Nov.-Zel. 2: 80, f. 7. 1854. Distribution: AUSTR 2. Orthotrichum cambrense Bosanquet & F.Lara, Cryptog. Bryol. 33: 330. 2012. Distribution: Eur. Orthotrichum casasianum F.Lara, Garilleti & Mazimpaka, J. Bryol. 21: 47, f. 1–9, 11–12, 15–16. 1999. Distribution: Eur. Orthotrichum columbicum Mitt., J. Proc. Linn. Soc. Bot. 8: 24. 1864. Distribution: Am 1, Eur. Orthotrichum comosum F.Lara, R.Medina & Garilleti, Taxon 62: 1142. 2013. Distribution: Eur, AFR 1, AFR 5. Orthotrichum confusum R.Medina, F.Lara & Garilleti, Taxon 61: 1191. 2012. Distribution: Am 1. Orthotrichum consimile Mitt., J. Linn. Soc. Bot. 8: 24. 1864. Distribution: Am 1. Orthotrichum consobrinum Cardot, Bull. Herb. Boissier sér. 2 8: 336. 1908. Distribution: Eur, AS 1, AS 2, AS 5. Orthotrichum coulteri Mitt., J. Proc. Linn. Soc. Bot. 8: 25. 1864. Distribution: Am 1. Orthotrichum crassifolium Hook.f. & Wilson, London J. Bot. 3: 546. 1844. Distribution: Am 6, AFR 4, AUSTR 2, ANT. Orthotrichum crassifolium subsp. acutum (Müll.Hal.) B.Goffinet, Monogr. Syst. Bot. Missouri Bot. Gard. 98: 288. 2004. Distribution: AFR 4. Orthotrichum crenulatum Mitt., J. Proc. Linn. Soc. Bot. Suppl. 1: 48. 1859. Distribution: Eur, AS 1, AS 3, AS 5. 380 F. Lara et al. Orthotrichum crispifolium Broth., Symb. Sin. 4: 69. 1929. Distribution: AS 3. Orthotrichum cucullatum F.Lara, R.Medina & Garilleti, Taxon 62: 1143. 2013. Distribution: Am 1. Orthotrichum cupulatum Brid., Muscol. Recent. 2: 25. 1801. Distribution: Am 1, Am 2, Eur AFR 1, AFR 5, AS 1, AS 3, AS 5, AUSTR 1, AUSTR 2. Orthotrichum cupulatum var. austroamericanum Lewinsky, Lindbergia 10: 69, f. 1-11. 1984 [1985]. Distribution: Am 4, Am 6. Orthotrichum cupulatum var. austrocupulatum (Dixon & Sainsbury) Lewinsky, J. Hattori Bot. Lab. 56: 433. 1984. Distribution: AUSTR 2. Orthotrichum cupulatum var. fuscum (Venturi) Boulay, Musc. France Mousses 329. 1884. Distribution: Eur. Orthotrichum cupulatum var. riparium Huebener, Muscol. Germ. 381. 1833. Distribution: Am 1, Eur, AS 1, AS 5. Orthotrichum dagestanicum Fedosov & Ignatova, Arctoa 19: 69. 2010. Distribution: Eur, AS 2. Orthotrichum denticulatum Lewinsky, Bot. Tidsskr. 72: 75. 1978. Distribution: AFR 2, AFR 5. Orthotrichum diaphanum Brid., Muscol. Recent. 2: 29. 1801. Distribution: Am 1, Am 2, Am 4, Eur, AFR 1, AFR 2, AFR 4, AFR 5, AS 3, AS 5, OC. Orthotrichum diaphanum var. podocarpi (Müll.Hal.) Lewinsky, Lindbergia 10: 72. 1984 [1985]. Distribution: Am 4, Am 6. Orthotrichum erubescens Müll.Hal., Nuovo Giorn. Bot. Ital. n.s. 4: 260. 1897. Distribution: AS 2. Orthotrichum euryphyllum Venturi, Bot. Centralbl. 44: 417. 1890. Distribution: Am 1. Orthotrichum flowersii Vitt, Bryologist 74: 159, f. 1-8. 1971. Distribution: Am 1. Orthotrichum franciscanum F.Lara, R.Medina & Garilleti, Taxon 62: 1144. 2013. Distribution: Am 1. Orthotrichum freyanum B.Goffinet, W.R. Buck & M.A.Wall, Nova Hedwigia Beih. 131: 6. 2007. Distribution: Am 6. Orthotrichum gigantosporum Lewinsky, Lindbergia 14: 183. 1988 [1989]. Distribution: Am 6. Orthotrichum griffithii Mitt. ex Dixon, J. Bot. 49: 140, 513 f. 2. 1911. Distribution: AS 2, AS 3. Orthotrichum hallii Sull. & Lesq., Icon. Musc. Suppl. 63, pl. 45. 1874. Distribution: Am 1, AS 1. Orthotrichum handiense F.Lara, Garilleti & Mazimpaka, Bryologist 102: 619, f. 1-15. 1999 [2000]. Distribution: AFR 5. Orthotrichum hispanicum F.Lara, Garilleti & Mazimpaka, J. Bryol. 22: 263, f. 1-19. 2000. Distribution: Eur, AS 3, AS 5. Orthotrichum hooglandii E.B.Bartram, Rev. Bryol. Lichénol. 30: 194. 1962. Distribution: AS 4. Orthotrichum ibukiense Toyama, J. Jap. Bot. 14: 620. 1938. Distribution: AS 2. Orthotrichum inclinatum Müll.Hal., Flora 68: 419. 1885. Distribution: Am 6. Orthotrichum jetteae B.H.Allen, Monogr. Syst. Bot. Missouri Bot. Gard. 90: 639. 2002. Distribution: AS 2. Orthotrichum karoo F.Lara, Garilleti & Mazimpaka, Bryologist 112: 195. 2009. Distribution: AFR 4. Orthotrichum kellmanii D.H.Norris, Shevock & B.Goffinet, Bryologist 107: 210, f. 2. 2004. Distribution: Am 1. Lewinskya gen. nov. Orthotrichum laxum Lewinsky, Lindbergia 24: 29, f. 1-2. 1999. Distribution: AS 2. Orthotrichum lescurii Austin, Musci Appalach. 163. 1870. Distribution: Am 1. Orthotrichum liliputanum Broth. in Herzog, Biblioth. Bot. 87: 63. 1916. Distribution: Am 4. Orthotrichum macrocephalum F.Lara, Garilleti & Mazimpaka, Bryologist 97: 402, f. 1-20. 1994. Distribution: Eur, AFR 1, AS 5. Orthotrichum mazimpakanum Garilleti & F.Lara, Bryologist 114: 351. 2011. Distribution: Am 1. Orthotrichum meyenianum Hampe, Syn. Musc. Frond. 2: 639. 1851. Distribution: AS 4. Orthotrichum microcarpum De Not., Erb. Crittog. Ital. 19: 915. 1863. Distribution: Eur, AS 5. Orthotrichum moravicum Plášek & Sawicki, Bryologist 112: 329. 2009. Distribution: Eur. Orthotrichum norrisii F.Lara, R.Medina & Garilleti, Bryologist 111: 670, f. 1-2. 2008. Distribution: Am 1. Orthotrichum notabile Lewinsky, Lindbergia 20: 102. 1995 [1996]. Distribution: AS 2. Orthotrichum ohioense Sull. & Lesq., Musci Appalach. 169. 1870. Distribution: Am 1. Orthotrichum pallens Bruch ex Brid., Bryol. Univ. 1: 788. 1827. Distribution: Am 1, Am 2, Am 4, Eur, AFR 1, AFR 5, AS 1, AS 3, AS 5. Orthotrichum pallens var. crenulatum Vitt, Bryologist 74: 161, f. 9-12. 1971. Distribution: Am 1. Orthotrichum pallens var. johnseniae Vitt, Bryologist 74: 162, f. 13-16. 1971. Distribution: Am 1. Orthotrichum pamiricum Plášek & Sawicki, Turk. J. Bot. 38: 755. 2014. Distribution: AS 3. 381 Orthotrichum parvulum Mitt., J. Linn. Soc. Bot. 8: 25. 1864. Distribution: Am 1. Orthotrichum patens Bruch & Brid., Bryol. Univ. 1: 787. 1827. Distribution: Eur, AS 5. Orthotrichum pellucidum Lindb., Öfvers. Förh. Kongl. Svenska Vetensk.-Akad. 23: 549. 1866 [1867]. Distribution: Am 1, Eur, AS 1. Orthotrichum penicillatum Mitt., J. Linn. Soc. Bot. 12: 185. 1869. Distribution: Am 4. Orthotrichum perexiguum Dusén ex Lewinsky, Lindbergia 10: 89, f. 142. 1984 [1985]. Distribution: Am 6. Orthotrichum persimile F.Lara, R.Medina & Garilleti, Taxon 61: 1193. 2012. Distribution: Am 1. Orthotrichum philibertii Venturi, Rev. Bryol. 5: 45. 1878. Distribution: Eur, AFR 1, AS 5. Orthotrichum pilosissimum R.Medina, F.Lara & Garilleti, Bryologist 114: 317. 2011. Distribution: Am 1. Orthotrichum pulchellum Brunt., Bot. Guide Northumberland & Durham 2: 23. 1807. Distribution: Am 1, Eur. Orthotrichum pumilum Sw. ex anon., Monthly Rev. 34: 538. 1801. Distribution: Am 1, Eur, AFR 1, AFR 5, AS 1, AS 2, AS 3, AS 5. Orthotrichum pungens Mitt., J. Linn. Soc. Bot. 12: 184. 1869. Distribution: Am 4. Orthotrichum pusillum Mitt., J. Linn. Soc. Bot. 8: 25. 1864. Distribution: Am 1. Orthotrichum revolutum Müll.Hal., Nuovo Giorn. Bot. Ital. n.s. 4: 261. 1897. Distribution: AS 2. Orthotrichum rivulare Turner, Muscol. Hibern. Spic. 96, pl. 8. 1804. Distribution: Am 1, Eur, AS 1. Orthotrichum rogeri Brid., Muscol. Recent. Suppl. 2: 9. 1812. Distribution: Eur, AS 5. 382 F. Lara et al. Orthotrichum scanicum Grönvall, Bidr. Känned. Nord. Orthotrichum & Ulota 13. 1885. Distribution: Eur, AFR 1, AS 3, AS 5. Orthotrichum schimperi Hammar, Monogr. Orthotrich. Ulot. Suec. 9. 1852. Distribution: Am 1, Eur, AFR 1, AFR 5, AS 5. Orthotrichum schofieldii (B.C.Tan & Y.Jia) B.H.Allen, Monogr. Syst. Bot. Missouri Bot. Gard. 90: 639. 2002. Distribution: AS 2. Orthotrichum sharpii H.Rob., Bryologist 67: 449, f. 16-22. 1964. Distribution: Am 2. Orthotrichum shevockii Lewinsky & D.H.Norris, Bryologist 101: 435. 1998. Distribution: Am 1. Orthotrichum sinuosum Lewinsky, J. Hattori Bot. Lab. 72: 73, f. 42. 1992. Distribution: AS 2. Orthotrichum spiculatum F.Lara, Garilleti & Mazimpaka, Bot. J. Linn. Soc. 155: 477. 2007. Distribution: Am 6. Orthotrichum sprucei Mont., London J. Bot. 4: 186. 1845. Distribution: Eur, AS 1, AS 5. Orthotrichum stellatum Brid., Bryol. Univ. 1: 274. 1826. Distribution: Am 1, Eur, AS 5. Orthotrichum stramineum Hornsch. ex Brid., Bryol. Univ. 1: 789. 1827. Distribution: Am 1, Eur, AFR 1, AS 3, AS 5. Orthotrichum strangulatum P.Beauv., Prodr. Aethéogam. 81. 1805. Distribution: Am 1. Orthotrichum subexsertum Schimp. ex Müll.Hal., Bot. Zeitung (Berlin) 16: 164. 1858. Distribution: AFR 2, AFR 4. Orthotrichum subpumilum E.B.Bartram ex Lewinsky J. Hattori Bot. Lab. 72: 59, f. 34 1992. Distribution: AS 2. Orthotrichum tenellum Bruch ex Brid., Bryol. Univ. 1: 786. 1827. Distribution: Eur, AFR 1, AFR 5, AS 5. Orthotrichum trachymitrium Mitt., J. Linn. Soc. Bot. 12: 183. 1869. Distribution: Am 4. Orthotrichum transvaaliense Rehmann ex Sim, Trans. Roy. Soc. South Africa 15: 276. 1926. Distribution: AFR 4. Orthotrichum tristriatum Lewinsky, Lindbergia 15: 70. 1989 [1990]. Distribution: Am 6. Orthotrichum underwoodii F.Lara, Garilleti & Mazimpaka, Bryologist 104: 268. 2001. Distribution: Am 1. Orthotrichum urnaceum Müll.Hal., Trudy Imp. S.-Peterburgsk. Bot. Sada 10: 259. 1887. Distribution: Eur. Orthotrichum urnigerum Myrin, Coroll. Fl. Upsal. 71. 1833. Distribution: Eur, AS 3. Orthotrichum vermiferum Lewinsky, Lindbergia 24: 33, f. 1. 1999. Distribution: AS 2. Orthotrichum vittii F.Lara, Garilleti & Mazimpaka, Bryologist 102: 53, f. 1-22. 1999. Distribution: Eur, AFR 1, AS.