Phytotaxa 220 (2): 127–142 www.mapress.com/phytotaxa/ Copyright © 2015 Magnolia Press ISSN 1179-3155 (print edition) Article PHYTOTAXA ISSN 1179-3163 (online edition) http://dx.doi.org/10.11646/phytotaxa.220.2.3 Notes on Early Land Plants Today. 72. Infrageneric classification and new combinations, new names, new synonyms in Frullania (Marchantiophyta) JÖRN HENTSCHEL1, MATT VON KONRAT2, LARS SÖDERSTRÖM3*, ANDERS HAGBORG2, JUAN LARRAÍN2, PHIANGPHAK SUKKHARAK4, JAIME URIBE5 & LI ZHANG6 1 Department of Systematic Botany with Herbarium Haussknecht and Botanical Garden, Friedrich Schiller University, Fürstengraben 1, 07737 Jena, Germany; j.hentschel@uni-jena.de 2 Gantz Family Collections Center, The Field Museum, 1400 South Lake Shore Drive, Chicago, IL 60605–2496, USA; mvonkonrat@ fieldmuseum.org, hagborg@pobox.com, jlarrain@fieldmuseum.org 3 Department of Biology, Norwegian University of Science and Technology, N-7491 Trondheim, Norway; lars.soderstrom@ntnu.no 4 Department of Biology, Faculty of Science, Burapha University, Mueang, 20131 Chonburi, Thailand; ppsukk@gmail.com 5 Instituto de Ciencias Naturales. Universidad Nacional de Colombia. Apartado 7495, Bogotá D.C., Colombia; juribem@unal.edu.co 6 Shenzhen Key Laboratory of Southern Subtropical Plant Diversity, Fairylake Botanical Garden, 160 Xianhu Rd., Liantang, Shenzhen 518004, Guangdong, China; zhangli@scib.ac.cn *corresponding author Abstract The liverwort family Frullaniaceae is circumscribed to include the single genus, Frullania, and has a complex and confusing taxonomical history. An overview is provided for the infrageneric classification adopted for the forthcoming worldwide checklist based on recent morphological and molecular studies. The genus is preliminarily subdivided into eleven subgenera, F. subg. Chonanthelia, F. subg. Diastaloba, F. subg. Diversitextae, F. subg. Frullania, F. subg. Homotropantha, F. subg. Mammillosae, F. subg. Meteoriopsis, F. subg. Microfrullania, F. subg. Saccophora, F. subg. Steerea, and F. subg. Thyopsiella, as well as several sections. However, some of the subgeneric divisions are only doubtfully recognized because there is still insufficient knowledge to arrive at a decision regarding their systematic status and placement. The taxa may be either originally not well described, not restudied recently or molecular analyses point to a more complex internal structure of the respective group. Fourteen new synonyms, eight new combinations, five new names, eight lectotypifications, and one new variety are provided. Introduction Frullania Raddi (1818: 9) is worldwide in distribution and is one of the largest and taxonomically most complex genera of leafy liverworts with more than 2,000 published names ascribed to the genus (von Konrat et al. 2010). The current contribution follows the series of Early Land Plant Today Notes and is similar in content, structure and format to Söderström et al. (2015) that likewise deals with Plagiochila (Dumortier 1831: 42) Dumortier (1835: 14). The current paper provides only an overview of the infrageneric classification adopted as part of the forthcoming worldwide checklist of liverworts (Söderström et al., in press). We do not attempt here to provide a full synopsis of the over 30 sections and subsections that have been historically described. The paper also includes several new synonyms, new combinations, new names and lectotypifications. Schusterella Hattori et al. (1972: 330), Amphijubula Schuster (1970: 298) and Steerea Hattori & Kamimura (1971: 429) have been considered segregate genera of Frullania, but are now considered synonyms of this genus (Engel 1978, Hattori & Mizutani 1982, Stotler & Crandall-Stotler 1987, Schuster 1992, von Konrat et al. 2006, 2011b, Hentschel et al. 2009). Neohattoria Kamimura (1962: 218) has also historically been placed in Frullania (Hattori & Mizutani 1982, Stotler & Crandall-Stotler 1987). However, the authors are actively investigating recently collected material identified as Neohattoria using molecular tools to test this relationship. Whereas the circumscription of Frullaniaceae can be deemed sufficiently clarified, the subgeneric classification of Frullania is still a matter of much controversy (e.g., Hentschel et al. 2009, Uribe 2011). The content of the paper is in two parts. First, an infrageneric classification is provided, including only those subgenera and sections that are recognized in the forthcoming liverwort worldwide checklist (Söderström et al., in Accepted by Christine Cargill: 6 May 2015; published: 20 Jul. 2015 Licensed under a Creative Commons Attribution License http://creativecommons.org/licenses/by/3.0 127 press). The second part focuses on new synonyms or new names at the species level and below associated with miscellaneous names. Both parts have novel nomenclatural elements. The format of this note follows Söderström et al. (2012). Fourteen new synonyms, eight new combinations, five new names, eight lectotypifications, and one new variety are provided. Infrageneric structure of Frullania Over the last five decades, there have been abundant changes and modifications to the subgeneric classification of Frullania, in part due to the establishment of supposedly allied genera, and the contractions and expansions of generic, subgeneric and sectional concepts. Consequently, this has led to a complex and confusing classification. Attempts to circumscribe subgenera and sections within the whole distributional range of the genus have been very limited (Hattori 1980b, Yuzawa 1991, Hentschel et al. 2009). There have been various proposals for a subdivision of Frullania into natural species groups (e.g., Spruce 1884, Verdoorn 1930, Vanden Berghen 1976, Hattori 1976, Stotler & CrandallStotler 1987, Yuzawa 1991, Schuster 1992). As a result, these authors established more than fifteen subgenera and over thirty sections and subsections based on morphology. Recently, Hentschel et al. (2009) presented the most comprehensive molecular phylogeny of Frullania to date. According to the molecular data several robust lineages within Frullania could be assigned to morphologically circumscribed subgenera and provide evidence for the monophyly of a suite of currently accepted taxa. The results point to a general congruence of morphological subgeneric classifications and the molecular topologies. However, in some cases previous subgeneric assignment for a few taxa, e.g., F. subg. Diastaloba Spruce (1884: 55) and F. subg. Microfrullania (Schuster 1970: 280) Schuster (1992: 34), based on morphological evidence is not supported by the molecular data and further modifications and refinements of the morphology-based subgeneric concepts are necessary to arrive at a more comprehensible and natural subdivision. Although the analysis of Hentschel et al. (2009) is based on a rather large dataset in terms of the number of accessions investigated as well as the number of molecular markers utilized, the taxon sampling is still inadequate to revise all supraspecific taxa, especially the many published sections and subsections. Hentschel et al. (2009) have already provided strong evidence that several sections and subsections are polyphyletic, e.g., F. sect. Ornithocephalae Verdoorn (1930: 50), F. sect. Integristipulae Verdoorn (1930: 49), and F. subsect. Ericoides Schuster (1985: 371) and many are suspected synonyms, e.g., F. sect. Coniferae Schuster (1985: 372). Due to the fact that several key taxa and type specimens, e.g., F. ciliata Lindenb. et Gottsche in Gottsche et al. (1847: 775), F. clemensiana Verdoorn (1932: 493), F. diversitexta Stephani (1897: 89), F. errans Verdoorn (1930: 59), F. junghuhniana Gottsche in Gottsche et al. (1845: 444), F. mammillosa Hattori (1977 [1978]: 424), F. subtilissima (Nees ex Montagne 1840: 333) Lindenb. in Gottsche et al. (1845: 443), F. ternatensis Gottsche in Gottsche et al. (1846: 465), and F. vaginata (Swartz 1781: 35) Dumortier (1835: 13), have not been studied yet, we have adopted a rather conservative approach for the circumscription of the subgenera and sections included in the forthcoming worldwide checklist of hornworts and liverworts (Söderström et al., in press). The following annotated synopsis of infrageneric ranks basically reflects a consensus of an informal Frullania Working Group that met at the Field Museum of Natural History in Chicago in 2009 and 2012. The subgenera outlined below reflect the infrageneric classification adopted for the forthcoming worldwide checklist. Discussion is provided to support that classification, which is outside the auspices of the checklist. References to articles in the nomenclature code follows McNeill et al. (2012). Frullania subg. Chonanthelia Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 8, 1884 (Spruce 1884). Lectotype (Hattori 1982: 261):—Frullania gibbosa Nees in Gottsche et al. (1845: 411). Note:—Frullania subg. Chonanthelia was monographed by Yuzawa (1991), who proposed several new infrageneric entities. Besides a few exceptions, most of his categories are natural and reflected in the topologies presented by Hentschel et al. (2009). However, due to the placement of F. gibbosa in the respective analyses and the use of F. sect. Chonanthelia as a putative autonym of F. subg. Chonanthelia without formal description or combining it (e.g., Yuzawa 1991) several nomenclatural refinements are necessary. For the forthcoming worldwide checklist we adopt three sections, namely F. sect. Cladocarpicae, F. sect. Chonanthelia and F. sect. Pluricarinatae. 128 • Phytotaxa 220 (2) © 2015 Magnolia Press HENTSCHEL ET AL. Frullania (subg. Chonanthelia) sect. Cladocarpicae Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 11, 1884 (Spruce 1884). Lectotype (Yuzawa 1991: 245):—Frullania brachyclada Spruce (1884: 11), nom. illeg. (ICN Art. 53.1, non Lehm. 1844) [≡ Frullania tunguraguana Clark & Frye (1952: 133)]. = Frullania sect. Acrocarpicae Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 11, 1884 (Spruce 1884), syn. nov. Lectotype (here designated):—Frullania arecae (Sprengel 1821: 99) Gottsche (1863: 236) [= Frullania obscura (Swartz 1806: 1869) Dumortier (1835: 13), cf. Gradstein (2012)]. Note:—This lectotypification makes the section synonymous with F. sect. Cladocarpicae (cf. Hentschel et al. 2009). Frullania (subg. Chonanthelia) sect. Chonanthelia Yuzawa ex Hentschel et von Konrat, sect. nov. Holotype (Hattori 1982: 161):—Frullania gibbosa Nees in Gottsche et al. (1845: 411). Note:—Yuzawa (1991) and others have used the name F. sect. Chonanthelia as a putative autonym below F. subg. Chonanthelia. However, the establishment of any subdivision of a genus under a subdivision that does not include the type of the genus, does not create an autonym (cf. ICN Art. 22.1). We here validate the name by referring to Yuzawa’s English description of F. sect. Chonanthelia (Yuzawa 1991: 200) and the selection of a holotype made here. Frullania (subg. Chonanthelia) sect. Pluricarinatae (Yuzawa, Mues et S.Hatt.) Hentschel et von Konrat, comb. nov. Basionym:—Frullania ser. Pluricarinatae Yuzawa, Mues et S.Hatt., J. Hattori Bot. Lab. 63: 428, 1987 (Yuzawa et al. 1987). Type (ICN Art. 22.6):—Frullania pluricarinata Gottsche (1864: 168). Note:—The new section is created to host the taxa Hentschel et al. (2009) erroneously referred to F. sect. Chonanthelia as we have not been able to find any existing sectional name that can be applied. Frullania subg. Diastaloba Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 55, 1884 (Spruce 1884). Lectotype (Hattori 1982: 235):—Frullania subtilissima (Nees ex Montagne 1840: 333) Lindenb. in Gottsche et al. (1845: 443). Note:—This subgenus is characterized basically by the shape, position and orientation of the leaf lobule (Spruce 1884). According to the molecular data at hand, the subgenus includes at least four genetically distinct lineages, provisionally named “Diastaloba I–IV” (Hentschel et al. 2009). Furthermore, the Diastaloba-like habit reoccurs in F. subg. Microfrullania, which makes the discrimination between both subgenera even more challenging (Schuster 1992). Although much progress has been made towards a refined circumscription of F. subg. Microfrullania (e.g., von Konrat et al. 2006, Hentschel et al. 2009, von Konrat et al. 2010, 2011a, 2012, 2013), the taxonomic problems concerning the classification of F. subg. Diastaloba have not yet been satisfactorily solved. Frullania subg. Diastaloba was typified by Hattori (1982) with F. subtilissima, a rare and enigmatic species from Guyana. Based on preliminary observations, including the study of relevant type specimens, a morphological characterization of the corresponding lineages seems possible. A taxonomical revision is currently underway and will lead to an extensive reclassification of this by far most complex subgenus. We hereby recognize the four informal clades that correspond to “Diastaloba I, II, III and IV” and have attempted to place species within these units for the forthcoming worldwide liverwort checklist (Söderström et al., in press). Not all purported taxa of these groups have been investigated using molecular tools. However, attempted groupings have been made based on morphological similarity and purported relationships described by previous workers, especially the many publications by S. Hattori on the genus. Since we do not yet know in which of the four lineages the type, F. subtilissima, belongs, we refrain from giving them formal sectional names. Below a brief discussion of these four natural units is presented. Diastaloba I corresponds to three well supported clades which in Hentschel et al. (2009) included the type species of F. sect. Graciles Verdoorn (1930: 110) [type F. gracilis (Reinwardt et al. 1824: 221) Nees in Gottsche et al. (1845: 452)], F. sect. Serratae Verdoorn (1930: 85) [type F. serrata Gottsche in Gottsche et al. (1845: 453)] and F. sect. Inconditum von Konrat et al. (2010: 480) [type F. hodgsoniae von Konrat et al. (2010: 492)] (as Rostratae). Although not included in any molecular study, F. sect. Vaginatae Verdoorn (1930: 141) [type F. vaginata (Swartz 1781: 35) Dumortier (1835: 13)], F. subsect. Ternatenses Hattori (1976: 466) [type F. ternatensis Gottsche in Gottsche et al. (1846: 465)], F. sect. Sinuatae Schuster (1985: 371) [type F. sinuata Sande Lacoste (1854: 424)], F. sect. Pycnophyllae Schuster (1985: 370) [type F. pycnophylla Hattori (1973a: 60) (= F. curvistipula Stephani (1911: 541)] and F. sect. Curvistipulae Schuster (1991: 144) [type F. curvistipula] seem to belong here. INFRAGENERIC CLASSIFICATION OF FRULLANIA Phytotaxa 220 (2) © 2015 Magnolia Press • 129 Diastaloba II includes the type species of one section in Hentschel et al. (2009), F. sect. Lucidae Verdoorn (1930: 125) [type F. repandistipula Sande Lacoste (1854 [1855]: 422)]. Although not included in any molecular study, F. sect. Ocellatae Schuster (1985: 371) [type F. ocellata Hattori & Kamimura (1973: 531)] seems to belong here. Diastaloba III comprises two species from Madagascar, F. usambarana Schiffn. ex Stephani (1894: 160) and F. grossiclava Stephani (1910: 384); neither of these represent a type of any subgeneric taxon. We were not able to find any existing name that would be possible to apply on this group. Diastaloba IV corresponds to a group of distinctive plants that have strong microphyllous branches and long, narrow lobules oblique to the stem, e.g., F. hypoleuca Nees in Gottsche et al. (1843: 471) and F. obcordata (Lehmann 1834: 51) Lehm. et Lindenb. in Gottsche et al. (1845: 447) [= F. caulisequa (Nees 1833: 373) Montagne (1839: 51)]. As with the previous group, none of the taxa included by Hentschel et al. (2009) in this group is a type of any subgeneric taxon, and we have no existing name that we think may be applicable to it. Frullania subg. Diversitextae (Kamim.) S.Hatt., J. Hattori Bot. Lab. 59: 154, 1985 (Hattori & Lin 1985). Basionym:— Frullania subsect. Diversitextae Kamim., J. Hattori Bot. Lab. 24: 80, 1961 (Kamimura 1961). Type (ICN Art. 22.6):— Frullania diversitexta Stephani (1897: 89). Note:—We treat F. subg. Diversitextae as circumscribed by Hattori & Lin (1985) as monotypic and including only F. diversitexta. Kamimura (1961) first diagnosed F. subg. Diversitextae as a subsection of subg. F. sect. Diastaloba. Hattori & Lin (1985) raised it to subgeneric rank recognizing its somewhat chimeric appearance; the combination of tuberculate perianths—a feature unique to F. subg. Frullania, and its otherwise Diastaloba-like habit. This species has not yet been included in a molecular study, and its phylogenetic position therefore remains unclear. Judging from a morphological point of view, especially the development of the first branch leaf and the first branch underleaf as well as the shape of the stylus, there are certain affinities with the “Diastaloba I”-clade of Hentschel et al. (2009). Frullania subg. Frullania, Trans. & Proc. Bot. Soc. Edinburgh 15: 8, 1884 (Spruce 1884). Lectotype (Evans 1918: 468):—Frullania major Raddi (1818: 9), nom. illeg. (ICN Art. 52.1; earlier name included) ≡ Frullania dilatata (Linnaeus 1753: 1133) Dumortier (1835: 13). Note:—Raddi (1818) included two species in his new genus Frullania, F. major (Raddi 1818: 9) and F. minor Raddi (1818: 10), both being illegitimate as they included earlier names. Raddi had Linnaeus’ (1753: 1134) synonymy confused treating the larger species [F. tamarisci (Linnaeus 1753) Dumortier (1835: 13)] as F. minor and the smaller species [F. dilatata (Linnaeus 1753: 1133) Dumortier (1835: 13)] as F. major. Evans (1918: 468) was the first to select a type. He selected F. dilatata which was included as a synonym of F. major in Raddi (1818). It is often argued that Evans used the American Code and thus that the typification was mechanical, which is not allowed by the International Code (cf. ICN Art. 10.5(b)). However, in this case he did not choose the species first mentioned by Raddi, but a synonym of it, and thus it cannot be a mechanical selection. Frye & Clark (1947: 736) lectotypified the genus with F. tamarisci. However, their synonymy included F. major, the only element of their synonymy possible to choose as lectotype. Thus, the lectotype of Frullania (and also the autonym) is F. major which has long been treated as a synonym of F. tamarisci (e.g., Frye & Clark 1947). Also, F. subg. Frullania was used for what is now F. subg. Thyopsiella while what we currently accept as F. subg. Frullania used to be named F. subg. Trachycolea Spruce (1884: 31). Frullania subg. Frullania accommodates about 250 accepted species and represents one of the most specious subgenera of Frullania, especially in the extratropical regions, and includes a large number of polymorphic species (Hentschel et al. 2009). The further subdivision in a number of sections and subsections as well as the occurrence of many rather polymorphic species makes this subgenus taxonomically fairly difficult (Schuster 1992, Hentschel et al. 2009). Hence, it is not surprising that the group remains unrevised on a global scale. Yuzawa (1991) regarded species of F. subg. Frullania as being closely related to species of F. subg. Chonanthelia, but the lineage proved to be distinct based on the morphological and molecular data at hand (e.g., Schuster 1992, Hentschel et al. 2009). Although the analysis of Hentschel et al. (2009) is based on a rather large dataset, the taxon sampling is still inadequate to revise all sections and subsections, but it is obvious that the sectional and subsectional assignment for several taxa based on morphological evidence is not supported by the molecular data. Key taxa to be investigated include the missing type species of supraspecific entities, e.g., F. errans Verdoorn (1930: 59), F. monocera (Hooker & Taylor 1845: 89) Gottsche et al. (1845: 418) and F. ornithocephala (Reinwardt et al. 1824: 216) Nees in Gottsche et al. 1845: 425, and more accessions of genetically complex species, like F. ericoides (Nees 1833: 346) Montagne (1839: 51), F. nepalensis (Sprengel 1827: 324) Lehm. in Gottsche et al. (1845: 422), and F. fugax (Hooker & Taylor 1845: 87) Gottsche et al. 130 • Phytotaxa 220 (2) © 2015 Magnolia Press HENTSCHEL ET AL. (1845: 445). Despite strong evidence that some sections and subsections likely will become synonyms (Hentschel et al. 2009), proposing formal synonymy now would only contribute to the already inflated sectional and subsectional classification due to some paraphyletic species such as F. ericoides Nees (1833: 346) Montagne (1839: 51). However, a suite of subclades can be recognized at present and are therefore adopted for the forthcoming worldwide checklist. Frullania (subg. Frullania) sect. Acutilobae Verd., Ann. Bryol., Suppl. 1: 44, 1930 (Verdoorn 1930). Lectotype (here designated):—Frullania monocera (Hooker & Taylor 1845: 89) Gottsche et al. (1845: 418). Note:—Schuster (1992) noted that the form of gynoecial axes in this section should be investigated citing that typical members, e.g. F. allanii Hodgson (1949: 371), bear gynoecia on unspecialised leafy axes that again branch, in contrast to the simple gynoecial branches of F. bonincola Hattori (1978: 551), which is purportedly allied to the section. Kamimura (1961) noted that F. sect. Acutilobae is a weak taxon and that the acute rostrum of the leaf-lobules was the only significant difference between F. sect. Acutilobae and F. sect. Trachycolea. However, Hattori (1983) stated that F. sect. Acutilobae was unique with the long piliferous beaks of the leaf-lobules. The molecular analysis by Hentschel et al. (2009) supports this classification. Frullania (subg. Frullania) sect. Australes Verd., Ann. Bryol., Suppl. 1: 58, 1930 (Verdoorn 1930). Lectotype (Hattori 1976: 463):—Frullania errans Verdoorn (1930: 59). Note:—Verdoorn (1930) first established F. sect. Australes as a section within F. subg. Frullania (as F. subg. Trachycolea). Later Hattori (1976) made a new combination raising the rank of F. sect. Australes to subgenus. Hentschel et al. (2009) showed molecular evidence that F. subg. Australes forms a robust subclade and is nested within a polytomous topology with several sections of F. subg. Frullania. We therefore prefer Verdoorn’s (1930) treatment of F. sect. Australes as a section of F. subg. Frullania. The majority of species of this lineage is distributed in Eastern Asia and Australasia, but according to the Hentschel et al. (2009) phylogeny the neotropical F. glomerata and African F. obscurifolia Mitten (1879: 400) also belong here. Frullania (subg. Frullania) sect. Frullania, Trans. & Proc. Bot. Soc. Edinburgh 15: 11, 1884 (Spruce 1884). Frullania (subg. Frullania) sect. Irregulares E.A.Hodgs. ex S.Hatt., J. Hattori Bot. Lab. 54: 143, 1983 (Hattori 1983). Holotype (Hattori 1983:143):—Frullania deplanata Mitten (1855: 161). Note:—This section was originally proposed by Hodgson (1949), but was invalid because a Latin diagnosis was omitted. Later, Hattori (1983: 143) validated the section. Hodgson (1949) included only a single species, F. deplanata. Frullania sect. Irregulares as currently circumscribed includes solely species from New Zealand (Hentschel et al. 2009). Frullania (subg. Frullania) sect. Planae R.M.Schust., Phytologia 57: 372, 1985 (Schuster 1985). Type (ICN Art. 22.6):—Frullania plana Sullivant (1849: 175). Note:—This section is seemingly monotypic (Schuster 1985, 1992). Schuster (1992) provided a detailed analysis comparing and distinguishing it from F. sect. Frullania (as F. sect. Trachycolea). Frullania subg. Homotropantha Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 35, 1884 (Spruce 1884). Lectotype (Kamimura 1961: 57):—Frullania replicata (Nees 1833: 369) Montagne (1840: 333) [= Frullania nodulosa (Reinwardt et al. 1824: 217) Nees in Gottsche et al. (1845: 433), cf. Kamimura 1961]. Note:—This subgenus was thoroughly revised by Hattori (1980b) and is here circumscribed to include three sections, F. sect. Remotilobae, F. sect. Fallaces and F. sect. Nodulosae. The latter two were incorporated in the molecular study of Hentschel et al. (2009) and resolved as monophyletic in a well supported sister relationship. Frullania (subg. Homotropantha) sect. Fallaces Verd., Rev. Bryol. Lichénol. 1: 112, 1928 (Verdoorn 1928a). Type (ICN Art. 22.6):—Frullania fallax Gottsche in Gottsche et al. (1845: 432). Note:—The integrity of F. sect. Fallaces seems to be supported by both the morphological evidence described by INFRAGENERIC CLASSIFICATION OF FRULLANIA Phytotaxa 220 (2) © 2015 Magnolia Press • 131 (Hattori 1980b) as well as the molecular analysis presented by Hentschel et al. (2009), although the type species has not yet been included in any molecular systematic study. Frullania fallax is regarded as most closely related to F. intermedia which is well known for its pronounced morphological plasticity (Hattori 1980b) and is possibly represented by a species complex. Frullania (subg. Homotropantha) sect. Nodulosae Verd., Rev. Bryol. Lichénol. 1: 116, 1928 (Verdoorn 1928a). Type (ICN Art. 22.6):—Frullania nodulosa (Reinwardt et al. 1824: 217) Nees in Gottsche et al. (1845: 433). Note:—Hattori (1980b) has used the name F. sect. Homotropantha as a putative autonym of F. subg. Homotropantha and included F. sect. Nodulosae in its synonymy. However, the establishment of any subdivision of a genus under a subdivision that does not include the type of the genus, does not create an autonym (cf. ICN Art. 22.1). Thus the oldest available name must be used. Frullania (subg. Homotropantha) sect. Remotilobae Verd., Rev. Bryol. Lichénol. 1: 119, 1928 (Verdoorn 1928a). Type (ICN Art. 22.6):—Frullania remotiloba Stephani (1894: 152). Note:—This section was established by Verdoorn (1928a) to accommodate F. remotiloba. Later Verdoorn (1930) added F. heteromorpha Schiffner (1890: 38). Some doubts were expressed by Hattori (1980b) regarding the sectional placement of F. heteromorpha, because the species shows some similarities with members of F. sect. Fallaces. According to Hattori (1980b, 1982) F. remotiloba may be considered as a ancestral species of F. subg. Homotropantha closely related to F. subg. Diastaloba. None of the respective species have been available for a molecular phylogenetic analysis yet. Frullania subg. Mammillosae S.Hatt., J. Hattori Bot. Lab. 60: 226, 1986 (Hattori 1986a). Type (ICN Art. 22.6):— Frullania mammillosa Hattori (1977 [1978]: 424). Note:—Representatives of this subgenus exhibit a Diastaloba-like habit, but are otherwise characterized by the development of stem-leaves with apiculate or pilose apices (Hattori 1986a). The phylogenetic affiliation of this subgenus remains unclear, because a taxonomic revision of F. subg. Diastaloba is still lacking. Until now, no species currently placed in F. subg. Mammillosae have been investigated by molecular methods. Frullania subg. Meteoriopsis Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 37, 1884 (Spruce 1884). Lectotype (Uribe & Gradstein 2003):—Frullania peruviana Gottsche in Gottsche et al. (1846: 465). Note:—The taxonomic history of this subgenus is fairly complicated and its current circumscription is strongly influenced by recent taxonomical efforts in F. subg. Diastaloba and F. subg. Thyopsiella. Spruce (1884) established F. subg. Thyopsiella to accommodate species with a prostrate to ascending habit while species with a pendent growth were assigned to F. subg. Meteoriopsis Spruce. Hattori (1972a) doubted the taxonomic relevance of the habit and synonymized F. subg. Meteoriopsis with F. subg. Thyopsiella (using the name F. subg. Frullania, cf. above). Spruce’s (1884) treatment is supported by both the morphological evidence presented by Uribe (2008) as well as the molecular study by Hentschel et al. (2009) where the types of the two subgenera, F. peruviana and F. tamarisci, are located in different clades. In the present circumscription F. subg. Meteoriopsis includes species with a pendent (F. sects. Intumescentes and Meteoriopsis), and a prostrate growth form (F. sect. Obtusilobae). Ongoing taxonomic studies have revealed that several taxa formerly assigned to F. subg. Meteoriopsis as circumscribed here, are indeed members of the specious “Diastaloba I”-clade of Hentschel et al. (2009) and closely related to F. apiculata (Reinwardt et al. 1824: 222) Nees in Gottsche et al. (1845: 452), F. serrata Gottsche in Gottsche et al. (1845: 453) and F. subdentata Stephani (1911: 545). Currently no extra-neotropical member of F. sects. Intumescentes and Meteoriopsis is known because most of the Asiatic species formerly placed in F. subg. Meteoriopsis exclusive of F. sect. Obtusilobae are presumably more closely related to F. subg. Diastaloba. Frullania (subg. Meteoriopsis) sect. Meteoriopsis Uribe, von Konrat et Hentschel., sect. nov. Type:—Frullania peruviana Gottsche in Gottsche et al. 1846: 465). 132 • Phytotaxa 220 (2) © 2015 Magnolia Press HENTSCHEL ET AL. Plants with deeply cordate leaves, with two large auricles on both sides of the base, which are strongly convoluted around the stem. Note:—Frullania sect. Meteoriopsis was formerly used as a putative autonym below F. subg. Meteoriopsis (e.g., Kamimura 1961, Hattori 1975b, Gradstein 1999). However, the establishment of any subdivision of a genus under a subdivision that does not include the type of the genus, does not create an autonym (cf. ICN Art. 22.1). Frullania sect. Vaginatae Verdoorn (1930: 141) was placed into synonymy of F. sect. Meteoriopsis by Kamimura (1961). However, representatives of F. sect. Vaginatae are assignable to the “Diastaloba I”-clade of Hentschel et al. (2009) and therefore excluded from F. sect. Meteoriopsis. The section is circumscribed here to include solely the taxa enumerated by Uribe (2008) and the two characters used to describe the section above exclude taxa of F. sect. Intumescentes and F. sect. Obtusilobae. Frullania (subg. Meteoriopsis) sect. Intumescentes R.M.Schust., Phytologia 57: 370, 1985 (Schuster 1985). Type (ICN Art. 22.6):—Frullania intumescens (Lehmann 1834: 52) Lehm. et Lindenb. in Gottsche et al. (1845: 460). Frullania (subg. Meteoriopsis) sect. Obtusilobae Verd., Ann. Bryol., Suppl. 1: 81, 1930 (Verdoorn 1930). Lectotype (Hattori 1986a: 205):—Frullania meyeniana Lindenb. in Gottsche et al. (1845: 455). Frullania subg. Microfrullania (R.M.Schust.) R.M.Schust., Hepat. Anthocerotae N. Amer. 5: 34, 1992. Basionym:— Neohattoria sect. Microfrullania R.M.Schust., J. Hattori Bot. Lab. 33: 280, 1970 (Schuster 1970). Type (Schuster 1970: 280):—Neohattoria parhamii Schuster (1963: 243) (≡ Frullania parhamii (R.M.Schust.) R.M.Schust. ex von Konrat, L.Söderstr. et A.Hagborg in Söderström et al. (2011: 407)]. Note:—This subgenus has been investigated by von Konrat et al. (e.g., 2006, 2010, 2011a, 2012, 2013). An even more comprehensive understanding might be expected when F. neocaledonica J.J.Engel in Engel & Smith Merrill (1999: 344) or the markedly polymorphous F. junghuhniana Gottsche in Gottsche et al. (1845: 444), and F. rostrata (Hooker & Taylor 1845: 87) Gottsche et al. (1845: 445) also are included. We recognize the three sections below as belonging to this subgenus. Frullania (subg. Microfrullania) sect. Amphijubula (R.M.Schust.) von Konrat, Hentschel, Heinrichs et Braggins, Bryologist 114: 53, 2011 (von Konrat et al. 2011a). Basionym:—Amphijubula R.M.Schust., J. Hattori Bot. Lab. 33: 298 (Schuster 1970). Type (Schuster 1970):—Amphijubula spruceana Schuster (1970: 301) [= Frullania microcaulis Gola (1922 [1923]: 172), cf. von Konrat et al. (2011a)]. Frullania (subg. Microfrullania) sect. Microfrullania (R.M.Schust.) von Konrat et Hentschel, comb. nov. Basionym:—Neohattoria sect. Microfrullania R.M.Schust., J. Hattori Bot. Lab. 33: 288, 1970 (Schuster 1970). Type:—Neohattoria parhamii Schuster (1963: 243) [≡ Frullania parhamii (R.M.Schust. 1963: 243) R.M.Schust. ex von Konrat et al. in Söderström et al. (2011: 407)]. Note:—Schuster (1991) used the name F. sect. Microfrullania as a putative autonym of F. subg. Microfrullania. However, the establishment of any subdivision of a genus under a subdivision that does not include the type of the genus, does not create an autonym (cf. ICN Art. 22.1). Hence, we here validate the section. Besides the type species, this section accommodates F. chevalieri (Schuster 1970: 289) Schuster (1992: 34), F. microscopica Pearson (1922: 33), and F. neocaledonica J.J.Engel in Engel and Smith Merrill (1999: 344). Frullania (subg. Microfrullania) sect. Regulares Verd., Ann. Bryol., Suppl. 1: 133, 1930 (Verdoorn 1930). Lectotype (Hattori 1976):—Frullania junghuhniana Gottsche in Gottsche et al. (1845: 444). Note:—Verdoorn (1930) originally described F. sect. Regulares as a section of F. subg. Diastaloba. Hentschel et al. (2009) showed that the F. rostrata subclade of F. subg. Microfrullania corresponds to F. sect. Regulares. However, the type of F. sect. Regulares has not been included in any molecular analyses. Frullania subg. Saccophora Verd., Ann. Bryol. 2: 121, 1928 [1929] (Verdoorn 1928b). Lectotype (Hattori 1973c):— Frullania sublignosa Stephani (1894: 148). INFRAGENERIC CLASSIFICATION OF FRULLANIA Phytotaxa 220 (2) © 2015 Magnolia Press • 133 Note:—The two subgenera, F. subg. Fusiorielligerae (Verdoorn 1930: 91) Hattori (1972b [1973]: 124) and F. subg. Saccophora (e.g., Hattori 1976, 1982, 1986b) have both been used to refer to a small group of allied taxa. However, Schuster (1992) combined both of these subgenera with reference to several shared morphological features. This treatment is adopted for the forthcoming worldwide checklist. There has been a long-lasting debate about the systematic position of this small and rather anomalous group (e.g., Stotler 1969, Hattori 1973b, 1975a, Schuster 1992, Gradstein & Costa 2003), characterized by rather small lobules, broadly inserted leaf lobes and the initial branch leaf with two saccate lobes inserted below the branch origin (cf. Schuster 1992). Although F. gaudichaudii (Nees & Montagne 1836: 64) Nees et Mont. in Gottsche et al. (1845: 435) was included in the molecular analysis by Hentschel et al. (2009), the study failed to resolve the systematic position of F. subg. Saccophora. Therefore, a more comprehensive study is needed. Frullania subg. Steerea (S.Hatt. et Kamim.) R.M.Schust., Hepat. Anthocerotae N. Amer. 5: 32, 1992 (Schuster 1992). Basionym:—Steerea S.Hatt. et Kamim., J. Hattori Bot. Lab. 34: 429, 1971 (Hattori & Kamimura 1971). Holotype:—Steerea mastigophoroides Hattori & Kamimura (1971: 429) [= Frullania clemensiana Verdoorn (1932: 493), cf. Hattori 1975d]. Note:—With only S. mastigophoroides (= F. clemensiana) included, Steerea has been regarded close to but distinct from Frullania (e.g., Hattori & Kamimura 1971, 1976), but is now considered a synonym of this genus (Schuster 1992). The phylogenetic affiliation of this puzzling species known from the mossy forests of Mt. Kinabalu remains unclear, since it never has been investigated by molecular methods. Frullania subg. Thyopsiella Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 41, 1884 (Spruce 1884). Lectotype (Uribe & Gradstein 2003: 195):—Frullania tamarisci (Linnaeus 1753: 1134) Dumortier (1835: 13). Note:—Uribe and Gradstein (2003) lectotypified this subgenus with F. tamarisci, a taxon erroneously regarded as the generitype by many earlier authors. Thus, the subgenus used to be treated as F. subg. Frullania as discussed above. Whereas the circumscription of this subgenus can be deemed sufficiently clarified (Uribe & Gradstein 2003, Hentschel et al. 2009), the taxonomy of the highly polymorphic F. tamarisci is still not completely understood and currently a matter of intense investigation (e.g., Vilnet et al. 2014). Nomenclatural novelties of miscellaneous Frullania names Below are new synonyms, new names and new combinations for miscellaneous Frullania names at the rank of species and below. Frullania (subg. Meteoriopsis sect. Intumescentes) ambronnii Steph., Biblioth. Bot. 87: 242, 1916 (Stephani 1916). Type:—BOLIVIA, Tablas, Herzog 4536, Herb. F. Stephani no. 13533 (G-00066976, lectotype here designated). Note:—The lectotype was annotated as type by both R. Stotler (August 1968) and Y. Yuzawa (23 November 1987), but they did not annotate the other syntype (Herzog 4569). = Frullania harpantha Herzog, Beih. Bot. Centralbl., Abt. 2. 61: 571, 1942 (Herzog 1942), syn. nov. Type:—COLOMBIA. Cundinamarca: Tequendama, 1929, Troll 2140 (JE!, holotype; JE!, isotype). Note:—Raymond Stotler annotated the type as a synonym of Frullania ambronnii in 1969, but according to our data the synonymization has not been published yet. Frullania (subg. Thyopsiella) acicularis Hentschel et von Konrat, nom. nov. pro Frullania tamarisci var. azorica J.-P.Frahm, Trop. Bryol. 27: 102, 2006 (Frahm 2006). Type:—PORTUGAL. Azores: Terceira, Serra de Santa Barbara, 22 August 2004, Frahm Az316 (BONN, holotype; LIS, isotype). Blocking name:—Frullania azorica Sim-Sim, Sérgio, Mues et Kraut, Cryptog. Bryol. Lichénol. 16: 112, 1995 (Sim-Sim et al. 1995). The epithet refers to the long needlelike leaf apices. Frullania (subg. Frullania sect. Frullania) ericoides var. verrucosa (Kamim.) Hentschel et von Konrat, comb. nov. Basionym:—Frullania squarrosa var. verrucosa Kamim., J. Hattori Bot. Lab. 24: 19, 1961 (Kamimura 1961). 134 • Phytotaxa 220 (2) © 2015 Magnolia Press HENTSCHEL ET AL. Type:—JAPAN. Kochi: Tosa, 20 February 1948, Kamimura 1321 (NICH-46849, isotype). Note:—Kamimura (1961) noted that the most diagnostic character of this variety is the verruculose cuticle of the leaf cells together with the propagules, which sometimes are seen on the ventral surface of the leaves. Frullania ericoides (Nees 1833: 346) Montagne (1839: 51) is a polymorphic species and Hentschel et al. (2009) indicated that at least two independent lineages are present. Further investigation of the complex is required. Frullania (subg. Microfrullania) fertilis De Not., Mem. Reale Accad. Sci. Torino (ser. 2) 16: 235, 1855 (De Notaris 1857). Type:—CHILE. Valparaiso: Puccio (FH). = Frullania magellanica var. diminutiva Herzog, Arch. Esc. Farm. Fac. Sci. Med. Cordoba 7: 29, 1938 (Herzog & Hosseus 1938), syn. nov. Type:—CHILE. Valdivia: Quitaluto, Herzog 679 ex p. (JE!, lectotype here designated); Villarica: Pucón, Herzog 204!; Chiloé: Llanquihué, Petrohué, Herzog 522). Note:—Only two of the three syntypes mentioned by Herzog and Hosseus (1938) are located at JE. One of them, Hosseus 204, was revised by G. G. Hässel de Menéndez in 1981 as belonging to F. fertilis. Due to the fact that Hosseus 679 ex p. is fertile (c.per., c.spor, ♂) the second syntype was chosen as the lectotype. Since F. magellanica frequently has been confused with F. fertilis, Hässel de Menéndez & Rubies (2009) considered several reports of F. magellanica F.Weber et Nees in Gottsche et al. (1845: 446) and its varieties doubtful, including the records of F. magellanica var. diminutiva by Herzog & Hosseus (1938). Engel (1978), who had not seen the type specimen of F. magellanica cited F. fertilis as its synonym. A detailed account of both species is presented by Hässel de Menéndez (1983). To decide on the status of F. magellanica subsp. tristaniana (S.W.Arnell 1958: 9) Váňa et J.J.Engel (2013: 59) further investigations are needed. Frullania (subg. Frullania sect. Australes) fugax (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat. 3: 445, 1845 (Gottsche et al. 1845). Basionym:—Jungermannia fugax Hook.f. et Taylor, London J. Bot. 4: 87, 1845 (Hooker & Taylor 1845). Type:—NEW ZEALAND. On Rimelia (Parmelia) reticulata, 1844, J. D. Hooker (ex herb. Lehm., S!) (lectotype here designated [packet of lower right hand corner on sheet]). = Frullania yorkiana Steph., Sp. Hepat. (Stephani) 4: 558, 1911 (Stephani 1911), syn. nov. Type:—AUSTRALIA. Queensland: Cape York, August 1897, Micholitz, Herbarium F. Stephani no. 18236 (G-00066920, lectotype designated by Hattori 1979: 152). Note:— Hattori (1979) noted the similarity with F. fugax and stated that it possibly is a synonym. Frullania (subg. Diastaloba) gracilis (Reinw., Blume et Nees) Nees, Syn. Hepat. 3: 452, 1845 (Gottsche et al. 1845). Basionym:—Jungermannia gracilis Reinw., Blume et Nees, Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12(1): 221, 1824 [1825] (Reinwardt et al. 1824). Type:—INDONESIA. Java: s. loc. spec. (type not located; missing in STR and other searched herbaria). Note:—Frullania gracilis belongs to a polymorphous and taxonomically difficult species complex (e.g., Verdoorn 1929, Hattori 1974a, 1974b, 1975c) and needs further attention. Particularly, the type of Jungermannia gracilis has never been examined. = Frullania minor var. integribracteola S.Hatt., J. Hattori Bot. Lab. 39: 293, 1975 (Hattori 1975a), syn. nov. Type:—INDONESIA. Sumatra, Mt. Singgalang, July 1955, Meijer 1723 (L, holotype; NICH, isotype). = Frullania minor subsp. recurviloba S.Hatt., J. Hattori Bot. Lab. 39: 293, 1975 (Hattori 1975a), syn. nov. Type:—INDONESIA. Sumatra, Mt. Korinchi, 29 July 1956, Meijer B-9055 (L, holotype; NICH isotype). Frullania (subg. Frullania) jovetiana von Konrat et Hentschel, nom. nov. pro Frullania pseudericoides S.Hatt., Bull. Natl. Sci. Mus. Tokyo, B 12: 132, 1986 (Hattori 1986c), nom. illeg. (ICN Art. 53.1; hom. illeg. [non S.Hatt. 1982]). Type:—INDIA. Kurz 1409, Herbarium F. Stephani no. 15963 (G-00115327, holotype; NICH, isotype). Blocking name:—Frullania pseudericoides S.Hatt., J. Hattori Bot. Lab. 51: 256, 1982 (Hattori 1982). Note:—The type is stored in G under the herbarium name “Frullania jovetiana”. It is possible that the name in Hattori (1986c) is a mistake for what was intended to be this name. Frullania (subg. Meteoriopsis sect. Obtusilobae) kunzei var. maritima R.M.Schust. ex Hentschel et von Konrat, var. nov. Based on:—Frullania kunzei var. maritima R.M.Schust., J. Hattori Bot. Lab. 70: 145, 1991 (Schuster 1991), nom. inval. (ICN Art. 37.7; no herbarium indicated). Type:—USA. Florida: Collier Co., Collier-Seminole State Park, R.M.Schuster 76-390 (F!, holotype). INFRAGENERIC CLASSIFICATION OF FRULLANIA Phytotaxa 220 (2) © 2015 Magnolia Press • 135 Frullania (subg. Frullania) laeviperiantha X.L.Bai et C.Gao ex Hentschel et von Konrat, sp. nov. Based on:— Frullania laeviperiantha X.L.Bai et C.Gao, Nova Hedwigia 70: 135, 2000 (Bai & Gao 2000), nom. inval. (ICN Art. 37.7; no herbarium indicated). Type:—CHINA. Yunnan: Ruili, 8 July 1977, Li Xing-jiang 89 (HKAS, holotype; IFSBH, isotype). Frullania (subg. Meteoriopsis sect. Intumescentes) macrocephala (Lehm.) Lehm. et Lindenb., Syn. Hepat. 3: 460, 1845 (Gottsche et al. 1845). Basionym:—Jungermannia macrocephala Lehm., Nov. Stirp. Pug. 5: 20, 1833 (Lehmann 1833). Type:—PERU. s. loc. Kunze s. n. (W, lectotype designated by Stotler 1969). = Frullania repanda Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 175, 1864 (Gottsche 1864), syn. nov. Type:—COLOMBIA. Ad Tequendamam in montibus Fusagasuganis, Triana 2 (COL, lectotype here designated). Frullania (subg. Frullania sect. Acutilobae) monocera (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat. 3: 418, 1845 (Gottsche et al. 1845). Basionym:—Jungermannia monocera Hook.f. et Taylor, London J. Bot. 4: 89, 1845 (Hooker & Taylor 1845). Type:—AUSTRALIA. Tasmania: Terra Van Diemen, misit Taylor 1847 (herb. Lehmannianum) (S-B143254!, lectotype here designated; FH!, isolectotype). Note:—Frullania monocera belongs to a species complex not yet fully understood (e.g., Hattori 1973a, 1979, 1980a). Frullania monocera var. acutiloba (Mitt.) Hentschel et von Konrat, comb. nov. Basionym:—Frullania acutiloba Mitt., J. Proc. Linn. Soc. Bot. 5: 120, 1861 (Mitten 1861). Type:—INDIA. Nilgiri Mts., Perrottet (NY, lectotype Hattori 1973a [see also Hattori 1975b]). Frullania monocera var. subhampeana (E.A.Hodgs.) Hentschel et von Konrat, comb. nov. Basionym:—Frullania subhampeana E.A.Hodgs., Trans. & Proc. Roy. Soc. New Zealand 77: 370, 1949 (Hodgson 1949). Type:—NEW ZEALAND. Wairoa, Maungapoike Falls, December 1937, E. A. Hodgson 1472 (MPN!, lectotype designated by Hattori 1979: 146). Note:—Frullania subhampeana has a complicated history of synonymy and recognition. Hodgson (1972: 110) regarded the taxon as a synonym of F. novae-zelandiae Colenso (1887: 296), a species Hattori (1979) placed in the synonymy of F. hampeana Nees in Gottsche et al. (1845: 426) which he synonymized with F. monocera at the same time. However, Hattori (1979, 1983) recognized F. subhampeana as separate from F. monocera, but commented that they were closely related. Frullania monocera var. undulata (Kamim.) Hentschel et von Konrat, comb. nov. Basionym:—Frullania undulata Kamimura, J. Hattori Bot. Lab. 24: 50 (Kamimura 1961). Type:—JAPAN. Kôchi: Mt. Kajigamori, July 1950, Hr 1385 (NICH, holotype). Frullania (subg. Frullania) multituberculata Hentschel et von Konrat, nom. nov. pro F. kalimantanensis Piippo et S.Hatt., J. Hattori Bot. Lab. 72: 117, 1992 (Piippo & Tan 1992), nom. illeg. (ICN Art. 53.1; hom. illeg. [non S.Hatt. 1986]). Type:—PHILIPPINES. Luzon: Rizal Prov., Tanay, 1977, Alvarez, Saprid & Guerrero 0-771187 (CAHUP, holotype; FH, H, NICH, isotypes). Blocking name:—Frullania kalimantanensis S.Hatt., Bull. Natl. Sci. Mus., Tokyo, B 12: 127, 1986 (Hattori 1986c). Note:—The epithet refers to the tuberculate perianths. Frullania (subg. Frullania) nepalensis (Spreng.) Lehm., Nov. Stirp. Pug. 4: 19, 1832 (Lehmann 1832). Basionym:— Jungermannia nepalensis Spreng., Syst. Veg. (ed. 16) [Sprengel] 4: 324, 1827 (Sprengel 1827). Type:—NEPAL. s. loc. spec., s. n. & s. d., ex hb. Kunze (S-B28164, lectotype here designated). = Frullania grevilleana var. densa Schiffn. ex Verd., Ned. Kruidk. Arch. (ser. 3) 42: 487, 1932 (Verdoorn 1932), syn. nov. Type:—INDIA. Sikkim: Kurseong, October 1898, Decoly (non vidi, type not located; syn. fide description). = Frullania grevilleana var. gebaueri Verd., Ann. Bryol. 5: 164, 1932 (Dixon et al. 1932), syn. nov. Type:—INDIA. Darjeeling: Kataphar, April 1915, A. K. Gebauer (W, herb. Verdoorn [FH?], isotypes). Frullania (subg. Chonanthelia sect. Cladocarpicae) obscura var. spiniloba (Steph.) Hentschel et von Konrat, comb. nov. Basionym:—Frullania spiniloba Steph., Sp. Hepat. (Stephani) 4: 336, 1910 (Stephani 1910). Type:— ECUADOR. loc. et coll. ign., ex hb. Müller s. n. (G, lectotype designated by Yuzawa 1991) ≡ Frullania arecae var. spiniloba (Steph.) Yuzawa, J. Hattori Bot. Lab. 70: 269, 1991 (Yuzawa 1991). Note:—Gradstein (2012) showed that F. arecae is a synonym of F. obscura, but did not transfer the variety. 136 • Phytotaxa 220 (2) © 2015 Magnolia Press HENTSCHEL ET AL. Frullania (subg. Frullania) obtusangula Hentschel et von Konrat, nom. nov. pro Frullania formosae Steph., Sp. Hepat. (Stephani) 6: 539, 1924 (Stephani 1924), nom. illeg. (ICN Art. 53.1; hom. illeg.). Type:—TAIWAN. s. loc. spec., 1903, Faurie 44 (G, lectotype designated by Hattori & Lin 1985: 134). Blocking name:—Frullania formosa Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 46, 1884 (Spruce 1884). Note:—Kamimura (1961) treated the species as a synonym of F. tamsuina Stephani (1910: 444), but he probably examined the syntype from Hawaii instead of the lectotype (cf. Hattori & Lin 1985). The epithet refers to the bluntly toothed underleaves. Frullania (subg. Meteoriopsis sect. Intumescentes) paradoxa Lehm. et Lindenb., Syn. Hepat. 3: 462, 1845 (Gottsche et al. 1845). Type:—Jubula tamarisci β paradoxa, s. loc. spec. [Tropical America] Hampe s. n. (W, lectotype designated by Stotler 1969: 455). = Frullania flexicaulis Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 53, 1884 (Spruce 1884), syn. nov. Type:—ECUADOR. Mt. Mulmul, Jameson s. n. (MANCH, lectotype designated by Stotler 1969: 464). Frullania (subg. Frullania) plicata Hentschel et von Konrat, nom. nov. pro Frullania acutiloba Gerola, Lav. Bot. Ist. Bot. Univ. Padova 12: 477, 1947 (Gerola 1947). Type:—ETHIOPIA. Socorà, 5 November 1937; Irgalem, 25 November 1937 (syntypes, non vidi). Blocking name:—Frullania acutiloba Mitt., J. Proc. Linn. Soc., Bot. 5: 120, 1861 (Mitten 1861). Frullania (subg. Frullania) setchellii Pearson, Univ. Calif. Publ. Bot. 10: 326, 1923 (Pearson 1923). Type:—NEW ZEALAND. North Island: near Waiotapu, 1904, W. A. Setchell 159 (UC!). = Frullania engelii S.Hatt., J. Hattori Bot. Lab. 54: 143, 1983 (Hattori 1983), syn. nov. Type:—AUSTRALIA. Tasmania: Florentine River, near Gordon River Road, 5 March 1977, J. J. Engel 15080, (NICH!, holotype; F!, isotype). Note:—The type material of F. engelii and specimens from Tasmania appear morphological identical to those of F. setchellii. Frullania (subg. Diastaloba) sinuata Sande Lac., Ned. Kruidk. Arch. 3: 424, 1855 (Sande Lacoste 1854 [1855]). Type (cf. Hattori 1986b):—INDONESIA. Java: s. loc. spec., inter Plagiochilam abietinam, Junghuhn s. n. & s. d., Herb. Lugd. Bat. 910.291-442 (L). = Frullania falsisinuata S.Hatt. et Piippo, Acta Bot. Fenn. 133: 28, 1986 (Hattori & Piippo 1986), syn. nov. Type:—PAPUA NEW GUINEA. Northern Province: Isuarava, 15 January 1936, Carr 15391 (L-HLB 937.162-27, holotype; NICH-431958, isotype). Note:—Frullania falsisinuata was established as a nomen novum for F. sinuata subsp. novoguinensis S.Hatt. (Hattori 1975a: 301). = Frullania falsisinuata var. crispidentata S.Hatt. et Piippo, Acta Bot. Fenn. 133: 29, 1986 (Hattori & Piippo 1986), syn. nov. Type:— PAPUA NEW GUINEA. Morobe: Mt. Sarawaket Southern Range, 1981 Koponen 31586a. (NICH-400031, holotype; LAE, H, isotypes). Jubula hutchinsiae (Hook.) Dumort., Syll. Jungerm. Europ.: 36, 1831 (Dumortier 1831). Basionym:—Jungermannia hutchinsiae Hook., Brit. Jungerm.: tab. 1, 1812 (Hooker 1812). Type:—IRELAND. Bantry, Hutchins (BM, lectotype; H-2313, isolectotype). = Frullania hutchinsiae β compacta Carrington, Trans. Bot. Soc. Edinburgh 7: 456, 1863 (Carrington 1863), syn. nov. Type:—IRELAND. Eagle’s Nest; Cromaglan (MANCH, syntypes). Acknowledgement The Early Land Plants Today project (ELPT) has been generously supported in part by the Global Biological Information Facility (GBIF) Seed Money Award No.2007-41, activities facilitated in part by funding from the Biodiversity Synthesis Center of the Encyclopedia of Life (BioSynC), partial funding from the National Science Foundation (Award No’s 0749762, 0949136, 1145898, and 1115002), the Warwick Foundation, and the Negaunee Foundation. We also acknowledge the support of the Museum Collection Spending Fund, administered by The Field Museum, as well as curatorial support provided by Yarency Rodriguez and Lucia Kawasaki (The Field Museum). 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