Frullania

Hentschel, Jörn, Konrat, Matt Von, Söderström, Lars, Hagborg, Anders, Larraín, Juan, Sukkharak, Phiangphak, Uribe, Jaime & Zhang, Li, 2015, Notes on Early Land Plants Today. 72. Infrageneric classification and new combinations, new names, new synonyms in Frullania (Marchantiophyta), Phytotaxa 220 (2), pp. 127-142 : 131-133

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

( 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).

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.