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Animadversions on the Moss Genus Cratoneuron (Sull.) Spruce

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Animadversions on the Moss Genus Cratoneuron (Sull.) Spruce fourn. Haltori BOI. Lab. No. 6 7: 203- 242 (Dec. 1989) ANIMADVERSIONS ON THE MOSS GENUS CRATONEURON (SULL.) SPRUCE R YSZARD OCHYRA 1 ABSTRACT. A review of taxonomic problems in Craloneuron (Sull. in Gray) Spruce is provided and all taxa that had previously been included in the genus are discussed at length. Cralonellron, since its inception, has been an unnatural and polyphyletic taxon and all the species of the genus, except C. jilicinum (Hedw.) Spruce, are transferred to other genera, leaving Craloneuron monotypic. Hypnumfallax Brid., Craloneuron punae C. M uell. , C.formosanum Broth., C. procerum (Dix .) Broth. and C. subcurvicaule P. Varde are reduced to synonymy with C.jilicinum. One distinct variety, C.jilicinum var. alrovirens (Brid.) Ochyra, is recognized and it includes the plants that have frequently been named C. jilicinum var. fallax (Brid .) G. Roth by most workers, and Hypnum vallis-clausae Brid. and Amb/ys/egium formianum Fior.-Mazz. are identical to this variety. The monotypic family Cratoneuraceae Moenk. is restored to accommodate Cratoneuron and it is considered to be related to the Leskeaceae. The taxonomic status of Hypnum curvicaule Jur., a species that for a long time has been treated as C. Jilicinum var. curvicaule (Jur.) Moenk., is discussed at length. This taxon is retained in the Amblystegiaceae in the new genus Callialaria Ochyra as C. curvicaulis (lur.) Ochyra. Craloneuron commulalum (Hedw.) G . Roth and C. decipiens (De No!.) Loeske are transferred to the newly erected genus Paluslriella Ochyra that is allied to Helodium (Sull.) Warnst. and its relatives as P. commulata (Hedw.) Ochyra and P. decipiens (De Not.) Ochyra. Several varieties of the former C. commulalum and C. decipiens are given formal names in Pa/uslriella, including P. commulala var. diaslrophylla (Brid.) Ochyra, P. c. var.falcala (Brid.) Ochyra, P. c. var.flucluans (B., S. & G .) Ochyra, P. c. var. irrorala (Mikut.) Ochyra, P. c. va r. minor (Lesq.) Ochyra, P. c. va r. pseudodecipiens (Amann) Ochyra, P. c. var. plychodioides (G. Roth) Ochyra, P. c. var. su/cala (Lindb.) Ochyra and P. decipiens var. napaeiformis (Schiffn.) Ochyra, while Cratoneuron commutalovirescens Amann and C. su/caloirrigalllm Meyl. are tentatively reduced to the status of varieties within P. commutata as P. c. var. commulalovirescens (Amann) Ochyra and P. c. var. su/calOirrigata (Meyl.) Ochyra. A brief assessment of the Thuidiaceae is presented and its subfamily Helodioideae Fleisch. is raised to family rank as Helodiaceae (Fleisch.) Ochyra. Apart from He/odium and Paluslriella it includes Bryochenea Gao & Chang and Actinothuidium (Besch.) Broth., while Bryonoguchia Iwats. & Inoue and Hylocomiopsis Card. are excluded from the fam il y. A key for separation of the genera of the Helodiaceae is given. Other heterogeneous elements in Craloneuron are a lso discussed. C. perplicalum (Dusen) Broth. and C. submersum Herz. are identical to Drepanocfadus /ongifolius (Mitt.) Broth. ex Par.; C. sordidum (c. Muell.) Broth. and C. drepanocfadioides Broth. in Dryg. are inseparable from Drepanocfadus sendlneri (Schimp.) Warnst. ; C. mendozense Herz. is placed in synonymy with Drepanocfadus aduncus (Hedw.) Warns!.; and Hypnum filicinum var. minus Wils. & Hook. f., Craloneuron kerguelense (Mitt.) Broth. and C. arclicum Steere are considered to be synonymous with Pseudoleskea chilensis (Lor.) Ochyra. The conspecificity of the latter species has considerably extended the range of P. chilensis that is now established as a bipolar disjunct. BRIEF HISTORICAL A CCOUNT Sullivant (\856) distinguished Cratoneuron as a section of the then all­ encompassmg Hypnum Hedw. to accommodate Hypnum filicinum Hedw. and H. I Laboratory of Bryology, Institute of Botany, Polish Academy of Sciences, Lubicz 46, PL-31-5 12 Krak6w, Poland. 204 Journ. Haltori Bot. Lab. No. 67 1 989 commutalum Hedw., two widely distributed species that are associated with wet or otherwise moist habitats. The section was subsequently elevated to subgeneric rank within Hypnum by Schimper (1860) and Spruce (1867) promoted it to the generic status. There is, admittedly, no description of the genus in Spruce's (1867) publication but Sullivant is cited for the generic name in the genus heading and this is an indirect reference to Sullivant's (1856) work. Since the genus was originally composed of two species, Grout (1931) lectotypified Cratoneuron (Sull. in Gray) Spruce with C.jilicinum (Hedw.) Spruce. The principal criteria for its recognition were the leaves with a strong costa, inflated and pellucid alar cells and the presence of paraphyllia on the surface of the stems and branches. Until the end of the nineteenth century Craloneuron was not accepted as an independent genus. Most bryologists considered it only as a section or subgenus of Hypnum (Schimper 1860, 1876; Hampe 1873; Limpricht 1876; Jaeger & Sauerbeck 1879; Hobkirk 1884; Boulay 1884; Lesquereux & James 1884; Hazslinszky 1885; Klinggraeff 1893) or Amblystegium B. , S. & G. (De Notaris 1867, 1869; Venturi & Bottini 1884). Finally, Roth (1899) resurrected Cratoneuron as a genus of its own and placed it in the newly established family Amblystegiaceae, along with Amblyslegium, Campylium (Sull.) Mitt., Drepanocladus (c. Muell.) G. Roth and Calliergon (Sull.) Kindb. He divided the genus into two sections: (1) sect. Filicina (Brid.) G . Roth (= Cratoneuron) including C.jilicinum, C.formianum (Fior.-Mazz.) G . Roth and C. curvicaule (Jur.) G. Roth and (2) sect. Sulcata G . Roth with C. commutalum (Hedw.) G . Roth, C. falcatum (Brid.) G. Roth, C. sulcatum (Lindb.) G. Roth and C. irrigatum (Zett.) G. Roth. Since then Cratoneuron has gained a wide acceptance of the bryologists who mostly ascribed to G. Roth its status as a genus distinct from all other amblystegiaceous mosses and the name has appeared in almost all floras, manuals, checklists and taxonomic revisions. A few authors have not accepted the placement of Cratoneuron in the Amblystegiaceae and in order to emphasize its distinctiveness they positioned it in the separate family Cratoneuraceae Moenk. (Moenkemeyer 1914, 1927; Podpera 1954; Pilous & Duda 1960; Schlyakov 1961; Abolin 1968; Melnichuk 1970). Since its inception, some 27 species have been validly and legitimately included in Cratoneuron. However, some of these have been reduced to the status of infraspecific taxa and in the result only 18 species still remain in the genus (Wijk et al. 1959, 1969). Closer examination of all species that are currently placed in Cratoneuron reveals that the genus is an increadibly artificial taxon that groups many unrelated species on the basis of superficial similarities of gametophytes, especially falcate leaves, strong costa, inflated and pellucid auric1es and the presence of paraphyllia. Most of these are concentrated around two leading species of the genus, namely C. jilicinum and C. commulalUm. A few extra-Holarctic taxa are poorly known and apparently they have never been examined and tested in order to establish their true relationship. Historically, two lines of interpretation of the interrelationships between C. jilicinum and C. commulalum can be recognized. Most workers followed the concept of Sullivant (1856) and considered these species as closely related taxa within the subdivision Cratoneuron R. OCHYRA: Animadversions on the moss genus Craloneuron 205 in Hypnum. Even prior to recognition of Cratoneuron, both C. filicinum and C. commutatum have often been placed together in various subdivisions of Hypnum which have been given different names, for instance sect. Filicoidea Brid. (Bridel 1801), sect. Filicina (BrideI1812, 1819, 1827; Hlibener 1833; Rabenhorst 1848), sect. Hypnafilicina (Bruch et al. 1854), subsect. Drepanophyllaria C. Muell. (Muller 1848- 1851), sect. Amphimalla Wallr. (Wallroth 1831), sect. Adunca Brid. (De Notaris 1838; Spruce 1849), "Gruppe" Hypna adunca (Girgensohn 1860). Adherents of the second approach considered C. filicinum and C. commutatum to be remotely related taxa and therefore placed them either in two different genera or in various sections within a larger genus. Most often C. filicinum was considered as a species of Amblystegium whereas C. commutatum was retained in the section or subgenus Cratoneuron within Hypnum (Milde 1869; Hartmann 1871; Husnot 1892-1894; Limpricht 1904; Paris 1904, 1905; Herzog 1906; Dixon & Jameson 1924). On the other hand, Lindberg (1879) and Braithwaite (1893- 1905) placed C. filicinum in the subgenus or section Euamblystegium (=Amblystegium) and C. commutatum in the subgenus or section Drepanocladus within the genus Amblystegium, whereas Kindberg (1894, 1896) kept C. filicinum in the section or subgenus Cratoneuron and established the new section and subgenus Alaria for C. commutatum within the genus Hypnum. Finally, some workers included C. filicinum in Hygroamblystegium Loeske that is a segregate of the large genus Amblystegium and left Cratoneuron oligotypic for C. commutatum and the relatives (Loeske 1907a, c; Brotherus 1908). Although the problem of the interrelationships of Cratoneuron species and seemingly related taxa of the Amblystegiaceae and Thuidiaceae was discussed by several authors (Loeske 1907a, c; Moenkemeyer 1911; Dietzow 1914; Fleischer 1923), the final conclusions seem to be unsatisfactory and the genus in the current understanding is very unnatural and polyphyletic. The present account is a new attempt to revise the old question of the span and affinity of Cratoneuron. A NEW GENERIC DEFINITION OF CRATONEURON On the basis of extensive
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