fourn. Haltori BOI. Lab. No. 6 7: 203- 242 (Dec. 1989)

ANIMADVERSIONS ON THE (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 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 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 (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 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, (Sull.) Mitt., (c. Muell.) G. Roth and (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 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 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 observations and detailed comparative study, I define Cratoneuron in much more restricted sense than have previous workers. With this narrower definition the genus may be viewed as circumscribed primarily by gametophytic characters. The extreme similarity of sporophytes, especially peristomes, among hypnobryalean mosses makes them of very limited value, or even useless, in taxonomic and phylogenetic considerations in this group of pleurocarps. Because Cratoneuron is lectotypified with C.filicinum (Grout 1931), the genus must include the lectotype species and the allied taxa. Thus, in the present circumscription it corresponds to some extent to section Filicina of Cratoneuron sensu Roth (1899), the only exception being C. curvicaule which I consider a generically distinct taxon.

Cratoneuron (Sull. in Gray) Spruce Cat. Musc. Amaz. And. 21. 1867. Hypnum Hedw. sect. Craloneuron Sull. in Gray, Man. Bot. No. U. S. ed. 2: 673 . 1856 (X). - Hypnum 206 lourn. Hattori Bot. Lab. No. 67 I 9 8 9

FIG. 1. Cratoneuronfilicinum (Hedw.) Spruce var.filicillum. 1- 5: leaves; 6- 9: outline of various paraphyllia; 10: cellular details of paraphyllium; 11: outline of pseudoparaphyllium; 12: alar cells; 13: mid-leaf cells at margins; 14: mid-leaf cells (all drawn from the lectotype - G). Scale bars: a = lOO Jlm (Figs. 6-14). b = I mm (Figs. 1- 5). subg. Cratoneuroll (Sull. in Gray) Schimp .• Syn. 6 13. 1869. - Amblystegium B., S. & G. sect. Cratoneuron (Sull. in Gray) De Not., Cronac. Briol. Ital. 2: 25. 1867 ["-um"]. Hypnum Hedw. sect. Filicoidea Brid., Musc. Rec. 2(2): 55. 1801 , syn. novo Lectotype species (chosen R. OCHYRA: Animadversions on the moss genus Cratoneuron 207 here): Hypnum jilicinum Hedw. Hypnum Hedw. sect. Filicina Brid., Spec. M usc. 2: 207. 1812 - Hypnum subsect. Filicina (Brid.) Rabenh., Deutsch. Krypt. FI. 2( 3): 277. 1848. - Cratoneuron (Sull. in Gray) Spruce sect. Filicina (Brid.) G . Roth, Hedwigia 38 Beib!. 1: 6. 1899, SYII. novo Lectotype species (selected here): Hypnumjilicinum Hedw. Hypnum Hedw. sect. Faflacia Brid., Spec. Musc. 2: 235. 18 12, 5yn. 1101'. Lectotype species (selected here): Hypnum fallax Brid. Hypnum Hedw. subsect. Drepanophyllaria C. Muell., Syn. 2: 418. 1851 , SYII. novo Lectotype species (selected here): Hypnum jilicillum Hedw. Drepanophyllaria C. Muell., Nuov. Giorn. Bot. Ita!. 3: 114. 1896, lIom. inval. sin. descr. gen.; ibidem 5: 204, 1898, nom. i//eg. incl. gen. prior. [Craloneuron (Sull. in Gray) Spruce)]. Craloneurum 1. Hag., K. Norsk. Vidensk. Selsk. Skr. (Trondheim) 1910(3): 14. 1910, or/hogr. pro Cra/oneuron (Sull. in Gray) Spruce. Plants medium-sized to fairly robust, very variable in habit, in loose to dense, often glossy tufts or mats, bright green, yellowish or brownish, often dark brownish-green with age. Stems wiry in texture, typically erect-ascending, sometimes prostrate or floating, irregularly or more typically pinnately but not complanately branched, rarely unbranched, usually densely tomentose for the greater part of their length, in particular in the prostrate or procumbent forms, with light brown, smooth and branched rhizoids occurring in clusters below the insertion of leaves along stems and branches, in transverse section rounded to elliptical, consisting of 3- 5 layers of small, thick-walled, yellow to brownish, sometimes reddish-brown cortical cells surrounding large, hyaline, thin-walled medullary cells and a distinct, small central strand; stem and branch apices mostly with hooked appearance from falcate leaves, seldom straight; paraphyllia few to many, scattered on the surface of stems and branches, occurring singly or in groups, often attached to the base of the costa, green, unbranched or occasionally bifid, foliose, variously lanceolate, ovate-lanceolate to triangular, plane, non-decurrent, serrulate to dentate, sometimes deeply eroso-dentate at margins; pseudoparaphyllia abundant, broadly ovate to suborbicular, sometimes shallowly bilobed or emarginate, serrulate to crenulate at margins; stem leaves larger and wider than branch leaves, close to distant, erect to spreading, all rigid and not or scarcely altered when dry, cordate-triangular to lanceolate, sometimes secund to fa\cate-secund, gradually or usually abruptly narrowed into a fa irly short, straight or curved acumen, plane or slightly concave, smooth or faintly striate; margins plane or often somewhat recurved at the extreme base, serrulate to serrate nearly all around; costa single, stout or seldom relatively slender, yellow to yellowish-brown, up to one-third the width of the leaf base, gradually tapering upwards, ending below acumen or excurrent as a stout, straight or oblique, smooth, cuspidate point, in transverse section biconvex and composed of uniformly small cells with strongly incrassate walls; angular cells enlarged, hyaline or yellow-brown, thin- or sometimes thick-walled, forming pellucid, inflated and decurrent auricles extending to costa and abruptly differentiated from the adjacent lamina cells; mid-leaf cells smooth or occasionally faintly prorate, firm-walled, shortly oblong to oblong-rhomboidal, mostly 2--4 times as long as wide, but often mixed in central part of leaf with narrower, long rectangular cells up to 6 times as long as wide. Dioicous. Perigonia axillary, bud-like, about I mm long, situated in groups along the stem; perigonial bracts broadly ovate, shortly acuminate, concave, sharply serrulate at margins above; outer bracts with distinct costa and lamina cells resembling those in vegetative leaves; inner perigonial bracts ecostate, with lamina cells short-rectangular and hyaline in the lower part; paraphyses lacking; antheridia short-stal ked, clavate. Perichaetia narrowly cylindric with rhizoids at the base; perichaetial leaves pale or yellowish, erect, sheathing the vaginula, lanceolate to oblong-lanceolate, gradually or suddenly long acuminate, strongly plicate; margins entire below, 208 10urn. J-Iattori Bol. Lab. No. 67 I 9 8 9 sharply serrate in the acumen; costa si ngle, ceasing in the acumen, plane, becoming indistinct or lacking in the innermost perichaetialleaves; lamina cells firm-walled , oblong-hexagonal above, becoming linear-rectangular, thin-walled below; vaginula cylindric, naked or with a few hyaline paraphyses. Setae elongate, 25-40 mm long, smooth, fl exuose, sinistrorse above, dark red to purple below, yellowish- or reddish-brown above; capsules oblong-ellipsoid to cylindrical from a well-developed and disti nct neck, inclined to horizontal, strongly curved, asymmetric and gibbous to almost symmetric, smooth, brown to reddish-brown, strongly arcuate and constricted below the mouth when dry and empty; operculum convex-conic, acute to stoutly rostellate; calyptra cucullate; annulus of 1-2 rows of small cells, separating; exothecial cells firm-walled with distinct collenchymatous thickenings, mostly isodiametric, quadrate to short rectangular, becoming smaller, thin-walled, rounded-hexagonal in 6-8 tiers below the orifice; stomata numerous in the neck, superficial, pale, with long, narrowly elliptic pores; the exostome teeth 16, lanceolate, yellowish brown below to somewhat paler above, on the front surface with a distinct median zig-zag line and dense cross-striolations to above middle, fine ly papillose above and more coarsely papillose at the apex, moderately bordered below and wider bordered above, on the back surface with well-developed, projecting lamellae; the endostome teeth 16, pale yellowish to almost hyaline, with a hi gh, almost smooth basal membrane of about half the length of the teeth giving rise to broad, keeled, sometimes narrowly perforate, papillose segments; cilia 2- 3, well-developed, as long as the segments or shorter, roughly papillose, nodose. Spores spherical, 14-22/lm in diameter, brownish, finely papillose. Type species: Cratoneuron jilicillum (J-I edw.) Spruce.

Cratoneuron filicinum (Hedw.) Spruce var. filicinum Cal. Musc. Amaz. And. 21. 1867 . - Hypnum .filicinum Hedw ., Spec. Muse. 285, 761 5- /0. 1801. - Stereodon jilicillus (J-Iedw.) Mitl., J. Linn. Soc. Bol. 8: 43. 1864. - Amblystegium jilicillum (J-Iedw.) De Nol., Cronac. Bri o!. Ita!. 2: 25. 1867. - Thuidium jilicinum (Hedw.) Kindb., Bih. K. Svensk. Vel. Ak. Hand!. 7(9): 10. 1883. - Hygroamblystegiunl jilicil/um (Hedw.) Loeske, Hedwigia 46: 313. 1907 (V I). Type: In palustribus scaturigines ad Bieniz, simi libusque locis Chemnitzii im Zeisigwalde; in rupibus humectatis Franconiae, Austriae; in addition Hedwig (180 I) provided referenccs to Dillenius (1741), Linne (1753), Weiss (1770), Necker (1771) and Weber (1778) [Lectotype (selected here): .. a me lectum ... Lipsiae ad scatur retro Bienitz lulio" - G-HedwigjSchwaegrichen!; this specimens was illustrated by Hedwig (1801 , pI. 76, f. 5) and some details of this specimen are illustrated on Fig. I in the present paper]. Synonyms: Hypnllm fallax Brid., Muse., Rec. 2(2}: 66, 21 I. 180 1. - Hypnum jilicinum Hedw. var. fallax (Brid.) Brid., Bryol. Univ. 2: 531. 1827. - Amblystegium irriguum (Hook. & Wils.) B., S. & G . var. fallax (Brid.) Schimp., Syn. 594. 1860 - Amblystegium fallax (Brid.) Milde, Bryol. Si1es. 325. 1869. - Amblystegium jilicinllm (Hedw.) De Nol. subsp. fallax (Brid.) Lindb., Musci Scand. 32. 1879. - Thuidium fallax (Brid.) Kindb., Bih. K. Svensk. Vel. Ak. Hand!. 7(9): 10. 1883. - Hypnum vallis-clwlsae Brid. var. fallax (Brid.) Boul., Musci n. France 51. 1884. - H. jilicinum Hedw. subsp. fr"lax (Brid.) Bryhn, K. Norsk. Vid . Selsk. Skrifl. 1892: 220. 1893. - Amblystegium jilicinum (Hedw.) De Nol. var. fallax (Brid.) Husn., Mus. Gall. 362. 1894. - Hygroamhlystegiumfallax (Brid.) Loeske, Moosfl. Harz. 298 . 1903. - Cratoneuron filicinum (Hedw.) Spruce var. fallax (Brid.) G . Roth, Eur. Laubm. 2: 532. 1904. - Hygroamblystegium filicinum (J-Iedw.) Loeske va r. fallax (Brid.) Roell, Jahrb. Ak. Wi ss. Erfurt n. ser. 41 : 149. 1915. - Cratoneuronjilicinum (Hedw.) Spruce subsp. fallax (Brid.) Giac., Alti Isl. Bol. Univ. Lab. Critl. Pavia ser. 5, 4: 259. 1947. Type: In Alpium Sancnsium uliginosis, ubi legi, et in Silesia, unde a revcr. Starkio missum accepi, habitat [Lectotype (selected here): "Hypnum fallax Alpes dc Gessenen 1796" - B- Brid! Syntype: "J-Iypnum fa llax Brid . Stark" - LE ' - the specimen is identical to Hygroam­ blystegium fiuviatile (Hedw.) Loeske!!!], syn. novo Cratoneuron punae C. Mull., Nuov. Giorn. Bol. Ita!. n. ser. 4: 162. 1897. Type: Boli via, Papa Chacra in Puna alia: P. G . Lorentz Ig. 26 Majo 1873 sterile [Lectotype (selected here): " Herb. V. F. Brotherus. R. OCHYRA: Animadversions on the moss genus Cratoneuron 209

Cratoneuron Punae C. M. Argentinia leg. P. G . Lorentz" - H-Broth!], syn. novo Cratoneuron procerum (Dix.) Broth. in Engl. & Prantl, Nat. Pflanzenfam. ed. 2, 11: 334. 1925. - Hygroamblyslegium procerum Dix., Smiths. Misc. Coll. 69(2): 23, 11 . 1918 . Type: vicinity ofThika, British East Africa, alt. about 1.350 meters, Sept. 6 and 7, 1909, No. 1140 .... A few stems were found mixed with No. 1139 (Philonolis speirophylla) [Lectotype (chosen here): No. 1140 - BM-Dix!; isolectotypes: F!, FH!, MO!, NY!, pc! Syntype: No. 1139 - BM-Dix!], syn. novo Craloneuron formosanum Broth., Ann. Bryol. I: 22. 1928. Type: Formosa: Prov. Taityn, Onae, ad rupes (1 . Suzuki). Taparon (E. Matsuda) [Lectotype (selected here): "2894. Cratoneuron formosanum Broth. n. sp. Formosa, prov. Taityn, Onae, on rock, 21 / 11 1926 leg. 1. Suzuki corn. H. Sasaoka" - H-Broth!], syn. novo [Craloneuronformosicum Broth. ex Sak., Bot. Mag. Tokyo 55: 210, 10.1 941 orlhogr.pro C.formosanum]. Craloneuron subcurvicaule P. Varde, Ark. Bot. ser. 2, 3: 185, 29. 1955. Type: [Tanzania, Kilimanjaro Mts] W. slope of Mawenzi, 4700 m, on a moist rock festooned with icicles (l274d) [Holotype: pc-Varde!; isotypes: s!, ups!], syn. novo Through the reduction of the aforementioned species names to the synonymy with C. jilicinum, the genus Cratoneuron is left monotypic. C. jilicinum is very wide­ spread and locally abundant, especially in basic habitats, throughout the Holarctic. In addition, it occurs at some disjunct and geographically isolated stations in the high mountains in East Africa and South America as well as in New Zealand. The species is exceedingly malleable and polymorphic. Consequently, it has been burdened with numerous infraspecific taxa, but they integrade to such an extent that they can be hardly recognized as separate taxa. This polymorphism may be partially genetic, but to a great degree is induced by differences in environment, particularly in water and nutri­ ents supply and illumination. Of the many various infraspecific taxa of C. .filicinum which have been taxonomically segregated at one time or another, the following two are generally recognized as varieties, and sometimes they are also given the separate specific status. One of these, C. jilicinum var. curvicaule (Jur.) Moenk., is here con­ sidered as a distinct species that must be segregated into a genus of its own which is remotely related to Cratoneuron, and both taxa have been associated with one an­ other on the basis of the ostensible similarity of leaves which is doubtless a result of perfect evolutionary convergence. The second morphologically distinct expression of C. jilicinum is generally called C. jilicinum var. fallax (Brid.) G. Roth. The plants referred to this variety are robust and coarse, with stems hardly radiculose and owing to the denuded foliage in older parts appearing hispid and bristly because of persistent costae that remain attached to stems and branches after erosion of leaf laminae and have erect, straight and lanceolate leaves with very strong and thick, 100-200.um wide at base, costae that are long excurrent as stout, cuspidate points. Bridel (1801) described Hypnumfallax from material collected by himself in the Alps and by Starke in Silesia. Examination of both syntypes revealed that the specimen from the Alps is truly Cratoneuron, while the specimen from Silesia is actually Hygroamblystegium fluviatile (Hedw.) Loeske. On the other hand, Bridel's specimen, which is here selected as the lectotype of Hypnum fallax (see above), does not differ materially from other expressions of C. jilicinum that are commonly found in wet or moist habitats in the mountains and it is characterized by percurrent to subpercurrent costa that is moderately, 60-80/lm wide at the base (Fig. 2). As a result I interpret the type collection of H.fallax as representing 210 Journ. Hallori Bot. Lab. No. 67 1 98 9

a typical phase of C. jilicinum. The referred to this variety is sufficiently distinct morphologically and ecologically and does merit recognition as a separate variety of C. jilicinum. This taxon has often been considered as a separate species and for the first time it was described by Bridel (1812) as Hypnum vallis-clausae (Fig. 2). Unfortunately, it has been variously interpreted by subsequent bryologists, but generally it was considered to be identical to Hypnum fallax (Renauld 1885; Loeske 1908) or (Limpricht 1904). Later, Bridel (1827) described additionally H. vallis-clausae var. atrovirens (Fig. 2), a variety that differs from the type variety only in plant colour, a point which is unlikely to give rise to recognition of such modification as a separate taxon. However, the name of this variety is the only available name that can be used for plants that have been commonly referred to as var.fallax by numerous bryologists. Accordingly, the following new combination and synonymy are necessary:

Cratoneuron filicinum (Hedw.) Spruce var. atrovirens (Brid.) Ochyra, comb. novo Basionym: Hypnum vallis-clausae Brid . var. atro-virens Brid., Bryol. Univ. 2: 534. 1827. - H. filicinum subsp. alro-virens (Brid.) Husn., Fl. Mouss. N. Ouest cd. 2: 159. 1882. - Amblyslegillm vallis-clausae (Brid.) Husn. var. alro-virens (Brid.) Corb., Mem. Soc. Sc. Nat. Cherbourg 24: 311 . 1889. Type: In aquis fluentibus prope Falaise Normandiae. D. de Brcbi sson [Holotype: " Hypnum Vallis Clausae f3 atro-virens. In aquis puris Armoricae prope Falaise Berbison [sic!] 1823 L. Pylaie dedit" - B-Brid'). Hypnum "aI/is-clausal' Brid ., Spec. Muse. 2: 238. 1812. - H. clallsaevallis Guer. in Steud., Nomencl. Bot. 2: 209. 1824 nom. nudo in synon. - H.filicinum Hedw. var. vallis-clallsae (Brid.) Hampe, Flora 20: 274. 1837 . - H. filicinum Hedw. subsp. val/is-clausae (Brid.) Boul., Muscin . France 50. 1884. - Amblyslegium filicinum (Hedw.) De Not. var. vallis-clausae (Brid.) Dix ., Stud. Handb. Brit. Moss. 446. 1896. Amhlystegillm val/is-c/ausae (Brid .) Husn., Musc. Gall. 2: 361 , 103. 1893 . Type: In aquis fluentibus Vallis clausae in Gallo-Provincia Guerin legit et communicavit [Holotype: " Hypnum Vallis-Clausae. Hypno Seligeri proximum. Hypnum Selige ri? var. eleganter-pinnati? Vauclusae aut. 1803. e fonte Vauclusae 1803, [illegible), donavit Guerin 1803" - Il-Brid!), syn. novo Amblyslegium formianllm Fior.-Mazz., Alii Ac. Pont. Nuov. Lincci 27: 101 , 3 f 1--4. 1874. - Hypnum formianum (Fior.-Mazz.) Schimp., Syn. ed. 2: 741. 1876. - Cralonellron formianum (Fior.-Mazz.) G . Roth, Hedwigia 38 Beibl. I: 6.1899. - C.filicilllll11 (Hedw.) Spruce var.formial1l1m (Fior.-Mazz.) Latz., Beih. Bot. Centralbl. 48(2): 500. 1931. Type: In Formia, terra di Lavoro nella provincia di NapoIi, nella sorgente passalo pOl11e, onde derivan l'acque i molini [Isotype: KRAM-B (L. Rabenborst, Bryotheca Europaea No. 1293»), syn. novo The species that was originally described by Fiorini-Mazzanti (1874) as Amblystegiumformianum from material collected from Italy also belongs to this variety. Shortly after its description Geheeb (1874) suggested the identity of this moss with Amblystegiumfallax. The type material of A.formianum indeed perfectly matches the type of the present variety of Cratoneuron jilicinum. A more complete discussion of C. jilicinum will be given separately when the full study is finally produced, with special consideration of its increadible variability, a formal list of synonyms, ecology and geographical distribution. The present account aims at discovery of the proper systematic disposition of this bizarre moss genus. As defined here, Cratoneuron is a sharply isolated and rigidly circumscribed genus. With very little experience one can easily recognize it on the following bases: (I) the R. OCHYRA: Animadversions on the moss genus Cratoneuron 211

FIG . 2. Cratoneuronjilicinum (Hedw.) Spruce var.jilicinum (Figs. 1- 3) and var. atrovirens (Brid.) Ochyra (Figs. 4-16). 1- 8: leves; 9- 11 : paraphyllia; 12- 13: alar cells; 14-15: mid-leaf cells; 16: mid-leaf cells at margin (1 - 3 drawn from the lectotype of Hypnumfallax; 4-6, 10-11, 13 , 15- 16 drawn from the holotype of Hypnum vallis-clausae var. atro-virens; 7- 9, 12, 14 drawn from the holotype of Hypnum vallis-clausae - all at B-Bridel). Scale bars: a = 100 Ilm (Figs. 9-16). b = I mm (Figs. 1- 8) . 212 Journ. Hattori Bot. Lab. No. 67 I 989 presence of mostly numerous, foliose, green, sharply serrulate to serrate paraphyllia on the surface of the stems and branches; (2) rigid and wiry texture of the plants; (3) pseudoparaphyllia that are broadly ovate to suborbicular, acuminate to emarginate; (4) stem and branches densely tomentose for the greater part of their length with light brown, smooth rhizoids arising in fascicles below the insertion of leaves; (5) stem and branch apices mostly with hooked appearance from falcate leaves; (6) rhomboidal to oblong-rhomboidal, smooth, firm-walled lamina cells that are mostly 2-4 times as long as wide; (7) enlarged, hyaline angular cells forming inflated decurrent auricles extending to the costa; (8) cordate-triangular to lanceolate leaves, contracted at the base and distinctly serrulate to serrate almost all around; (9) single, stout costa, percurrent to long excurrent as a stout, cuspidate point. The systematic disposition of Cratoneuron is difficult to discern for numerous reasons. The perfectly hypnaceous peristome indicate obviously its close relationship with hypnobryalean mosses, but its further affinity within is rather obscure and unclear. The genus has historically been placed in the then all-encompassing Hypnaceae in the proximity of taxa that Roth (1899) segregated into the Amblystegiaceae. The latter placement has been unanimously accepted by bryologists and Cratoneuron has been firmly established as a member of this "aquatic" moss family that is a rather ecological assemblage of various remotely related taxa (Ochyra 1986). Another view of the relationship of Cratoneuron was presented by Moenkemeyer (1914), who established the monotypic family Cratoneuraceae to accommodate this genus, while all amblystegioid genera he retained in the Hypnaceae. This concept has not been widely accepted and the Cratoneuraceae are recognized only in a few handbooks of European mosses (Moenkemeyer 1927; Podpera 1954; Pilous & Duda 1960; Schlyakov 1961; Bertsch 1966; Abolin 1968; Melnichuk 1970; Petrov 1975; Orban & Vajda 1983). Finally, Kindberg (1883) presented the entirely different point of view regarding the affinity of C. filicinum itself. He transferred this species together with what is here called var. atrovirens to Thuidium and stressed that this realignment seems to be most natural ("Obgleich diese beide Arten [i.e. Thuidium filicinum and Th . fallax] keine Blattpapillen haben, ist doch wohl ihre Stellung in dieser Gattung [i.e. Thuidium] natiirlicher, als in Amblystegium oder Hypnum"). In addition, Thuidium was placed by Kindberg (1883) in the Leskeaceae along with Leskea Hedw., Anomodon Hook. & Tay!., Myurella B., S. & G. and Heterocladium B., S. & G ., genera that are presently mostly segregated into the Thuidiaceae. Although discovery of Cratoneuron's position is encumbered with difficulties that are parallel to those with other genera of hypnobryalean mosses, the relationship of this genus does not seem to be nebulous. In my opinion the neglected concept of Kindberg (1883) from the past century should be seriously taken into account for several reasons. The Thuidiaceae and Leskeaceae are considered to be rather closely related families and Buck and Vitt (1986) placed them in the superfamily Leskeacanae which is recognized by more-or-Iess abundant paraphyllia and predominantly papillose and short lamina cells. Of these, the presence of paraphyllia appears to be crucial for the alliance of Cratoneuron with this group of families, at least with the Thuidiaceae R. OCHYRA: Animadversions on the moss genus Craloneuron 213 and Leskeaceae. As I emphasized elsewhere (Ochyra 1987), paraphyllia are the structures that are generally vital in phylogenetic considerations, albeit their value has been hitherto underestimated. Paraphyllia are known to occur in several families of hypnobryalean mosses including Leskeaceae, Thuidiaceae, Hypnobartlettiaceae and Hylocomiaceae. More­ over, they have been recorded in some genera of the Amblystegiaceae, which, however, proved to be misplaced there and consequently have been transferred to other families of both isobryalean and hypnobryalean mosses (Ochyra 1987, 1988). The removal of the genera with paraphyllia from the Amblystegiaceae left it a more natural entity that, however, still needs a refinement. If the paraphyllia were absent Cratoneuron could be easily wedged in the Amblystegiaceae. It shares many features with some genera of this family, especially with Amblystegium and in particular with its segregate Hygroamblystegium Loeske, including leaf shape, strong and percurrent to long excurrent costa and short-celled leaf areolation. In addition, pellucid and inflated, hyaline angular cells are commonly found in other genera of the Amblystegiaceae, for instance in Calliergon (Sull. in Gray) Kindb. or Drepanocladus (C. Muel!.) G. Roth, and the plants are associated with wet or otherwise moist habitats. These similarities are very suggestive and it is tempting to place Cratoneuron in the Amblystegiaceae as it has been done in the past by most bryologists. However, I interpret this striking and superficial resemblance as the result of convergent evolution of aquatic plants. All aquatic mosses, especially those growing in rheophytic ha bitats, have a strong tendency for stoutness of costae and this is evidently a structural adaptation to an aquatic environment (Vitt & Glime 1984). As well, the pellucid, hyaline angular cells a re known in various groups of aquatics, not necessarily closely related, for instance in Brachythecium rivulare B., S. & G . from the Brachytheciaceae and Fontinalis dalecarlica B., S. & G . of the Fontinalaceae. Hedeniis (1987) has shown the different ontogeny of alar cells in the Calliergon--Drepanocladus complex and Cratoneuron, thus confirming their distant relationship. On the other hand, Cratoneuron is habitually very distinct from the members of the Amblystegiaceae in the rigid and wiry texture of the plants, firm-walled lamina cells and, most of all, in having leafy paraphyllia. In terms of traditional families, Cratoneuron is doubtless nearest to the Thuidiaceae-Leskeaceae complex. It has in common with these families short and relatively firm-walled lamina cells, similar texture of leaves, stems and branches as well as paraphyllia. The latter, however, are unlike those found in both families. In the Thuidiaceae the paraphyllia are short-celled, strongly papillose, filamentous and mostly branched and this structure of the paraphyllia is coupled with the strong papillosity of the lamina cells. Such a combinatiCU1 of characters makes Cratoneuron rather distantly related to this family. The Leskeaceae has been variously defined in the past and the family has been a convenient repository for various genera. Recently Crum (1983) and Buck and Vitt (1986) defined this family in a more restricted sense than has often been accepted. The latter authors included in it only Leskea Hedw. and Pseudoleskea 8., S. & G. and their near relatives that are united by their strong tendency towards cells papillose 214 Journ. Hattori Bot. Lab. No. 67 I 9 89 over the lumina and the presence of paraphyllia. The latter are mostly filiform to narrowly lanceolate and are totally unlike those in Cratoneuron. Buck and Vitt (1986) suggest that a tendency toward the papillosity of leaf cells in this family is secondarily derived as a response to xerophytism. Thus, the characters of Cratoneuron also do not fit into the concept of the Leskeaceae. The characters of Cratoneuron make it a rather bizarre genus. The foliose and unbranched paraphyllia are really a unique feature in the Hypnales. The only genus in this order that possesses similar paraphyllia is Miehea Ochyra, a monotypic genus recently described as endemic to the Himalayan region (Ochyra 1989). rt doubtless belongs to the Hylocomiaceae and possesses several ancestral features, of which the most important are unicostate leaves and leafy paraphyllia. Miehea can be immediately recognized from Cratoneuron by (1) the presence of the longitudinal lamellae on stems and branches; (2) the sympodial branching pattern typical of the Hylocomiaceae; (3) deeply concave and strongly plicate, broadly ovate to almost circular leaves; (4) oblong-linear to Iinear-Aexuose lamina cells that are 7- 10 times as long as wide; and (5) basal cells with strongly incrassate and porose walls and with a characteristic orange to golden-brown colour. These features are entirely lacking in Cratoneuron and this precludes any closer alliance of both genera, despite the striking similarity of the paraphyllia. Inclusion of CralOneuron in any of the traditionally recognized families could only contribute to their heterogeneity. Therefore the restoration of the monotypic family Cratoneuraceae appears justified. Cratoneuraceae Moenk. in Pascher, Siisswass. Fl. Deutsch. Oesterr. Schweiz 14: 116. 1914. Type genus: Cratoneuron (Sull. in Gray) Spruce. Synonym: Amblystegiaceae G . Roth subfam. Cratoneuroideae (Moenk. in Pascher) Ochyra & Szmajda, Fragm. Flor. Geobot. 24: 96. 1978 comb. in val. basioll. non citato Although the Cratoneuraceae occupy the isolated position in the Leskeacanae, it seems to be more related to the Leskeaceae than to any other family of this superfamily. Most probably both families evolved from a common leskeoid stock and the present differences between them are associated with their evolution in various environments. The Leskeaceae are mostly associated with xeric environmental niches and consequently they developed thick-walled and often papillose lamina cells, whereas the Cratoneuraceae evolved in more-or-less hydric environmental niches and in the result they lost the tendency for papillosity of the lamina cells. It is worth noting that Pseudoleskea chilensis (Lor.) Ochyra, a species widely distributed in the cool temperate zone in the Southern Hemisphere and recently discovered in the Arctic in the Northern Hemisphere (see discussion on the identity of Cratoneuron arcticum below), strikingly resembles C. filicinum. Some populations of this moss growing in wet habitats have been very often misnamed as C. filicinum. The only real difference between both taxa is the lack of pellucid angular cells and the filiform paraphyllia in P. chilensis, whereas the texture of the plants, leaf shape and areolation and the costa condition are exactly the same. This similarity may not be the result of convergence, but it can be interpreted as a consequence of their closer relationship. R. OCHYRA: Animadversions on the moss genus Cratoneuron 215

THE TAXONOMIC STATUS OF HYPNUM CURV/CAULE Hypnum curvicaule (Figs. 3 & 4) was described by J uratzka (1864) from material collected at several altimontane sites in the Alps. This author compared it to H . filicinum and found that both species have similar overall appearance and leaf a reolation but the new species is at once recognized by the total lack of the tomentum of rhizoids and paraphyllia on the stems and branches. This comment and the true similarity of gametophytes of both H. curvicaule and H . filicinum have undoubtedly told upon evaluation of this moss by subsequent bryologists. The general consensus of opinion has placed H. curvicaule near H. filicinum, although various authors present very dif­ ferent views regarding its status. In older treatments, mostly until the beginning of the present century, this taxon was very often treated as a separate species and associated with Hypnum (Schimper 1876; Jaeger & Sauerbeck 1879; Breidler 1891 ; Husnot 1892- 1894; Hagen 1904), Amblystegium (Dixon & Jameson 1896, 1924; Braithwaite 1893- 1905; Matouschek 190 I; Paris 1904; Dalla Torre & Sarntheim 1904; Limpricht 1904; Sebille 1905; Moller 1907a, b), Cratoneuron (Roth 1899, 1905; Abramova et al. 1961 ; Kuc 1973), (De Notaris 1867, 1869; Venturi & Bottini 1884) or Hygroamblystegium (Loeske 1907b, c). Other authors, however, considered H. curvicaule only as a variety or subspecies of Hypnum filicinum in all possible nomencIatural combinations of this name as visible in the list of homotypic synonyms below (Molendo 1875; Berggren 1875; Venturi 1881 ; Lindberg 1879; Kindberg 1894, 1896; Warnstorf 1906; Moenkemeyer 1911, 1927; Brotherus 1923; Jensen 1939) and Amann (1919) placed it as a subspecies in Amblyslegium fallax (Brid.) Milde. Nowadays, the varietal status Hypnum curvicaule is accepted practically by all bryologists and the name Cratoneuronfilicinum var. curvicaule (Jur.) Moenk. appears in almost all floras, manuals and checklists of mosses (Podpera 1954; Pavletic 1955, 1968; Lazarenko 1955; Szafran 1961, 1963; Kuc 1963; Nyholm 1965; Papp 1967; Zerov & Partika 1975; Ochyra & Szmajda 1978; Smith 1978; Duell 1985). It should be noted, however, that this name does not appear to be the oldest available name for this moss in such nomencIatural combination. Molendo (1865) described from the Alps in Allgau Hypnum filicinum var. supraalpinum. Later, Molendo (1875) commented on this taxon as follows: " ... meiner var. "supraalpinum" sehr wenig verschieden und steht dem H. filicinum sogar niiher (all Molendo's italics), als Bridels Hypnumfallax". Alas, this name has remained totally forgotten in bryology and Podpera (1954) reduced it to the form status of Cratoneuronfilicinum var. curvicaule, though a new nomencIatural combination is here evidently necessary if this taxon has to be retained at varietal level. The systematic position of H. curvicaule has been very perplexing to systematists, though otherwise the moss may be very readily known. The sporophyte of H. curvicaule is still unknown and consequently the uncertainty as to its correct systematic position has continued. Certainly, without the benefit of sporophyte material opinions on the exact systematic position of any moss are conjectural. Nevertheless, despite the sterile condition of H. curvicaule, its general systematic position (i.e. order) should be obvious 216 Journ. Hattori Bot. Lab. No. 67 t 9 8 9 from gametophyte characters alone. This moss is circumscribed by a relatively strong, single costa, the presence of well-marked alar cells, linear and thin-walled lamina cells and ovate-lanceolate leaves that are coupled with the typically pleurocarpous growth-form of gametophytes. On the basis of these characters it can be easily assigned to the suborder Hypnineae of Hypnales as defined by Buck and Vitt (1986). Indeed Hypnum curvicaule shows striking superficial simi larity to Cratoneuron filicinum, especially in having (I) cordate-triangular leaves; (2) serrulate leaf margins; (3) pellucid and inflated alar cells; (4) costa vanishing below the apex; and (5) dioecious sexual condition. This superficial resemblance I interpret as evolutionary convergence of plants growing in similar, wet or moist, environmental niches. On the other hand, the following set of characters differentiates Hypnum curvicaule from Cratoneuron filicinum: (I) plants soft in texture; (2) stem and branch apices straight and acute, very seldom slightly curved; (3) stems and branches without the tomentum of rhizoids or very occasionally with some scattered rhizoids along stems; (4) paraphyllia lacking; (5) leaves concave, erect and straight, never falcate; (6) lamina cells linear-rhomboidal, 6-10 times as long as wide. This array of characters, in particular the lack of paraphyllia and leaf areolation, exclude any closer relationship of Hypnum curvicaule with Cratoneuron. At the same time, it indicates the close affinity of this moss with the Amblystegiaceae, especially to Drepanocladus and Calliergon. Both genera are characterized by having unicostate leaves and well-developed alar cells, at least in many species. In the traditional comprehension both genera are doubtless very heterogeneous entities and need a reinterpretation. Tuomikoski and Koponen's (1979) attempt of a generic re-arrangement of species of the Calliergon-Drepanocladus complex seems to be unsatisfactory and newly circumscribed genera appear to be heterogeneous as well. Hypnum curvicaule does not fit into the concept of Calliergon in its traditional understanding (Karczmarz 1971) because serrulate leaf margins and long acuminate leaves are unknown in this genus. Species of Drepanocladus are basically characterized by having entire leaf margins, with exception of D. uncinatus (Hedw.) Warnst. and the D. fluitans complex. The former is frequently split in the separate genus Loeske and is easily known by its strongly falcate-secund, gradually long-subulate leaves from a lanceolate base that are often slightly to strongly plicate. The genus occupy a rather isolated position within the Amblystegiaceae and obviously H. curvicaule is very different gametophytically from it. Species of the D.fluitans complex form a quite natural group that is characterized by falcate to falcato-secund, more-or-Iess serrulate leaves with distinct, pellucid alar cells and frequent presence of a group of functionless nematogen cells at the leaf apex. They are sometimes segregated into the genus Warnstorjia Loeske. All species of this complex are strongly variable and polymorphic aquatics. [n their typical appearance they are totally unlike H. curvicaule and are immediately recognized by leaf shape and frequent clusters of rhizoids at the back of the leaf tip arising from nematogen cells. However, some extreme habitat modifications are morphologically very different from typical plants and their interpretation is a source of constant confusion. A classic R. O CHYRA: Animadversions on the moss genus Cratoneuron 217 example is here Hypnum pseudostramineum C. MueH. whose systematic evaluation differs radically in various bryologists. Some authors consider it to be merely a phenotypic phase of D.fluitans (Hedw.) Warnst. (Moenkemeyer 1914, 1927), whereas it is given the separate species status by the others (Nyholm 1965), and additionally some authors segregate it into the separate genus Calliergidium (Ren.) Grout (Crum & Anderson 1981). The moss is recognized by its entire, ovate-lanceolate, not falcate leaves that are acuminate to a narrow, blunt or rounded tip and in this form of leaves it approaches species of Calliergon. However, all evidence seems to indicate that such plants are merely habitat modifications of D. fluitans that are created as a result of flooding. Submerged leaves on the stem unmistakably show D.fluitans, while emergent ones are Calliergon-like. The genus Calliergidium was established under a misapprehension rather than true diagnostic combination of stable characters. Anyway, H. curvicaule is a very different species immediately recognized by the strongly serrulate leaf margins, the lack of nematogen cells and sharply acuminate leaves. Because this enigmatic species cannot be accommodated to any known moss genera, it is deemed necessary to establish a new genus for it. The generic name Callialaria refers to its beautiful, suddenly dilated, hyaline or orange-brown alar cells forming decurrencies.

Callialaria Ochyra, gen. novo Amblystegiacearum Plantae paludosae, mediocres, lutescenti-virides vel fuscescentes, molles, parce subnitentes. Caulis prostratus vel ascendens, irregulariter pinnatim ramificatus vel subsimplex, 5 vel pluria cm longus. nudus vel parce radiculosus, apicem rectus et acutus. nonnumquam leniter incurvus, paraphylliis nullis, pseudoparaphylliis foliosis. lanceolatis. Folia caulinaria et ramealia similia, erecta vel erecto-patentia, laxe imbricata, concava, ovato- vel obcordato-Ianceolata. subito fere in apiculum breviusculum acuminata. haud sulcata, toto margine pIano minute serrulata. Costa solitaria, lutescens, subteres, ante apicem evanida. Cellulae laminae unistratosae. laeves, pellucidae, oblongo-rhomboidales, basi parum laxiores, ad angulos subito valde di latatae, hyalinae vel aurantiae, excavatae et decurrentes. Dioicum. Perigonia et sporophyta ignota. Relatively slender to medium-sized, soft plants. in deep, loose or dense mats, somewhat lustrous, green or, more often, yellow-green to yellowish-golden above, brownish below. Stems 2- S, occasionally up to IS, cm long, creeping or ascending, fiexuose, without tomentum of rhizoids or sometimes with scattered, brownish, smooth radicles, sparcely, irregularly or more-or-less pinnately branched, in cross section elliptic or indistinctly rounded-pentagonal, consisting of 2- 3 rows of small, yellowish-brown or brown on old stems, moderately thick-walled cortical cells surrounding 3--4 rows of large, hyaline, thin-walled medullary cells and small central strand; branches up to I.S cm long, slender, almost julaceous; stem and branch apices obtuse or acute, straight or occasionally slightly curved; paraphyllia lacking; pseudoparaphyllia around branch primordia, foliose, acuminate, sharply denticul ate all around. Stem leaves soft, crowded, loosely imbricate, erect when dry, erecto-patent when moist, straight or sometimes with subsecund apices, decurrent, broadly ovate to ovate-lanceolate from a cordate base, abruptly narrowed in a short to relatively long, slender point, smooth or indistinctly striate, 0.9- 1.3 mm long, 0.4-0.6 mm wide; leaf margins plane, sharply to indistinctly serrulate almost throughout; costa single, yellow, slender, 4S- 60 /lm wide at base, in transverse section biconvex composed of 3--4 layers of almost uniform, larger and moderately thick-walled cells, tapering upwards, vanishing well below or sometimes in the base of filiform acumen; lamina cells unistratose, 218 10urn. Hattori Bot. Lab. No. 67 I 9 8 9

FIG. 3. Callialaria curvicaulis (Jur.) Ochyra. I: habit; 2: portion of branch when moist; 3 ~9: stem leaves; 10~ 13: branch leaves (all frawn from the lectotype of Hypllum curvicaule - w). Scale bars: a = 1 mm (Figs. 3- 13), b = 0.5 cm (Fig. I) & c = 1 mm (Fig. 2). R. OCHYRA : Animadversions on the moss genus Craloneuron 219 pellucid, smooth or slightly prorulose, oblong-rhomboidal to linear-rhomboidal, 5- 7 J.lm wide, 35-50 J.lm long, straight or flexuose, mostly attenuate at ends, 6-10 times as long as wide, thin- to moderately firm-walled; alar cells abruptly differentiated, large, rectangular or oval, hyaline to orange-brown, thin-walled fonning wide, inflated and decurrent auric1es reaching the costa; branch leaves similar in shape to stem leaves but smaller, 0.5-0.8 mm long, 0.2-0.3 mm wide, smooth, straight or somewhat secund, serrulate all around, costa vanishing in mid-leaf, lamina cells oblong to rhomboidal, up to 5 times as long as wide, basal cells larger, yellowish, in 1 or 2 rows across the insertion, alar cells scarcely differentiated. Dioicous. Perigonia unknown. Perichaetia small, axillary along the stem; perichaetial bracts slender, ecostate, sharply serrulate at the apex; outer bracts ovate, bluntly acuminate, inner bracts lanceolate, long acuminate; archegonia few among filiform, hyaline paraphyses. Sporophytes unknown. Type species: Callialaria curvicaulis (lUL) Ochyra, comb. novo Basionym: Hypnum curvicaule JUL , Verh. Bot. Zool. Ges. Wien 14: 103 . 1864. - Limnobium curvicaule (JUL) De Not., Cronae. Briol. Ital. 2: 28. 1867. - Amblyslegium filicinum (Hedw.) De Not. vaL curvicaule (Jur.) Mol., Jahresber. NatUTh . VeL Passau 10: 238. 1875. - HYPllumfilicinum Hcdw. var. curvicaule (JUL) BerggL, K. Svcnsk. Vct. Ak . Handl. 13(7): 86. 1875. - Amblyslegium filicillllm (Hedw.) De Not. subsp. curvicaule (JUL) Lindb., Musei Seand. 32. 1879. - Hypnum /ilicillum Hedw. subsp. curvicaule (Jur.) Kindb., Canad. Rec. Se. 6: 74. 1894. - Amblyslegium curvicaule (JUL) Dix., Stud. Handb. Brit. Moss. 447. 1896. - Craloneuroll curvicaule (Jur.) G. Roth, Hedwigia 38 Beibl. I: 6. 1899. - Hygroamblyslegium curvicaule (JUT.) Loeske, Hedwigia 46: 313. 1907 (VI). CralOneuroll filicillum (Hedw.) Spruce var. curvicaule (Jur.). Moenk, Hedwigia 50: 267. 1911. - AmblY.l'legiumfallax (Brid.) Milde subsp. curvicaule (Jur.) Amann, FI. Mouss. Suisse 2: 337. 1919. - Cralolleuroll filicinum (Hedw.) Spruce subsp. cllrvicallle (lur.) Broth., Laubm. Fennosk. 470. 1923. Type: Auf dem Manhardt in den julischen Alpen (9. August 1841 Sendtncr im Hbr. v. Tommasini); Kalkfelswiinde auf dClll grossen Priel in Ober6sterreieh bei 6800' (8. August 1861 Or. Schicdemayr); auf der Hohe des Unterberges bei Salzburg (14. September 1858 Fr. Bartseh); auf der Heukuppe der Raxalpe in Niederosterreich und a uf der Kuppe des Watzmann in Baicrn (Dr. H. W. Reiehardt). Ich [i.e. J. Juratzka] sammelte es auf der Raxalpc und auf dem Sehnecberg in Nicderosterreich. dann a uf dem Pyrgas in Oberosterreich in Hohen von 5-6000 Fuss [Leetotype (selected here): " Hypnum curviea ule Jur. n. sp. Bayern. Zwisehen Geroll auf der Kuppc des Watzmanns. 8. 1863 Or. H. W. Reiehardt" w!; isolectotype: H-SOL' , w-Juratzka' All syntypes are deposited at Wand have been seen and examined d uring the course of the present study]. Additional selected specimens seen: AUSTRIA. Steiermark: Liegnitzh6he near Schladming, ca 1900m, 30 August 1877, Breidler S. 11 . (w); Eisenhut near Turrach, ca 2300 m, 16 July 1878, Breidler S. 11 . (w); Hochwildstelle near Sehladming, 2470 m, 4 August 1870, Breidler S. n. (w); Tamischbaehturm near Hieftau, 2030 m, 4 August 1895, Baumgarlller S. 11. (G, KRAM, w); Sehiedeck near Sehladming, 2200 m, 13 September 1893 , Baumgarlller S. Il. (w); Kl. Koppenkarstein, 31 August 1897, Ballmgarlner S. n. (w); Koppenkarstein near Sehladming Gletsehers, ca 2700 m, Ballmgarlller S. n. (w); Hoehgolling near Sehladming, 5 September 1895, Ballmgarlner S. n. (w); Vordenberg, 27 July 189 1, Ballmgartner S. Il. (w); Lungauer Kalk near Sehladming, 10 September 1895, Ballmgarlner S. n. (w). Niederosterreieh: Gr. Otscher, 1800 m, 28 July 1896, Ballmgarlller S. n. (w); Ochsenboden at Schneeberg, ca 1800m, 4 August 1918, Ballmgarlller S.II. (w); Hoehkar in Schneegruben, 17 July 1896, Baumgartller S. n. (w); near Damboekhaus, ca 1800 m, Baumgartner S. Il. (w). Ober6sterreieh: Gr. Priel, 1800-1900 m, 10 September 1923, Baumgarlner S. n. (w); Waschenegg near Windisch-Darsten, ca 2100m, 6 September 1923, Baumgarlller S. Il. (w). Kiirnthen: Moll, 1865, Molendo S. Il. (w); Glockner-Gruppe, Leitertal, 2500-2600 m, 26 September 1905, Baumgarlner S. Il. (w); Gopnitzfall near Heiligenb1ut, ca 1400m, 28 September 1905, Baumgartller S. Il. (w). Salzburg: Radstiittcr-Tauern, Zehnerkarspitz, ca 2300 m, 4 Septemeer 1897, Baumgarlner S . 11 . (w); Gassteingipfel, 31 August 1867, Lorelllz S. n. (w). Tirol: Martellthale, ca 2330 m, 8 August 1864, LorenlZ S . n. (w). SWITZERLAND. Retisehen Alpen, Val d'Agnelli, 2700m, 19 July 1868, Pfe/fer S. n. (w); Zerveila an der Curatelsehap in Adula, 16 August 1867 , Pfeffer & Holler S. n. (w); Calanda, ca 2000 m, Pfeifer S. 11 . (w); near Grindenwald, 23 August 1906, Culmalltl S. Il. (w). YUGOSLAVIA. Slovenia: Mangart, ca 2500 m, 10 September 1898, Baumgartller S. n. 220 Journ. Hattori Bot. Lab . No. 67 198 9

cflq~ {}IJ&V~ ~ QU ~ CQO, ~ ~IB- vO~~Q_-~-

1 ~)()N~·.cr' "'i , , f ,"",=; 2;

. curvicaulis (Jur. ) Ochyra. 6' . mid-leafI.' transverse ce II s,. 7' . sectIOn mld-. I.ea 0f cells at mar gin FIG..4 Callia/ana. . 3--4' alar ce,lls' 5: leIaf bar apex 100.u,. or all figures. pseudopa raphylhumthe ,lectotype . _ w) . Sca e ~ (all drawn from m R. OCHYRA : Animadversions on the moss genus Cratoneuron 221

(w); Eishoh1en irn Ternovaner-Walde [ =Trnovska surna], October 1902, Loit/esherger s. n. (w) and 12 February 1903, Baumgartner s. n. (w). WEST G ERMANY. Bayern: Frauenalpen near Parten-Kirchen, August 1874 m Amo/d s. n. (w). CZECHOSLOVAKIA. Belanskc Tatry: Zdiarski Vidla, ca 1880 m, August 1947, Pi/oils 303 (BRNM, OP, PRC, w); Grota Alabastrowa, Zmuda 1325 (KRAM). POLA D. Tatry Wysokie: near Siklawica, 13 October 1910, Racihorski s. n. (KRAM); Szpiglasowa Przelecz in Dolina Pieciu Stawow Polskich, Lisowski 64626 (KRAM); Tatry Zachodnie: Kopa Magory, 1600 m, Li~'o\Vski 238 (KR~M); Swistowka, 1867, Rehmann .1'.11. (w). SOVIET UNION. Kazakhstan: Kungej-Alatau Mts, Kurrnitinka, 3300 m, 18 August 1958, Lisowski s. n. (KRAM) . MONGOLIA . Mongolian Altai Mts., Turgen somon, Mt. Turgen-Ula, 6 July 1974, Tsegmed s. 11. (KRAM). CHINA. Yunnan: " inter fluvios Lu-Djiang (Salween) et Djiou-Djiang (lrawadi or. sup.), in regione alpina retro montem Gomba-Ia supra Tschamutong, in fontibus ad lacum Tsukue, 3825 rn, Hallde/­ Mazzelli 9540 (G, H-Broth, w). INDIA . Himalayas, near Manh Pass, Spiti, 4830 m, Badhwar 824 (BM-Dixon). Callia/aria curvicaulis is a very distinct taxon of the Amblystegiaceae that is unlikely to be mistaken for any other moss species and genus. It is an altimontane moss occurring at an elevation of abqve 1800 m and only occasionally it was found at lower altitudes of 1600 m in the Tatra Mountains of Poland and 1400 m in Salzburg of Austria. The species grows in wet to moist habitats, mostly on calcareous ground, seldom on gneiss and schist rocks. C. curvicaulis seems to be widespread and abundant in appropriate habitats throughout the Alps, especially in Austria, but I have also seen collections from Slovenia of Yugoslavia,. Switzerland and West Germany. There are also numerous records of the moss from Italy (Venturi 1881; Dalla Torre & Sarntheim 1904) and France (Sebille 1905). Furthermore, C. curvicaulis occurs in the Tatra Mountains of Poland and Czechoslovakia and from outside Europe I discovered collections from Kazakhstan in the Soviet Union, Mongolia, China and the Himalayas of India where it was collected at an elevation of 4830 m and this is the highest elevation for it. The species has often been recorded in bryological literature, among others from Scandinavia (Nyholm 1965), the British Isles (Smith 1978), the Pyrenees (Casas Sicart 1986), the East Carpathians (Zerov & Partika 1975) and the South Carpathians (Papp 1967) as well as from Svalbard (Kuc 1963), Novaya Zemlya (Savicz 1936) and Axel Heiberg Island in the Canadian Arctic Archipelago (K uc 1973). Herzog (1961a) reported it from Bolivia. All these records are badly in need of verification and only verified material can be used for phytogeographic consideration. It proved that some collections of this species were simply misnamed and actually belong to other taxa, for instance specimens from Svalbard reported by Kuc (1963) and from Novaya Zemlya cited by Savicz (l936) are Drepanocladus aduncus, while Herzog's (1916a) specimens from Bolivia are Cratoneuron jilicinum. At present C. curvicaulis must be considered as a Eurasiatic oreophyte occurring at high elevations in the mountains in the southern part of the Holarctic.

TAXONOMIC STATUS OF CRATONEURON SECT. SULCATA Roth (1899) established section Sulcata of Cratoneuron to accommodate four species, namely C. commutatum (Hedw.) G . Roth. (Fig. 5), C. Jalcatum (Brid.) G. Roth, C. sulcatum (Lindb.) G. Roth and C. irrigatum (Zett.) G . Roth. These species are very closely related and in modern are considered as habitat expressions 222 Journ. Hattori Bot. Lab. No. 67 I 989 of the exceedingly polymorphous and protean C. commutatum. Usually they are not recognized at any taxonomic rank (Crum & Anderson 1981) or are given the status of varieties or forms within C. commutatum (Podpera 1954; Szafran 1961, 1963; Nyholm 1965; Smith 1978). Like with other aquatic mosses, C. commutatum exhibits the polimorphism that is doubtless conditioned by varying degrees of submersion (D avy de Virville 1921 ) and consequently most, if not all, of its habitat expressions are worthless taxonomically. The second distinct species of section Sulcata is C. decipiens (De Not.) Loeske (Fig. 6) . It was originally described by De Notaris (1869) as Thuidium decipiens and subsequently transferred to Cratoneuron by Loeske (1903). The species is readily recognized from C. commutatum by its distinctly papillose lamina cells, although some workers reported some intermediate forms between both species (Blumrich 1917). In addition, the following species fit well into the concept of section Sulcata of Craloneuron. Grout (1931) described from Montana in the United States C. papil/osum [Type: Columbia Falls, Montana, Oct. 10, 1895, R. S. Williams no. 441. Holotype: NY!; isotype: DUKE-Grout!]. Later Grout (1934) proposed the substitute name C. williamsii for this species to replace the nomenclaturall y illegitimate former name. Although Wijk et al. (1959) maintained it as the distinct species, Podpera ( 1954) considered the North American species to be identical to C. commulalum fo . janzenii (Loeske) Podp. Having examined the type material of this species [ agree with Podpera and can confirm that this species does not merit recognition at the species level. The same refers also to two species described from Switzerland by Amann (1919) and Meylan (1930) as C. commulatovirescens and C. sulcaloirrigatum, respectively. Judging from their descriptions alone, they appear to be merely phenotypic expressions of C. commutatum and therefore I tentatively reduce them herein to the status of varieties within this species. Species of section Sulcata are usually distinguishable as such at a glance by the following: (I) stem leaves cordate-triangular, sometimes very broadly so, much narrowed at the insertion, gradually or rapidly tapering into a long and usually curved acumen and deeply plicate; (2) branch leaves ovate-lanceolate, strongly falcate; (3) angular cells enlarged, hyaline to brownish, forming usually decurrencies extending to the costa; (4) leaf margins usually sharply serrulate to serrate; (5) stem and branches mostly densely radiculose; (6) lamina cells, at least in the upper part, narrow, prosenchymatous, oblong-rhomoboidal to linear-f1exuose, distinctly prorate or with tall conical papillae at distal ends; (7) paraphyllia filamentous to narrowly lanceolate, uniseriate or consisting of one row of elongate cells at the distal end and usually several rows of cells in the basal portion; (8) peristome perfectly hypnaceous. Some of the above mentioned features, including the falcate, cordate-triangular shape of the leaves, well-marked alar cells forming decurrencies, the presence of the tomentum of rhizoids on stems and branches and the association of plants with hydric habitats are really reminiscent of those found in Cratoneuron filicinum and seemingly confirm the pertinence of recognition Cratoneuron in its present comprehension and span. Nevertheless the basic dissimilarities between C. .filicinum and species of section R. OCHYRA: Animadversions on the moss genus Cratoneuron 223

Sulcata refer to the essential structural characters that are of special importance in phylogenetic consideration. Of these the most important are paraphyllia and leaf areolation that preclude any closer relationship of both taxa. As a result species of section Sulcata must be excluded from Cratoneuron and the Cratoneuraceae that are characterized by having foliose paraphyllia and short and smooth lamina cells. Because there is no reason for maintaining section Sulcata in Cratoneuron and no other genus seems suitable for the placement of it, [ prefer to describe a new genus to accommodate Hypnum commutatum and its relatives. The generic name is a diminutive derived from the Latin word meaning swamp to emphasize the close association of its species with wet and swampy habitats. For the reasons discussed below Palustriella is placed in the newly erected family Helodiaceae.

Palustriella Ochyra, gen. novo Helodiacearum Hypnum Hedw. sect. CommUlala Debat, Bull. Trim. Soc. Bot. Lyon 3: 55. 1885 nom. nud., syn. novo - Hypnum Hedw. sect. A/aria Kindb., Canad. Rec. Sc. 6: 74. 1894 nom. nud., syn. novo - Hypnum Hedw. subg. A/aria C. Muel!. in Kindb., Spec. Eur. N. Am. Bryin. 1: 126. 1897. Lectotype species (selected here): Hypnum commulatum Hedw. [= Palustriella commulata (Hedw.) Ochyra], syn. novo- Craloneuron (Sull. in Gray) Spruce sect. Sulcala G. Roth, Hedwigia 38 Beibl. I: 6.1899. Lectotype species (selected here): Craloneuron commulalum (Hedw.) G. Roth [= Palustriella commulala (Hedw.) OchyraJ, syn. novo Caulis dichotome divisus, cristato-pinnatus, erect us vel prostratus, frequenter dense tomentosus, paraphylliis multis, filiformibus vel anguste lanceolatis. Folia caulinaria triangulari-cordata vel cordato-deltoidea, remotiuscula, auriculata, decurrentia, pluries sulcata, anguste falciformia, tenui­ acuminata, marginibus serratis, planis; folia ramealia angustiora, ovato-lanceolata, dense conferta, falcato-secunda. Cellulae laminae oblongo-rhomboidales vel ftexuoso-lineares, papillosae; costa solitaria, crassa sub apicem evanida. Capsulae alte pedicellatae, horizontales vel cernuae, arcuatae, operculis convexo-acutis. Peristomium perfecte hypnaceum. Plants slender to robust, in loose or dense cushions or mats, not glossy, soft, bright to dark green, yellowish or brownish above, brown and often encrusted with lime below. Stems strong and stiff, erect or ascending, sometimes procumbent or floating, frequently complanately plumose and pinnate, sometimes irregularly branched, usually densely tomentose with light brown, slightly papillose and strongly branched rhizoids occurring along stems and branches, in transverse section elliptical, consisting of 3-5 layers of small, thick-walled, yellowish-brown to dark brown cortical cells surrounding larger, hyaline, thin-walled medullary cells, central strand lacking or very indistinct, composed of a few small cells; stem and branch tips with hooked appearance from falcate leaves; paraphyllia few to very many on the surface of the stems and branches, often attached to the base of the costa, unbranched, green, filiform and uniseriate or filamentous to narrowly lanceolate consisting of several rows of cells in the basal portion and uniseriate distally, serrate or dentate at margins; pseudoparaphyllia infrequent, lanceolate, long acuminate, serrate at margins; stem leaves crowded to relatively distant, erect-spreading to spreading, strongly falcato-secund, decurrent, cordate-triangular to widely deltoid or ovate-lanceolate, often strongly contracted at the insertion, gradually or abruptly narrowed to a long, slender, twisted or flexuose, circinate to almost straight, usually channelled acumen, strongly plicate; branch leaves shorter and narrower than the stem leaves, often homomallous, ovate-lanceolate, mostly long acuminate; margins plane or slightly recurved in the base, sharply serrate to denticulate below at the widest part of the base, serrulate to almost entire in the upper part of the leaf; costa si ngle, green, yellowish or yellow-brown, ending in 224 Journ. Hatlori Bot. Lab. No. 67 I 989 the acumen and usually extending 2/3 to 3/4 way up the leaf, never excurrent, in transverse section biconvex, consisting of several layers of almost uniform, relatively thick-walled cells; lamina cells with thin to incrassate walls, linear-flexuose to oblong rhomboidal above, smooth or prorate to distinctly papillose at the distal end, 6-15 times as long as wide, becoming shorter and wider, oblong-rhomboidal to rhomboidal, 3- 6 times as long as wide below; cells at the basal angles enlarged, lax, suddenly dilated, hyaline, yellowish or brownish forming decurrent, inflated auricles extending to costa. Dioicous. Perigonia axillary, but-like, less than I mm long, in groups along the stem; perigonial bracts ovate-lanceolate to broadly ovate, abruptly narrowed to an acumen, concave, indistinctly serrulate at margins above, ecostate, with oblong-rhomboidal to oblong-linear, smooth, thin- to firm-walled cells; antheridia few, clavate, short-stalked; paraphyses scarce, filiform , hyaline, usually exceeding the antheridia. Perichaetia narrowly cylindric, not or weakly tomentose at the base; outer perichaetial leaves ovate, broadly apiculate, sometimes ecostate, mostly with strong costa reaching the acumen; inner perichaetialleaves ovate-lanceolate, tapering to a long, often squarrose acumen, with a single costa ceasing in the acumen, distinctly ciliate at the base of the acumen at margins with fragile, uni seriate, short cilia when young, becoming narrowly lanceolate, pale, strongly plicate when mature, having strong, yellow costa, mostly entire margins and linear lamina cells becoming oblong-rhomboidal below and widened and brownish at the base; vaginula cylindric, dark brown, naked or with a few hyaline paraphyses. Setae elongate, 25-40 mm long, smooth, flexuose, si nistrorse, purple to dark brown below, reddish- or yellowish-brown above; capsules large, ± horizontal, mostly elliptic to subeylindric, symmetric or slightly asymmetric and gibbous, strongly curved and contracted below the mouth when dry, smooth, brown; operculum convex-conic with straight or slightly slanted, stout, acute point; ealyptra naked, eueullate; annulus separating of 1- 2 rows of small, thick-walled cells; exothecial cells thick-walled, distinctly collenchymatous, short rectangular to subrounded­ quadrate, beoming isodiametric, hexagonal in 6-8 rows below the mouth and oblate in 2-3 rows at the mouth; stomata numerous in the basal portion of the capsule, pale, superficial, bicellular, with short, round or elliptic pores; exostome teeth 16, lanceolate, brownish at the base, becoming paler, yellow to brownish-yellow above, on the outer surface with a distinct median zig-zag line, densely cross-striolate to above middle, finely papillose above and with scattered, coarse papillae at the apex, moderately bordered below and with a wider border in the upper half, on the inner surface with well-developed trabeculae; endostome teeth 16, with a yellow, smooth basal membrane of medium height; segments broad, keeled, narrowly perforate, slightly papillose, as long as or slightly shorter than the exostome teeth, cilia 2- 3, well-developed, as long as segments or shorter, strongly papillose, nodose. Spores spherical. brownish, 12- 25 ~m in diameter. slightly roughened. Type species: Po/uslriel/a cOl1ll1lulala (Hedw.) Ochyra, comh. no vo Basionym: HYPIIW1I ('ol1lllll/talUI1I Hedw., Spec. Musc. 284. 1801. - Slereodoll COI11I11Uta/u.l' (Hedw.) Milt.. J. Linn. Soc. Bot. 8: 43 . 1864. - AmhiY.I'lef(iulIl commutalul11 (Hedw.) DeNot.. Cronac. Bri ol. Ital. 2: 25. 1867. - Craloneuron commulalum (Hedw.) G. Roth. Hedwigia 38: Beibl. 1: 6. 1988. Type: In palustribus et ad rivulos. maxime regionum calcarearum montosarum Austriae, Franconiae. Hercyniae, Galliae. In addition, Hedwig (180 1) provided references to Hedwig (1797) and Vaillant (1723) [Lectotype (selected here): "Herr Frolich. von dcssen Giitigkeil ich diese Art erhiell, traf sie hiiufig in der algauer Siimpfen an"; this specimen was illustrated by Hedwig (1797) on plate 26 and the above locality data is quoted in this work; the left upper specimen on the sheet in Hedwig's personal herbarium perfectly corresponds to this illustration and is here selected as the lectotype - G-Hedwig/Schwaegrichen!; syntype: "Petersdf pr Vindob d. Bemaetz" - G-Hedwig/Schwaegrichen']. R. OCHYRA : Animadversions on the moss genus Cratoneuron 225

FIG. 5. Pa/us/riel/a commu/a/a (Hedw.) Ochyra. 1- 2: stem leaves; 3- 5: paraphyllia; 6: mid-leaf cells; 7: ciliate ma rgin of perichaetial leaf (all drawn from Ochyra 554/82 - KRAM) . Scale bars: a = 100j1m (Figs. 3- 5) and 50j1m (Figs. 6-7). b = I mm (Figs. 1- 2).

The following is a list of all currently accepted names in the genus Palustriella (Wijk et al. 1959, 1964, 1969) and the appropriate transfers and basionyms. r decided to include herein also all infraspecific taxa of P. commutata and P. decipiens at the varietal level in order to facilitate future study of variability of these protean mosses. There are also numerous names for forms, mostly used by European bryologists (Podpera 1954). These, however, are not given formal names in Palustriella because they are doubtless merely habitat expressions and certainly will be reduced to synonymy in the result of careful taxonomic investigation. 226 Journ. Hattori Bot. Lab. No. 67 I 989

Palustriella commutata (Hcdw.) Ochyra var. commutatovirescens (Amann) Ochyra, Slat. et comb. novo Basionym: Cralolleuroll cOmmulalo- l' ireSCI!IIS Amann , FI. Mouss. Suissc 2: 339. 1919. Palustriella commlltata var. diaJtrophylla (Brid.) Ochyra, comb. novo Basionym: HYPllum COnlmUlalum Hedw. var. diaslrophyllunI Brid., Bryo l. Univ. 2: 525. 1827. PaLlIstriella commlltata var. Jalcata (Brid.) Ochyra, comb. novo Basionym: Hypllum f a/calwn Brid., Mus. Rec. 2(3): 63, / I 6. 1801.

Palustriella commlltata var. jiuctuans (B ., S. & G ,) Ochyra, comb. fl OV. Basionym: H ypllum commUlalum Hedw. var.jiuc/ualls B., S. & G ., Bryol. Eur. 6: 115 608"1. 1854. var. irrorata (Mikut.) Ochyra, comb. novo Basionym: CralOlleurOIl COmmUlalum (Hedw.) G. Roth var. irroratum Mikut.. Bryoth. Bait. 187. 19 13. Palustriella commutata var. minor (Lcsq.) Ochyra, comb. novo Basionym: H yplIllm comnlulalUm Hedw . va r. millus Lesq., Mem. Soc. Sc. Nat. euchatel3(3): 49. 1846. Palllstriella commutata va r. pselldodecipiens (Amann) Ochyra, comb. novo Basionym : CralOlll!urOJ/ commulalum va r. pSl!udodecipiells Amann, Fl. Mouss. Suisse 2: 340. 1919. Palustriella commutata var. ptychodioides (G. Roth) Ochyra, comb. novo Basionym: Craloneuroll plychodioide.l' G . Roth , Hedwigia 49: 226, 8I 12. 19 10.

Palustriella commutata var. sulcata (Lindb.) Ochyra. comb. flOV. Basionym : H ypnum CO/lunufalum Hedw. va r. su/calum Lindb .. Oefv. K . Vet. Ak. Foerh. 23: 540. 1867. Palustriella commutata var. sulcatoirrigata (Mcyl.) Ochyra, stat. et comb. no vo Basionym: Crafolleurol1 su/cafo·irrigalul1l Mcyl. , Rev. Bryol. 3: 186. 193 1.

Palustriella commutata var. virescens (Schimp.) Ochyra, comb. fl OV. Basionym: H ypllum fa/calum Brid. va r. virescl!lls Schi mp .. Syn. cd . 2: 743. 1876. Palllstriella decipiens (Dc NOL) Ochyra, comb. novo Basionym: Thuidium decipiells De Not., Atti Univ. Genova I: 233. 1869.

Palllstriella decipiens var. napaeijormis (Schitfn.) Ochyra, comb. flOV. Basionym: H ypnum decipiens (De Not.) Limpr. in Cohn var. Ilapaeiforme SchifTn., Oesterr. Bot. Zeitschr. 58: 348. 1908 . Palustriella is a small and natural taxon, including only two species, which are widespread throughout the Holarctic in appropriate habitats. At least P. commutata seems to be circumpolar occurring in temperate and boreal zones in the Northern Hemisphere from Europe to Siberia and Japan and bicentrically in North America. The perfectly hypnaceous peristome obviously indicates that the genus is a hypnobryalean moss. Palustriella, as circumscribed here, bears doubtless great similarities to thuidioid mosses. The similarity of sterile plants of P. decipiens to Thuidium s. lato is such that the species has been originally described as a member of this genus. This salient external similarity is followed by excellent correspondence in essential structural features. It appears that Palustriella shares many character states with He/odium (Sull. in Gray) Warnst. and its allies. The presence in this genus of filamentous to filiform paraphyllia, striking leaf dimorphism, leaf plication and areolation and ciliate perichaetial leaves all support a placement of this genus in the R. OCHYRA : Animadversions on the moss genus Cralonellron 227

ca

FIG. 6. Palllstriella decipiens (De Not.) Ochyra. 1- 2: stem leaves; 3: paraphyllia; 4: upper lamina cells; 5: lower lamina cells (all drawn from Ochyra 1616/82). Scale bars: a = 100 Ilm (Fig. 3) and 50 Ilm (Figs. 4-5). b = I mm (Figs. 1- 2).

Thuidiaceae. The relationship of Cratoneuron to this family was suggested a long time ago by Loeske (1907a, 1910), and Fleischer (1923) simply stated that "Die Gattung He/odium is die phyletische Brucke zur Gattung Cratoneuron, inbesondere zu C. decipiens und mithin zu den Amblystegiaceen". The only other family of the Leskeacanae that is diagnosed by the presence of paraphyllia is the Hypnobartlettiaceae (Ochyra 1985, 1987). It includes four salient and sharply isolated genera with very restricted distribution that are all aquatics. The 228 Journ. Hattori Bot. Lab. No. 67 1 989 family is at once distinct from the Thuidiaceae by the lack of leaf dimorphism and predominantly variously bistratose lamina cells. On the other hand, the Thuidiaceae is a very large moss family that in the commonly accepted circumscription comprises some 17 genera divided into four subfamilies (Brotherus 1925) that are characterized by papillose leaf cells and abundant paraphyllia of various types. The family is doubtless unnatural and polyphyletic and so needs a refinement. Although Buck and Vitt (1986) excluded from it Anomodon Hook. & Tay\., Haplohymenium Doz. & Molk. and Herpetineuron (c. Muell.) Card. and placed them in the Anomodontaceae in the Leucodontales, it still cannot be precisely defined. Crum's (1983) proposal as to the generic composition of the Thuidiaceae also does not appear to be convincing or statisfactory. In my opinion the family Thuidiaceae s. sIr. includes primarily genera corresponding to the subfamily Euthuidioideae (= Thuidioideae) in Brotherus' (1925) circumscription and centered around Thuidium B., S. & G., although other genera of the former subfamily Anomodontoideae such as Haploc/adium C. Muell., Miyabaea Broth. and Claopodium (Lesq. & Jam.) Ren. & Card. apparently fit into the concept of the family. In such definition, the Thuidiaceae includes taxa having short, isodiametric, strongly papillose lamina cells and paraphyllia that are branched, filamentous, short-celled and strongly papillose. When considered as a whole, the constelation of character states present in the Thuidioideae is such that it cannot accommodate Helodium and its relatives (and inferentially Palustriella which is their closest kindred) that constitute the subfamily Helodioideae within the Thuidiaceae in their present circumscription (Fleischer 1923; Brotherus 1925). The comparison of diagnostic characters of both Thuidioideae and Helodioideae comfirms the distinctiveness of the latter as a very unique entity that is not especially closely related to the former. Therefore J conclude that the Helodioideae should be raised to family rank. Helodiaceae (Fleisch.) Ochyra, stat. et comb. novo Basionym: Thuidiaccac Schimp. subfam. Helodioideae Fleisch" Musci Ft. Buitcnzorg 4: 1499 . 1923. Type genus: He/odium (Sull. in Gray) Warnst. The Helodiaceae is a very distinct family that can be immediately distinguished from the Thuidiaceae by different shape ofparaphyllia. In both families these structures are very polymorphous, but those in the Thuidiaceae are always short-celled, strongly papillose and ending in a truncate cell with two papillae. The paraphyllia in the Helodiaceae are smooth and never papillose, acute and composed of elongate cells mostly with oblique walls. There are three different types of the paraphyllia in this family: (I) filiform-branched type found in He/odium (Fig. 7: 7) including paraphyllia that are composed of a single row of elongate cells; (2) lanceolate-ciliate type found in Actinothuidium (Besch.) Broth. (Fig. 7: 3) and Bryochenea Gao & Chang (Fig. 7: 5) including paraphyllia that are basically narrowly lanceolate with numerous acute, uniseriate cilia of various length at margins; (3) lanceolate type found in Palustriella (Figs. 5: 3 & 6: 3- 5) including unbranched structures composed of 2 to many rows of cells in the basal portion and uniseriate in the apex. R. OCHYRA: Animadversions on the moss genus CralOneuron 229

FIG. 7. Paraphyllia and lamina cells of BryonoKuchia mofkenhoeri (Sande Lac.) Iwats. & [n oue (Figs. I 2), Actil/otizuidium hookeri (MitL) Broth. (Figs. 3 4), Bryocizellea sacizalillensis (Lindb.) Gao & Chang (Figs. 5~ 6), Hefodium pafut/oslIm (Aus!.) Broth. (Figs. 7 8) and Hyfocomillm spfendens (Hedw.) B., S. & G. (Figs. 9~ 10). (l ~ 2 drawn from /nolle 281; 3 4 from Kanai 476; 5~6 from Watanabe 1225; 7 ~ 8 from Flowers s. 1/ . , 18.6. 193 1; 9~ 10 , from Ochyra 2797/ 79 ~ all in KRAM). Scale bar: 100 I'm (Figs. I, 3, 5, 7 & 9) and 50l/m (Figs. 2, 4, 8 & 10). 230 Journ. Hattori Bot. Lab. No. 67 I 9 8 9

The shape of the paraphyllia is coupled with characteristic leaf areolation of mostly prosenchymatous cells, at least in the upper part of the leaf, that are prorate or with distinct papillae at distal or proximal cell ends. As defined here, the Helodiaceae includes four genera that can be recognized in the following key: I. Plants autoicous; paraphyllia filiform-branched ...... Helodium I. Plants dioicous; paraphyllia lanceolate or lanceolatc-ciliatc ...... 2 2. Alar cells inflated in well-marked groups ...... Palustriella 2. Alar cells not differentiated ...... 3 3. Median cells of stem leaves isodiametric with single, spinose papillae ...... Bryochenea 3. Median cells of stem leaves prosenchymatous with 2- 3 papillae at cells ends ...... Actinothuidium T definitely exclude from the Helodiaceae the monotypic genus Bryonoguchia Twats. & Inoue that was placed in the subfamily Helodioideae by Fleischer (1923) and Brotherus (1925) as Tetracladium (Mitt.) Fleisch. B. moelkenboerti (Sande Lac.) Iwats. & Inoue, the only species of this genus, possesses irregularly leafy paraphyllia (Fig. 7: I) but filamentous at margins. The cells of the paraphyllia are partly papillose, and the filaments at margins are strongly papillose and short-celled, truncate and bipapillose at the apices and are exactly like those of Thuidium. The general appearance of this moss is also thuidioid, not helodioid. The bitypic genus Hylocomiopsis Card. is also excluded from the family and its peristome structure precludes closer affinity to the Thuidiaceae - Helodiaceae complex. The Helodiaceae are doubtless related to the Thuidiaceae and both families probably evolved from a common thuidioid stock. However, another very close relative of the family seems to be also the Hylocomiaceae, a family that has traditionally been associated with hypnoid mosses (Fleischer 1914, 1923; Brotherus 1925; Buck & Vitt 1986; Rohrer 1985a, b). In my opinion this family is not closely related to the Hypnacanae but it has an alliance with the superfamily Leskeacanae. The Helodiaceae and Hylocomiaceae share many important, structural characters that are essential for phylogenetic speculations and these taxa seem closer to each other than either family is to any of the families within the Hypnales. As presently understood (Rohrer 1985a, b) the Hylocomiaceae is an unnatural and polyphyletic taxon and most, if not all, of the taxa without paraphyllia should definitely by excluded from it. The paraphyllia in Hylocomium and leaf areolation (Fig. 7: 9- 10) of prosenchymatous cells are exactly the same in both families, and the types of paraphyllia in the Helodiaceae can be readily classified in the types of paraphyllia recognized for the Hylocomiaceae (Noguchi 1972). Of many structural characters separating both families, the most essential is the different costa condition. The Helodiaceae is without exception a unicostate taxon, while the Hylocomiaceae is predominantly a bicostate taxon. However, recent discovery in the Himalayan region the unicostate genus Miehea that is undoubtedly a hylocomioid moss (Ochyra \989) as well as the frequent occurrence of unicostate leaves in the genus Hylocomiastrum Broth., in particular in H. pyrenaicum (Spruce) Broth. and H. himalayanum (Mitt.) Borth., indicate that the Hylocomiaceae is a derived family that might have evolved from the unicostate ancestral type. As well , I showed R. OCHYRA: Animadversions on the moss genus CralOneuron 231 that one can visualize very easily evolution of branched paraphyllia from the un branched type that is found in Miehea (Ochyra 1989). Thus, I would be strongly inclined to accept the idea that the Helodiaceae and Hylocomiaceae are closely related families that have common ancestors. The problem of the relationship of the Hylocomiaceae is herein mentioned only incidentally and it will be discussed at length elsewhere.

SPECIES EXCLUDED FROM CRATONEURON AND PALUSTRIELLA Apart from the above discussed species the traditional concept of Cratoneuron placed in this genus several species known mostly from the Southern Hemisphere. All these species are associated with wet or otherwise moist habitats and indeed can imitate such species as Cratoneuronfilicinum or Palustriella commutata. This is, however, only the superficial similarity and the lack of paraphylIia or their different type precludes the closer affinity of these taxa with either Cratoneuron or Palustriella. They are actualIy identical to various species of aquatic mosses, mostly of the Amblystegiaceae, and the following is a brief assessment of the taxonomic status of these species. I. Cratoneuron perplicatum and C. submersum Dusen (1903) described Hypnum perplicatum from material collected from Tierra del Fuego. He alIied it to HypnumJalcatum Brid. from which it is very di stinct by "its longer, quite glabrous and scarcely decurrent leaves with doubly broader and excurrent nerve, and by the absence of paraphylla (sic!)". Since its description the species has remained a poorly known taxon that was merely mentioned by Cardot (1908) and Kiihnemann (1938) in their reviews of austral South American mosses. Brotherus (1909) transferred the species to Cratoneuron without comment, while Roth (1908) placed it in the Sendtneri-Gruppe of Drepanocladus. The plants are large and robust, up to 10 cm long, brownish- to golden-green, glossy and grow in wide, rather loose mats. They have characteristic hooked appearance owing to strongly falcate leaves that are up to 5 mm long and gradualIy long subulate from a lanceolate base and non-decurrent. The margins are entire and the costa is very strong, brown, 1/2- 1/3 the width of the base and long excurrent. The lamina cells are linear-ftexuose, firm-walIed and the alar cells are smalI and form an inconspicuous, non-decurrent group. These characters are sufficient to indicate inclusion of Cratoneuron perplicatum in Drepanocladus as Roth (1908) did. The entire leaf margins and small alar cells ally it to the type section of Drepanocladus. The species resembles in many respects D. capillifolius (Warnst.) Warnst. from the Northern Hemisphere but it is at once distinct in its smalI alar cells. The species is very distinct and it matches perfectly the type material of D. longifolius (Wils. ex Mitt.) Broth. ex Par. This species was described by Mitten (1869) as Amblystegium longifolium Wils. ex Mitt. from material collected from the Falkland Islands by Lechler and 1. D. Hooker. It is interesting to note that specimens gathered by 1. D. Hooker were originally named HypnumJalcatum (Wilson & Hooker l847a). The second species mentioned in the heading, Cratoneuron submersum Herz., was 232 Journ. Hattori Bot. Lab. No. 67 I 989 originally described by Herzog (1916a) from high elevations in the Andes of Bolivia. He considered it to be closely related to Cratoneuron fa/catum. This species also perfectly agrees in all characters with the type material of D. longifolius. Unlike other species of Drepanoc!adus, D. longi{o/ius shows astonishingly little morphological variation and it is a very distinct species that is widely distributed in southern South America and scattered at higher elevations in the northern Andes. The species is very distinct and in my opinion does deserve recognition as a separate species contrary to the statements of Dixon (1929), Sainsbury (1955) and Smirnova (1956). Drepanocladus longifolius (Wils. ex Mitt.) Broth. ex Par., ColI. 10. 1909. Hypnum perpii('(Jlum Duscn in Scott, Rep. Princeton Univ. Exp. Patagonia 1896--1899 Botany 8(1): 104, Fig. 26 & Tab. 11, fig. 7. 1903. - Drepallocladus perplicalus (Dusen) G. Roth, Hedwigia 48: 157. 1908. - Craloneuron perplicalllm (Dusen) Broth. in Engl. & Prantl, Nat. Pflanzenfarn. \(3): 1236. 1909. Type: Patagonia orientalis ad Cape Fairweather in paludosis [Lectotype (selected here): "Princeton Scientific Expedition to Patagonia 1896--1897 . P-45. Hypnurn perplicaturn Dus. Small brook near Cape Fairweather, Patagonia austr. coIl. J. B. Hatcher, August 7, 1896" - NY!; isolectotypes: H-Broth!, Nyl, s! Syntypes: (I) ditto, No. P-115 Nyl , (2 ) ditto, No. P-83 H-Broth!. SI , (3) ditto. No. P-9 - SI), syn. novo Crmonellron slIbmerswn Herz., Biblioth. Bot. 10\: 145. 1916. Type: [Bolivia) Untergetaucht in Moortumpeln des oberen Montehuaikotales, ca. 3900 rn, No. 2625 [Holotype: JE-Herz!; isotypes: BM!, FH-Fleisch!. H-Broth!, M! , s!), syn. novo 2. Cratoneuron sordidum and C. drepanocladioides M liller (1856) described C. sordidum (c. Muell.) Broth. as Hypnum sordidum from material collected from Mexico. In the original diagnosis he compared it to H. uncinatum and H. commUlatum with which it shares the overall appearance of the plants but differs in leaf shape. Later Mitten (1869) transferred the species to Amblystegium and placed it in the group of species that now constitute the genus Drepanoc!adus, and Brotherus (1908) suggested its alliance with Cratoneuron. Despite his clear statement that no original material was seen by himself, I located in his personal herbarium at Helsinki a specimen that is evidently a portion of the original collection. Although the material is rather scanty it was possible to examine all characters that are essential for correct identification of this species. The plants are relatively robust and golden-brown. The leaves are patent, strongly falcato-secund, ovate-lanceolate and gradually tapering into acumen. The costa is strong and extends into the acumen but it is not excurrent. The basal cells of leaves are rhomboidal, relatively firm-walled and porose, and the angular cells are rather thin-walled, pellucid and are differentiated into small auricles that do not reach the costa, while the upper lamina cells are linear-flexuose. No trace of paraphyllia on the stem could be detected. All these characters fit excellently into the concept of Drepanocladus sendtneri (Schimp.) Warnst. and accordingly C. sordidum must go into synonymy with this species. Cratoneuron drepanocladioides was described by Brotherus (1906) from material collected from the sub-Antarctic island of Possession (Crozet [slands) during the German South Polar Expedition of 1901 - 03 under the command of E. von Drygalski. The specific name of the moss alludes to the drepanocladioid habit of the plants and the salient overall similarity to Drepanocladus aduncus has been stressed in the R. OCHYRA: Animadversions on the moss genus CralOneuroll 233 protologue. The plants are medium-sized with nice golden tinge and irregularly branched. The leaves are strongly falcato-secund, with very distinct, small auricles that are very distant from the costa and are composed of yellowish-brown, incrassate and porose cells. The costa is strong and ceases in the acumen. The paraphyllia are entirely lacking. Like the preceding species, C. drepanocladioides cannot be separated from D. sendtneri and consequently both names must be considered synonymous. Drepanocladus sendtneri (Schimp.) Warns!., Beih. Bot. Centralbl. 13: 400. 1903. Hypnum sordidum C. MueH., Bot. Zeit. 14: 457. 1856. - Amblyslegium sordidum (c. Muell.) Mitt., J. Linn. Soc. Bot. 12:571. 1869. - Cratoneuron sordidum (C. Muell.) Broth. in Engl. & Prantl, Nat. Pflanzenfam. 1(3): 1032 . 1908 . Type: Mexico, unde Miquel habuit, qui nobiscum communicavit [Lectotype (selected here): "Specimen origin. Hypnum sordidum C. Mull. Mexico. communicavit Miquel ad C. Mull. Herb. Schliephacke" - H-Broth!], syn. novo CralOneuron drepanocladioides Broth. in Dryg., Deutsch. Sudpolar-Exp. 8: 94, 8 f 3. 1906. Type: Crozet-Gruppe: Possession-Insel (Vanhoffen) [Holotype: H-Broth!]. syn. novo 3. Craloneuron mendozense This species was described by Herzog (1916b) from a single gathering he made in Argentina in 1908. When describing this new species, this author stated that it differs in some details from the type of the genus Cratoneuron, expecially in the structure of the costa, but the material is too scanty and sterile to describe it as a new genus. The type material of C. mendozense includes medium-sized plants that are soft in texture, yellow-green and heavily encrusted with lime. The leaves are falcate-secund to almost straight, often rugose at the apex, erect-spreading, cordate at base and somewhat decurrent, gradually tapering to the long acuminate apex and have entire margins. The costa extends usually to mid-leaf, sometimes more above and reaches ca 50.um at base and is frequently branched in the upper part. The angular cells are enlarged, hyaline and form distinct, pellucid and inflated auricles that reach almost to the costa. The lamina cells are linear-rhomboidal, 45- 65.um long, and the paraphyllia are wanting. The above characters are all typical of Drepanocladus aduncus and C. mendozense must be reduced to synonymy with it. In fact Herzog originally intended to describe this material as a new species of Drepanoc/adus as visible on the label of the original material in his personal herbarium in Jena. The variable costa condition might suggest polygamus (B., S. & G.) Kanda that has additionally well-marked alar cells. However, this species can be easily distinguished by its straight, never falcate, leaves with channelled acumen. Drepanocladus aduncus (Hedw.) Warnst., Beih. Bot. Centralbl. 13: 400. 1903. Cratoneuron mendozense Herz., Hedwigia 57: 249. 1916. Type: An den Rii ndern kleiner Sturzbiiche bei Puente Inca (Prov. Mendoza), ca. 2900 m, mit Hygrodicrallum bolivianum f. brevl/olia und var. latifolium; leg. Th. Herzog, 1908 [Holotype: JE- Herz!]. , syn. novo 4. Hypnum filicinum var. minus, Cratoneuron kerguelense and C. arcticum Hypnum filicinum var. minus Wils. & Hook. f. in Hook. f. is a poorly known taxon and apparently it has never been studied since its erection (Vitt 1979). Until now it is held in the traditional nomenclatural combination and nobody transferred it to Cratoneuron. The variety was described by Wilson and Hooker (1847b) from 234 Journ. Hattori Bot. Lab. No. 67 I 989 material collected by J. D. Hooker from the Auckland Islands during the course of the Antarctic voyage on the ships Terror and Erebus in 1839- 1843 under the command of Captain lames C. Ross. In the original diagnosis Wilson and Hooker (1847b) characterized this variety as having slender stems and erect, lanceolate leaves but "though the habit is more delicate than that of British specimens, we are unable to separate this moss specifically from H. jilicinum, of which a perfectly intermediate variety occurs in the Falkland Islands". The plants used for the description of this variety are small, golden-brown, straight and shiny. The leaves are erect, lanceolate and gradually long acuminate and have entire or faintly serrulate margins. The costa is strong and percurrent. The upper cells are oblong-rhomboidal, while the basal cells are shortly oblong to subquadrate in the basal angles and thick-walled. For a long time the identity of this moss was puzzling to me but finally I found it fits into the concept of Pseudoleskea chilensis (Lof.) Ochyra. This species is one of the most confusing of antipodal mosses. Owing to its great variability it was described for over twenty-five times as a separate species or a variety in South America, South Africa, Antarctica and sub-Antarctica. Although P. chilensis has the strong and percurrent costa, it cannot be mistaken for Cratoneuron jilicinum, the lack of pellucid alar cells and lanceolate, never contracted at the base \eaves immediately di stinguishing it from that species. In addition, in many populations one can find easily filiform, uniseriate paraphyllia scattered on the stem that are totally different from the lea fy paraphyllia of C. jilicinum. The above remarks refer also to Cratoneuron kerguelense (Mitt.) Broth. This species was originally described by Mitten (1879) as Amblystegium kerguelense from material collected from the Kerguelen Islands during the Transit of Venus Expedition in 1874-75 by E. A. Eaton. In addition, Mitten (1879) cited in the protologue other specimens collected in this archipelago by J. D. Hooker and reported in his "Flora Antarctica" as Hypnum jilicinum and H. serpens (Wilson & Hooker I 847a). The type specimens of this species are inseparable from other sub-Antarctic and South American plants of Pseudoleskea chilensis and show the perfect correspondence in all critical and taxonomically important characters. Accordingly both Hypnum jilicinum var. minus and Amblystegium kerguelense are reduced to synonymy with P. chilensis. Cratoneuron arclicum is a poorly understood North American moss. Steere (1959) originally described it from several specimens collected in the Nearctic and the sterile material considerably hindered discovery of the proper placement for this distinct and unique moss. At the beginning Steere assumed the genus Lescuraea B., S. & G . could be the good placement for it but he left this idea at Lawton's suggestion. Finally he placed this moss in Craloneuron and it was rather as much from desperation as from conviction. Later Crum et al. (1965) synonymized C. arcticum with Amblyslegium varium (Hedw.) Lindb. but Brassard (1971) stated that C. arcticum is inseparable from C.jilicinum and stunted nature of the plants is doubtless conditioned by severe climatic conditions in the Arctic. On the other hand, Kuc (1973) maintained C. arclicum as a distinct species and according to him it was closely related to C. curvicaule. He reported one additional collection of this moss from Axel Heiberg Island in the R. OCHYRA: Animadversions on the moss genus Craloneuron 235

Canadian Arctic Archipelago, and Holmen and Scotter (1971) and Steere (1978) found it on the mainland in the northern part of the Northwest Territories in Canada and in Arctic Alaska, respectively. Although little can be added to the excellent description and illustration of C. arcticum, the problem of its disposition was compounded because the plants were known only in a barren condition. Luckily I have located two collections bearing fully mature capsules from Wrangel Island in Arctic Siberia (10 August 1985, Polozova s. n. - LE, KRAM) and from Banks Island in the Nearctic (Steere 63- 787 - NY). The species has perfect hypnaceous peristome which allies it to the hypnobryalean mosses. Un­ fortunately this is insufficient to decide the generic placement of a moss. Examination of many fully developed plants of C. arcticum from the Arctic led me to the conclusion that they are inseparable from the antipodal Pseudoleskea chilensis!!! I cannot find any discernible differences between arctic and antarctic populations and consequently I consider both species to be identical. The conspecificity of both species increased the number of bipolar moss species by one very distinct and peculiar species. This is very interesting and special case of the bipolar distribution pattern since the species is very widespread and abundant in the Southern Hemisphere, while it is very uncommon and scattered in the Northern Hemisphere. The only other similar case known to me is the distribution of Andreaea mutabilis Hook. f. & Wils. (M urray 1988). Details of the taxonomy and geographical distribution of P. chilensis will be presented el sewhere and herein I provide only the necessary synonymy that are associated with the problem of the heterogeneity of Cratoneuron. Pseudoleskea chilensis (Lor.) Ochyra, 1. Bryol. 14: 459. 1987. Hypnumfilicinum Hedw. var. minus Wils. & Hook. f. in Hook. f. ["-or"], Fl. Anla rct. 2: 14 1. 1844. Type: Lord Auckl and's group; in woods by the banks of streams [Lectotype (se lected here): " Hypnum fi licinum var. p mi nor Lord Auckland's Islands. Antaret. Exp. 1839-1843. Or Lya1l 26" - BM'; isolectotypes: BM-Hook! , BM- Wils (3 specimens)!, BR!, E!, H-SOL! , MA NC H-Spruce', MICH! Syntypes: (I ) ditto, Lya ll 33 - llM! , BM -Wils! , BM -Hook!, (2) ditto, Lyall 37 - BM-H ook! , BM-Wils!], syn. 11 0V. Amhlystegium kerxuelense Mitt., Phil. Trans. R. Soc. London 168: 37. 1879. - Cratoneuron kerguelense (Mitt.) Broth. in Engl. & Pra ntl, Nat. Pnanzenfam. ed. 2, 11 : 334. 1924. Type: Christmas Harbo ur, Hooker. Near Swain's Bay, Ealon [Lectotype (selected here): " Near Swain's Bay, Kerguelen. Rev. E. A. Eaton. Transit Venus Exp. 22. XII. 1874" - BM! Syntypes: (I) " No. 266. Hypnum filicinum var. 2 Kerguelen's Land. Antarct. Exp. 1839- 1843. J . O. H ." - BM-Hook! ; (2) " Hypnum serpens var. Kerguelen's La nd 752. Antarct. Exp. 1839- 1843 JOH 267" - BM-Hampe!, BM-Hook (2 specimens)!, BM-WILS! ; (3) "Hypnum serpens var. 2 Kerguelen's Land 753. Antarct. Exp. 1839- 1843 JOH 268" - BM! , BM-Hampe!, BM-Hook (2 specimens)! , NY-Mitt!], syn. novo Cratoneuron arelicum Steere, Nat. Mus. Canada Bull. 164: 97, I. 1960. Type: Canada, Ellesmere Island: in clay-shale, damp moss-tundra, calcareous, Cape Belknap, July 17, 1955, Schusler 35285 [Holotype: NY'], syn. 1I0V.

5. Cratoneuron la t ifolium , C. oedogonium and C. papil/osum Cratoneuron latifolium was originally described by Okamura (1915) as Hygroamblystegium latifolium Okam. and later Brotherus (1925) transferred it to Cratoneuron. The species is considered to be identical to Cratoneuron filicinum by Kanda (1975). Cratoneuron oedogonium, originally described by Muller (1897) from Bolivia, was 236 Journ. Hattori Bot. Lab. No. 67 I 9 8 9 transferred by Brotherus (1909) to Eurhynchium B., S. & G. However, I found that the best placement for this eurhynchioid moss characterized by distinctive heterophylly is in Kindbergia Ochyra (Schultze-Motel & Menzel 1987). Craloneuron papillosum Warnst. was described for the first time by Warnstorf (1915) from a single gathering from Japan. Reimers (1937) considered it synonymous with Haploc/adium microphyllum (Hedw.) Broth.

ACKNOWLEDGMENTS. I thank the directors and curators of the following herbaria for loaning specimens without which research of this nature would not be possible: B, BM , BR, E, F, FH, G, H, JE, LE, M, MANCH, MICH, MO, NY, PC, S, ups and w. As usual, my wife completed illustrations and I gratefully acknowledge her aid.

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