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Animadversions on the Moss Genus Cratoneuron (Sull.) Spruce

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fourn. Haltori BOI. Lab. No. 6 7: 203- 242 (Dec. 1989) ANIMADVERSIONS ON THE MOSS GENUS CRATONEURON (SULL.) SPRUCE R YSZARD OCHYRA 1 ABSTRACT. A review of taxonomic problems in Craloneuron (Sull. in Gray) Spruce is provided and all taxa that had previously been included in the genus are discussed at length. Cralonellron, since its inception, has been an unnatural and polyphyletic taxon and all the species of the genus, except C. jilicinum (Hedw.) Spruce, are transferred to other genera, leaving Craloneuron monotypic. Hypnumfallax Brid., Craloneuron punae C. M uell. , C.formosanum Broth., C. procerum (Dix .) Broth. and C. subcurvicaule P. Varde are reduced to synonymy with C.jilicinum. One distinct variety, C.jilicinum var. alrovirens (Brid.) Ochyra, is recognized and it includes the plants that have frequently been named C. jilicinum var. fallax (Brid .) G. Roth by most workers, and Hypnum vallis-clausae Brid. and Amb/ys/egium formianum Fior.-Mazz. are identical to this variety. The monotypic family Cratoneuraceae Moenk. is restored to accommodate Cratoneuron and it is considered to be related to the Leskeaceae. The taxonomic status of Hypnum curvicaule Jur., a species that for a long time has been treated as C. Jilicinum var. curvicaule (Jur.) Moenk., is discussed at length. This taxon is retained in the Amblystegiaceae in the new genus Callialaria Ochyra as C. curvicaulis (lur.) Ochyra. Craloneuron commulalum (Hedw.) G . Roth and C. decipiens (De No!.) Loeske are transferred to the newly erected genus Paluslriella Ochyra that is allied to Helodium (Sull.) Warnst. and its relatives as P. commulata (Hedw.) Ochyra and P. decipiens (De Not.) Ochyra. Several varieties of the former C. commulalum and C. decipiens are given formal names in Pa/uslriella, including P. commulala var. diaslrophylla (Brid.) Ochyra, P. c. var.falcala (Brid.) Ochyra, P. c. var.flucluans (B., S. & G .) Ochyra, P. c. var. irrorala (Mikut.) Ochyra, P. c. va r. minor (Lesq.) Ochyra, P. c. va r. pseudodecipiens (Amann) Ochyra, P. c. var. plychodioides (G. Roth) Ochyra, P. c. var. su/cala (Lindb.) Ochyra and P. decipiens var. napaeiformis (Schiffn.) Ochyra, while Cratoneuron commutalovirescens Amann and C. su/caloirrigalllm Meyl. are tentatively reduced to the status of varieties within P. commutata as P. c. var. commulalovirescens (Amann) Ochyra and P. c. var. su/calOirrigata (Meyl.) Ochyra. A brief assessment of the Thuidiaceae is presented and its subfamily Helodioideae Fleisch. is raised to family rank as Helodiaceae (Fleisch.) Ochyra. Apart from He/odium and Paluslriella it includes Bryochenea Gao & Chang and Actinothuidium (Besch.) Broth., while Bryonoguchia Iwats. & Inoue and Hylocomiopsis Card. are excluded from the fam il y. A key for separation of the genera of the Helodiaceae is given. Other heterogeneous elements in Craloneuron are a lso discussed. C. perplicalum (Dusen) Broth. and C. submersum Herz. are identical to Drepanocfadus /ongifolius (Mitt.) Broth. ex Par.; C. sordidum (c. Muell.) Broth. and C. drepanocfadioides Broth. in Dryg. are inseparable from Drepanocfadus sendlneri (Schimp.) Warnst. ; C. mendozense Herz. is placed in synonymy with Drepanocfadus aduncus (Hedw.) Warns!.; and Hypnum filicinum var. minus Wils. & Hook. f., Craloneuron kerguelense (Mitt.) Broth. and C. arclicum Steere are considered to be synonymous with Pseudoleskea chilensis (Lor.) Ochyra. The conspecificity of the latter species has considerably extended the range of P. chilensis that is now established as a bipolar disjunct. BRIEF HISTORICAL A CCOUNT Sullivant (\856) distinguished Cratoneuron as a section of the then all­ encompassmg Hypnum Hedw. to accommodate Hypnum filicinum Hedw. and H. I Laboratory of Bryology, Institute of Botany, Polish Academy of Sciences, Lubicz 46, PL-31-5 12 Krak6w, Poland. 204 Journ. Haltori Bot. Lab. No. 67 1 989 commutalum Hedw., two widely distributed species that are associated with wet or otherwise moist habitats. The section was subsequently elevated to subgeneric rank within Hypnum by Schimper (1860) and Spruce (1867) promoted it to the generic status. There is, admittedly, no description of the genus in Spruce's (1867) publication but Sullivant is cited for the generic name in the genus heading and this is an indirect reference to Sullivant's (1856) work. Since the genus was originally composed of two species, Grout (1931) lectotypified Cratoneuron (Sull. in Gray) Spruce with C.jilicinum (Hedw.) Spruce. The principal criteria for its recognition were the leaves with a strong costa, inflated and pellucid alar cells and the presence of paraphyllia on the surface of the stems and branches. Until the end of the nineteenth century Craloneuron was not accepted as an independent genus. Most bryologists considered it only as a section or subgenus of Hypnum (Schimper 1860, 1876; Hampe 1873; Limpricht 1876; Jaeger & Sauerbeck 1879; Hobkirk 1884; Boulay 1884; Lesquereux & James 1884; Hazslinszky 1885; Klinggraeff 1893) or Amblystegium B. , S. & G. (De Notaris 1867, 1869; Venturi & Bottini 1884). Finally, Roth (1899) resurrected Cratoneuron as a genus of its own and placed it in the newly established family Amblystegiaceae, along with Amblyslegium, Campylium (Sull.) Mitt., Drepanocladus (c. Muell.) G. Roth and Calliergon (Sull.) Kindb. He divided the genus into two sections: (1) sect. Filicina (Brid.) G . Roth (= Cratoneuron) including C.jilicinum, C.formianum (Fior.-Mazz.) G . Roth and C. curvicaule (Jur.) G. Roth and (2) sect. Sulcata G . Roth with C. commutalum (Hedw.) G . Roth, C. falcatum (Brid.) G. Roth, C. sulcatum (Lindb.) G. Roth and C. irrigatum (Zett.) G. Roth. Since then Cratoneuron has gained a wide acceptance of the bryologists who mostly ascribed to G. Roth its status as a genus distinct from all other amblystegiaceous mosses and the name has appeared in almost all floras, manuals, checklists and taxonomic revisions. A few authors have not accepted the placement of Cratoneuron in the Amblystegiaceae and in order to emphasize its distinctiveness they positioned it in the separate family Cratoneuraceae Moenk. (Moenkemeyer 1914, 1927; Podpera 1954; Pilous & Duda 1960; Schlyakov 1961; Abolin 1968; Melnichuk 1970). Since its inception, some 27 species have been validly and legitimately included in Cratoneuron. However, some of these have been reduced to the status of infraspecific taxa and in the result only 18 species still remain in the genus (Wijk et al. 1959, 1969). Closer examination of all species that are currently placed in Cratoneuron reveals that the genus is an increadibly artificial taxon that groups many unrelated species on the basis of superficial similarities of gametophytes, especially falcate leaves, strong costa, inflated and pellucid auric1es and the presence of paraphyllia. Most of these are concentrated around two leading species of the genus, namely C. jilicinum and C. commulalUm. A few extra-Holarctic taxa are poorly known and apparently they have never been examined and tested in order to establish their true relationship. Historically, two lines of interpretation of the interrelationships between C. jilicinum and C. commulalum can be recognized. Most workers followed the concept of Sullivant (1856) and considered these species as closely related taxa within the subdivision Cratoneuron R. OCHYRA: Animadversions on the moss genus Craloneuron 205 in Hypnum. Even prior to recognition of Cratoneuron, both C. filicinum and C. commutatum have often been placed together in various subdivisions of Hypnum which have been given different names, for instance sect. Filicoidea Brid. (Bridel 1801), sect. Filicina (BrideI1812, 1819, 1827; Hlibener 1833; Rabenhorst 1848), sect. Hypnafilicina (Bruch et al. 1854), subsect. Drepanophyllaria C. Muell. (Muller 1848- 1851), sect. Amphimalla Wallr. (Wallroth 1831), sect. Adunca Brid. (De Notaris 1838; Spruce 1849), "Gruppe" Hypna adunca (Girgensohn 1860). Adherents of the second approach considered C. filicinum and C. commutatum to be remotely related taxa and therefore placed them either in two different genera or in various sections within a larger genus. Most often C. filicinum was considered as a species of Amblystegium whereas C. commutatum was retained in the section or subgenus Cratoneuron within Hypnum (Milde 1869; Hartmann 1871; Husnot 1892-1894; Limpricht 1904; Paris 1904, 1905; Herzog 1906; Dixon & Jameson 1924). On the other hand, Lindberg (1879) and Braithwaite (1893- 1905) placed C. filicinum in the subgenus or section Euamblystegium (=Amblystegium) and C. commutatum in the subgenus or section Drepanocladus within the genus Amblystegium, whereas Kindberg (1894, 1896) kept C. filicinum in the section or subgenus Cratoneuron and established the new section and subgenus Alaria for C. commutatum within the genus Hypnum. Finally, some workers included C. filicinum in Hygroamblystegium Loeske that is a segregate of the large genus Amblystegium and left Cratoneuron oligotypic for C. commutatum and the relatives (Loeske 1907a, c; Brotherus 1908). Although the problem of the interrelationships of Cratoneuron species and seemingly related taxa of the Amblystegiaceae and Thuidiaceae was discussed by several authors (Loeske 1907a, c; Moenkemeyer 1911; Dietzow 1914; Fleischer 1923), the final conclusions seem to be unsatisfactory and the genus in the current understanding is very unnatural and polyphyletic. The present account is a new attempt to revise the old question of the span and affinity of Cratoneuron. A NEW GENERIC DEFINITION OF CRATONEURON On the basis of extensive
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    Catalog of the Colorado Flora: a Biodiversity Baseline Mosses: Weber and Wittmann, electronic version 11-Mar-00 Amblystegiaceae Amblystegium Bruch & Schimper, 1853 Amblystegium serpens (Hedwig) Bruch & Schimper var. juratzkanum (Schimper) Rau & Hervey WEBER73B. Amblystegium juratzkanum Schimper. Calliergon (Sullivant) Kindberg, 1894 Calliergon cordifolium (Hedwig) Kindberg WEBER73B; HERMA76. Calliergon giganteum (Schimper) Kindberg Larimer Co.: Pingree Park, 2960 msm, 25 Sept. 1980, [Rolston 80114), !Hermann. Calliergon megalophyllum Mikutowicz COLO specimen so reported is C. richardsonii, fide Crum. Calliergon richardsonii (Mitten) Kindberg WEBER73B. Campyliadelphus (Lindberg) Chopra, 1975 KANDA75 Campyliadelphus chrysophyllus (Bridel) Kanda HEDEN97. Campylium chrysophyllum (Bridel) J. Lange. WEBER63; WEBER73B; HEDEN97. Hypnum chrysophyllum Bridel. HEDEN97. Campyliadelphus stellatus (Hedwig) Kanda KANDA75. Campylium stellatum (Hedwig) C. Jensen. WEBER73B. Hypnum stellatum Hedwig. HEDEN97. Campylophyllum Fleischer, 1914 HEDEN97 Campylophyllum halleri (Hedwig) Fleischer HEDEN97. Nova Guinea 12, Bot. 2:123.1914. Campylium halleri (Hedwig) Lindberg. WEBER73B; HERMA76. Hypnum halleri Hedwig. HEDEN97. Campylophyllum hispidulum (Bridel) Hedenäs HEDEN97. Campylium hispidulum (Bridel) Mitten. WEBER63,73B; HEDEN97. Hypnum hispidulum Bridel. HEDEN97. Cratoneuron (Sullivant) Spruce, 1867 OCHYR89 Cratoneuron filicinum (Hedwig) Spruce WEBER73B. Drepanocladus (C. Müller) Roth, 1899 HEDEN97 Nomen conserv. Drepanocladus aduncus (Hedwig) Warnstorf WEBER73B.
  • AMBRA1 Controls Plant Development and Senescence in Physcomitrella Patens

    AMBRA1 Controls Plant Development and Senescence in Physcomitrella Patens

    Presentation type: Oral Presentation, Poster Presentation (underline the preferred type) AMBRA1 controls plant development and senescence in Physcomitrella patens. Alessandro Alboresi1, Jessica Ceccato1, Tomas Morosinotto1, Luisa Dalla Valle1. The first one should be the presenting/corresponding author (underlined) 1Dipartimento di Biologia, Università di Padova, Via Ugo Bassi 58/B, 35121, Padova ([email protected]; [email protected]; [email protected]) Autophagy is a universal mechanism that in plants control development, resistance to stresses and starvation. The role of autophagy is possible thanks to the programmed degradation of cell material that is delivered to the vacuole where hydrolases and proteases are localized. So far, many autophagy-related proteins (ATGs) have been identified. Some of them are universal, some are either specific to animals, plants or yeast. ATG protein complexes govern autophagosome initiation, nucleation, expansion, and maturation. In particular, the regulation of nucleation by the ATG6 (Beclin-1 in mammals) complex has not been well defined in plants. Here we described the study of the Activating Molecule in Beclin 1-Regulated Autophagy (AMBRA1) protein, recently identified in mice and then characterized in our department in zebrafish and in the non-vertebrate chordate Botryllus schlosseri. In animals AMBRA1 is a positive regulator of autophagy that binds Beclin-1 upon autophagic stimuli. AMBRA1 is a large intrinsically disordered protein, able to bind other regulatory partners involved in cell processes such as autophagy, apoptosis, cell proliferation, development and cancer. AMBRA1 sequence was found in plant genomes and we are studying its function in Physcomitrella patens where two lowly expressed genes are present, AMBRA1a and AMBRA1b.