Published by Maney Publishing (c) British Bryological Society J. varieties Frey, var. with varieties was muralis, strongly tion cluding have 21 Iberian the n more Schiffner lectotype. original study and incana name Geneva One (Amann, (Casas, (hair-point de (Schimper, = Bryol. The No In In The la June 50/52) varieties is of designated aestiva, certainty of the Frontera the type 1983). was or proved in probably purpose type B., baetica two var. them Peninsula 1981). description. muralis emarginate (G). (1992) is the have practice less Iberian 1847, The (1913) 1918), can original prompted S. specimen uncertain length, of cytotypes 1876), muralis, Smith aim widely obcordata & continuous. (pr. been T. be from to 17, Of (holotype of G., by more taxonomic Schimper Hedw. (Reimers, J. for muralis being distinguished Peninsula, (Musci, this because 275-283 warrant two these Hyeres, (Schimper, colour (1978) GUERRA, description Hedwig var. described each leaves, recognized The example, because of by var. is paper than localities University to T. and obcordata not is we the Hedw. other other in of noted aestiva muralis determine Even morphologically the leg., just terrasse 1941; the specific justified believe Herb. is (1801), baetica, knowledge the six of ): (Wijk to is 1876; only morphological var. (In and a the R. status of were in var. that varieties , present lectotype now habitat Brid. Boudier, of INTROOUCTION Hedw. (Schimp.) Oliva, Europe ponte M. continuous T. from baetica that du in but et recognition. Murcia, baetica (Newton, both the Casas the mentioned recognized part al., ROS muralis leaf Grand ex with 275 we that only the 'var. form'. of the Guadalaviar true var. Cordoba), of described Hedw., 1969). by by (Smith, 1988). in size, & were a Casas results and Limpr. T. var. different Tortula Murcia, other three aestiva its muralis very their taxonomic variation BM), Hotel, wide Oliva, 1968). a var. Correct muralis stem peculiar J. and Some as muralis: able comparative Tortula & var. of S. variable size; infraspecific can 1978; range from and T. was Oliva, samples is obcordata is a Spain Hedw., 1982), height, CARRI6N Reuter prope ploidy to distinctive baetica brevissima, comparative of be muralis determination therefore have hence, of var. also type status Nyholm, study samples of them the distinguished muralis Espana, the taxon, variation Valenciam rupestris and from size leg.) levels available been Sp. of var. Casas relevant taxonomic the are of taxa (Schimp.) papillosity the in Muse. of collected these obcordata both selected does as 1956; var. very df recognized, lectotype fa. Sevilla, study study its (n the & of type K. they described which of = appearance for Hispaniae, gypsophila Oliva, are capsule) T. difficult not characters taxa. 123.1801, easily F. 26/27 Frahm the based recogni display. study. variety, muralis and Schultz Limpr. in Moron as on fit many by other from BM. This var. and and the the the the in- on by its to & is .. Published by Maney Publishing (c) British Bryological Society in upper The hyde leaves glutaraldehyde 276 EM Anatomy above. calculated. aestiva 7.75 between in tome embedded muralis identical Rurales acetone and somewhat rubra separate ent baetica, intermedia character Size In included mucronifolia of Detail Similarly, (1980) F F Leaves Data Spurr the cell the or or shoots photographed 10c of biometric fJ-m with and Mitt. scanning were genus half transmission of varieties size After the as Hedw. obtained var. and (mean electron and to T. De resin the the the were of the stained 0.1 to smaller a leaf of in (Brid.) of geheebiaeopsis thos'e of (Lightowlers, made liquid the obcordata the haploid character leaf Tortula, washing variety Not. differentiate present M epoxy Schwaegr. each each the and cells washed study length and and electron cacodylate leaf obcordata mean microscope during nerve, given with using than De three with CO, 2 stained sample It leaf nerve electron T. resin. post-fixed cytotype baetica of study the = and Not. width uranyl also with and by those of vahliana a 12.2 a as the leaf sputtered were microscopy, taxa. and size JEOL Reichert 1985, Newton dehydration the (C. seen Sections with (mean between (see buffer var. has and distilled present RESULTS serves 8.25 cells fJ-m, and operated of microscopy, of acetate T. with Mull.) measured However, first shown in in muralis. the Appendix). 1986). toluidine T. JSM the J. (Schultz) subulata fJ-m. the at mean transverse involved length with (1968) 1 GUERRA princeps Jung n latter. % were order study to 4°C, METHODS middle water, similar = Broth. AND T-300 selected and other that These at in 26/27. osmic a distinguish width Two Ultramicrotome. = 250-300 submitted and 60 as made a it blue The (Fig. lead DISCUSSION of T. the 9.20 Hedw. Mont. fixed ET graded three ranges and from operated can The De taxa, kV. species values section, muralis adult importance the acid AL. = mean leaves citrate and 1) using Not., fJ-m, upper be 13.1 with length T.filaris samples A mean show and (Smith, of ethanol and in leaves are semifine to seen between thick length - mean has var. fJ-m). were glutaraldehyde, at a the and the before critical were leaf the means noticeably Reichert a and for 10-20 values been that certain were baetica gold in section (C. between same each series, cells cells fixed The of 1978) width width example, similarly transverse studying point the for there species Mull.) regarded kV. layer, taken for last has of of buffer in degree Jung is T. = upper - the lower Tortula of T. quite drying the each 3 11.75 T. been muralis, is are % Broth. as and from 5 impregnated material fixed muralis with muralis less Ultramicro- such between former cells glutaralde- sections mentioned by leaf than practically well of of distinct leaf observed used fJ-m) in 3 Kramer Spruce. overlap overlap a section in and cells differ- in which as 1000/0 those Zeiss as were var. var. was as and 30/0 the are T. T. of T. to in is a Published by Maney Publishing (c) British Bryological Society var. var. this 2c, the mamillae Fig. but these thickened taxa cells middle abaxial of baetica, (Fig. The The The In the d). obcordata 1. bear respect upper baetica they var. 2a, lack Relative cells leaf 10 nerve 14 abaxial 12 stereid 4 8 6 2 and epidermal with 3-4(5) b). in are baetica mamillae are considerably (Fig. third from (solid the the the In cell outline 10-12 numerous II more epidermal cells the upper number lower papillae. size 2c, of circles), the the cells, other 5 at d), (Fig. the in or is throughout other ventral the and third semicircular less the but o T. stereid throughout. o leaf, (Fig. increases uppermost two 2a), cells, 7 0 o TORTULA m. 0 middle two 0 cubical upper is 0 of var. taxa cells 0000 occasionally have than U-cfJ. -f) 2b). the which varieties. 9 cells o baetica eo the . part part third it considerably are (Fig. MURALIS leaf in ••• from In one • . -. is and 11 leaf the . in of usually markedly of the __ (open •• these ••• of the 2c, the mamilla the·upper The nerve the • other -- - - and 13 semicircular • the other leaf. VAR. d, circles). var. . leaf cells differences - elliptical • see leaf all Tortula outline. taxa in 15 ampullose baetica BAETICA (Fig. two and - arrows). the lack their W third are with = - muralis 17 taxa, lower thin 2a, width, from mamillae up considerably are walls to 20-30(40) are b). to the They walls with var. generally 19 the L the third, found the In = are base, length. muralis middle abaxial the 1(-2) are apex. and (Fig. w(~m) but thickened in (Fig. other not in fewer the ventral ampullose (squares), protuberant in var. 2a). to ampullose epidermal 2c, the walls two the baetica in In d). other cells, (Fig. base taxa var. T.m. 277 the are In in Published by Maney Publishing (c) British Bryological Society The ventral Laminal the usefulness papillae phylogenetic respect, mamillae and are Even and 278 Jarai-Komlodi, Bar mically 1984), Fig. baetica, Casas Tortula The = 2. var. highly middle sporoderm abaxial type 20 Light within et upper sculpturing, including IJ-m; cells is the baetica useful al., papillosity (Fig. (Fig. of that microscope significant in d, part. leaf differences surfaces, of 1990; foliar certain var. a implications 3a-d). 3e-h). shown the single features to 1975; Bar cells obcordata, Tortula Casas specific nerve papillosity = and micrographs genera 25 possess and The In Scott by the genus, IJ-m; between the the & the (Lewinsky, (Fig. alone (Steere, ornamentation middle upper rank. b, have Sergio, & of leaf processes var. var. single spores of Stone, the is 3d). present semi of part. the The baetica, considered leaf J. also muralis Pottiaceae var. 1940; GUERRA 1990). or fine var. Bar of most 1976; cells 1974; occasionally been transverse are baetica, middle strong = the baetica Kramer, is 20 and In frequent have made Stone, ET thus IJ-m. Pottiaceae Saito to general, recognized part. (Delgadillo, var. AL. evidence however, sections be and 4-5(6) identical Bar up 1980), 1988, obcordata. bifurcate & an type the = almost the of Hirohama, 25 important sometimes bi-or other to the 1989; IJ-m; of both (Mishler, is exine to 1975; support leaves. papillosity widely c, mamillae that exclusively trifurcate two surfaces var. On Carrion contributes Frey a, character of taxa muralis, both 1974; the Tortula exhibit recognized. 1986). the & in or treated of elevation et Kiirschner, the papillae dorsal middle the of the very muralis aI., Boros taxono- little adaxial In and a upper genus 1990; here high very part. and this var. Its its or of to & Published by Maney Publishing (c) British Bryological Society surface Fig. al., baetica, surface; processes electron-dense T. Saito princeps 3. 1990). Scanning and g, adaxial & var. nerve; of Hirohama obcordata, on the electron surface material b, the former var. of microscope basis adaxial (1974) baetica, nerve; species. of TORTULA of a surface e, perinic adaxial have secondary micrographs var. Surface-spore near muralis, previously surface MURALIS nature nerve; of patterns adaxial near leaf h, (McClymont V var. discriminated surfaces. AR. nerve; outlines surface obcordata, BAETICA of c, ornamentation a, var. near Tortula were & baetica, adaxial nerve; Larson, between also muralis surface. f, adaxial studied var. var. 1964; over T.muralis muralis, Bars surface; baetica, Carri6n in the = Tortula 10 abaxial adaxial d, main !.Lm. and 279 var. et Published by Maney Publishing (c) British Bryological Society 280 pilum-like, by are spores Brown or Perinic papillae that microscope pertain surfaces processes sporomorphs three b, Fig. and although Although Tortula nano-granulous proportionally Lewinsky 4. taxa. the other Transmission (Fig. electron-opaque & to are muralis shape of Nano-processes the and Lemmon, mature large all without conical some 4b, taxonomic were (1974), broader var. populations of higher see electron warts intrapopulational secondary baetica; elements poured spores (Larson, or previous arrow). 1980, in (n) rounded although or spinule-like var. microscope significance projections covering irregular c, of on baetica. var. 1981, studied are processes preparation. 1964; T. a muralis; and muratis formvar/carbon structural surface also J. micrographs Relative 1985), processes GUERRA nano-processes variation cover Reighard, are is of visible. d, debatable. of was very T. var. var. thickness perine internal the Nevertheless, interpretation muratis ET of of obcordata. variable. could was baetica As outer spores AL. taxonomic can 1967; coated of in observed, be polarity be exine Phascum surface showing var. are seen. (Fig. Bars Olesen distinguished. Baculate, (e) grid Lewinsky noteworthy baetica value. = was and 4a, is inner completely 1.3 proportionally and (Carrion displayed intine & b). f,Lm. inaccurate sporoderm gemmate, In Mogensen, viewed and These Perinic did (i) T. Nano-spinules on are are demonstrate muratis, et (Fig. by similar

elements variations under structure. the al., evident, because broader clavate, Tortula 4a-d).

1990) distal 1978; in the the the (p) a, J I I Published by Maney Publishing (c) British Bryological Society three var. the muralis, It Ecological muralis taxon Tortula On 7: Basionym: samples The Amann, staff We establish support Boros, material, 1971). Boudier, the baetica of the configuration assistance. as Tortula is On 104. var. a hypothesis France Budapest. genus Region interesting baetica special are distinct the of of A. should the J. muralis 1982. as P. muralis var. baetica which maintenance collected & the grateful var. (1918). other et to their Tortula. a (1988). Jarai-Komlodi, other behaviour This Tortula type has of de species, species. Dr muralis Electron obcordata retain of Suisse. Murcia that colonize Flore that and been taxonomic (Casas var. research hand, Martha the of Tortula hand, to in This, muralis leafpapillosity genetic ecological des var. sporoderm, varietal Cryptogamie obcordata curators Spain, T. observed and of and Mousses Microscopical & populations baetica M. brevissima walls and the together E. obcordata, was varietal var. TORTULA Oliva) DGICyT (1975). barriers Hedw. Newton status. were status characters var. supported and and characteristics de obcordata. growing and Bryol. ACKNOWLEDGMENTS (Casas seems la Guerra An Schiffner compared with baetica status rocks var. shown directors Suisse. CONCLUSIONS or MURALIS REFERENCES The T. isolate the atlas for SUMMARY from morphologically Lichenol. perhaps muralisvar. Service a & sufficient baetica mixed leaf in studied by revising for smaller of results Part & have (Pottiaceae, Oliva) by Therefore Spain nitro T. recent the Ros, VAR. with var. nerve of 2. T. are 9,219-230. muralis with be not Casas Lausanne. of Council philo muralis herbaria size to Guerra are European obtained identical (Project the obcordata stat. BAETICA T. regarded Murcia anatomy been baetica, warrant var. muralis Musci) English us completely of it & var. nov. intermediate is of sites. Oliva, var. the muralis found. & consulted PB90-0301-C02-01). considered in Culture baetica nouveau University from Ros. just two recognition upper that and var. baetica the manuscript Frequently spores. Acta taxa as These and taxa is recognition stat. muralis a overlapping (see pour and leaf distinct a biometric Bot. is for var. Akademiai form. described considered, that between nov. uncommon for Education Grant, data cells la of the Malacitana obcordata. in and bryoflore T. this the from technical order support muralis and loan of to taxon Kiad6, latter study in 1963, from var. var. that 281 the the all T. to of of de in Published by Maney Publishing (c) British Bryological Society Lightowlers, Hedwig, Delgadillo, Mishler, McClymont, Newton, Reighard, Lightowlers, Larson, Kramer, Frey, Frahm, Lewinsky, Smith, Saito, Nyholm, Schimper, Grant, Grant, Casas, Casas, Casas, Casas, Carrion, Brown, Brown, Reimers, Schiffner, Olesen, Brown, Scott, 282 new 41-47. to 245-303. Musci. 281-295. 265-276. Bryophytorum Iberica. stems Research 523-535. the Zechstein-Kyffhauser. Illinois, Hedw. spores Bryol. 7. bryologicis Prinkipo. longicollis Botany Spore (Huelva, W. K. G.A.M. C. C., A.J.E. exine C. C. V. V. R.C. R.C. J.P. P. D.A. Scanning R.C. J. B.D. M.E. E. W. J species & & H. (1981). J V. & in (1971). (1963). & .S., J. Sergio, & wall M.C. (1801). in Lichenol. W.P. .A. Am. (1956). (Pottiaceae, Hirohama, Kurschner, Missouri (1980). USA. P.]. Polytrichum & Cryptogamie P.J. J. Sergio, (1974). Oliva, ornamentation. & & (1941). Mogensen, (1913). (1964). Sevilla selected (1986). Council, (1978). Ann. & Guerra, W. & (1968). Mx. Frey, (1967). Lemmon, Lemmon, formation. tractante. from J. (1975). (1985). The Stone, Lemmon, (1986). Plant Electron The C., (1876). & Species Illustrated Bot. Bibliotheca R. Protoplasma Tortula K.K. An C. Bryophyta Further Larson, Tortula y W. mosses Botanical 11, Ontogeny Cros, The Asir Origin mosses. Cyto-taxonomy Light Cordoba). (1982). J. T. Stockholm. Speciation. (1990). H. electron Taxonomic I. 51, Musci). G.S. A Taxonomy (1983). 57-62. juniperinum B.E. B.E. Ed. & Naturhist. Bryol. Moss (1974). (1976). Muscorum Synopsis Am. synoptic Mountains Microscope. (1984). Notizbl. Hedw. R.M. 195-200. B.E. Ros, and of J. brevissima electron of D.A. (1978). Moss II, Aportaci6n (1985)~ (1980). Bryologist Acaulon Spain: 21. Garden Bryol. Adaptations. J. aus Flora and Moosflora. microscopical Nova electron Lichenol. 106, Stuttgart. R.M. & A (1981). Acta The Bot. Studies (1964). J. sect. Columbia Flora flora revision Bot. Mesopotamien and Brugues, comparative Hofnus. phylogeny muscorum Ultrastructure, Cramer, microscopic Phylogenetic 273-287. Ultrastructure an Hedw. frondosorum of mosses (Saudi 8,269-273. Hedwigia of Bot. (1990). 67, 11, J. fontiquerianum Rurales distribution Schiffn., Britain of annotated Gart. J. microscopic of Aperture 81,494-516. Tortula An Hattori in al 918-934. 11, 25-39. south GUERRA of Eugen Fennoscandia. Malacitana Arabian and 27,472-504. conocimiento Columbia Arabia). M. University of electron-microscopic Aloin Vaduz. study 63-70. Spore Berlin-Dahlem in De and Southern 51, europaeorum (1990). study Georgian P. ein muralis studies Tortula bot. checklist. aspects Ulmer, und descriptae Not. a, ohioense development of 411-433. of Ireland. of histochemistry morphology studies neues ET Aloinella J. Lab. sporogenesis the of the University 7, Kurdistan, sp. Datos Press, (Pottiaceae, Bryol. AL. of the Australia. (Musci: Hedw. 97-118. of subantarctic genus mosses: II. Stuttgart. de nov. exine 38, vorderasiatisches Treballs on Cambridge sporogenesis Ren. spores .. Musci. 15, la sobre New praemissa spore and 475-488. 13, , brioflora de Tortula 3: in 402-405. Leipzig. of structure in Pottiaceae). & Press, sowie Tortula. study and 25-31. Crossidium in Crossidium la York. of Britain. Academic the el Card. spores de Musci) germination Fasc. a Peninsula taxa genero species notes l'Institut moss, European University Hedw., Syrien, New of in de introductione Ph.D. 2. and Br. in spore Archidium. of in Trans. Andaluda on Swedish Wiistensteppenmoos Ditrichum the Acaulon York. of Syst. der Press, Antarct. stratification. the (Musci). asirense with Rhodos, germination Botanic Iberica. Tortula. Thesis, and Pottiaceae wall ostlichen species Press, moss Br. Bot. special London. bud Natural structure en Surv. (Pottiaceae) bryol. Monogr. Bryologist78, pallidum. noroccidental de de University 11, Cryptogamie J. Mytilini la Trematodon London. apical of Barcelona Holarktis. Bryol. Peninsula reference 189-208. by elementis Phascum Grana stages Bull. Science Soc. use in meri- Syst. und III. the 67, 14, im of of of 5, 5, a Published by Maney Publishing (c) British Bryological Society J. Wijk, 2235); et MUB Stone, ale Albacete, Almodovar, Munoz MUB Balsa Nabao, 3543); Stone, s.n.); Segura, Alange, de 3778). Cartagena, Steere, Cadiz, (Bryoth. (Bryoth. (Herb. (LISU Oliva Tortula GUERRA, ale Tortula Tortula Erro, (LISU and R. I.G. I.G. (Herb. W.C. del 660). Beira (Herb. 3779); Asturias, Murcia, key Munoz van s.n.); EI 340); muralis Melo Sergio Fuertes Romero MUB (1988). (1989). MUB Pino, R. s.n.); baetica to Sierra muralis Bosque, der, (1940). Biologia, Ros M. Australian Alta, Oliva Fuertes Leon, Minho, (LISU Oliva Murcia, Margadan, 1552); Acaulon Ros Revision Ros 3156); et Bullas, Estremadura, 901); Hedw. (Herb. Pravia, Tortula de (Bryoth. (Casas ale et Hedw. and Sortelha, (Herb. ApPENDIX Universidad Priaranza (Broth. Las Alcaraz, 1268); s.n.); ale s.n.); Sevilla, species. (LISU Granada, Freixieiro Murcia, granulosum, J. of Cartagena, in Salto var. Fuertes W.D. S. Agones, Lomas, & Phascum (Bryoth. North TORTULA var. CARRI6N, Oliva MUB Ribatejo, Cadiz, Murcia, Oliva) MUB J. muralis. & s.n.). carretera Me Lucero, Rio Setubal, del de Bryol. (SELECTED American Cartagena, Florschiitz, obcordata. s.n.); Sierra 10 Murcia, and a do 1279); Oliva Munoz Bierzo, 3775); 501); de new Guerra Jerez MURALIS Departamento MUB SPAIN: (LISU Cementerio 15,257-268. Cartagena, Acaulon Soutelo, PORTUGAL: San las Salamanca, Ros species Mor6n-Pruna, Serra Elvira, North Campus Murcia, (Herb. Cordoba, P.A. de Espineras, (Herb. Murcia, Antonio, Munoz 937); SPECIMENS & Fuente Almeria, (Bryoth. PORTUGAL: 148903); V in la in of da Ros. (1969). AR. Australia. Melo Frontera, the de Mexico. Varo de Arrabida, Oliva Cantabria, Isla calle San BAETICA Munoz Espinardo, Ribatejo, Acaulon Biologia (Herb. Aldeadavila de SPAIN: Jumilla, Index Entre Casas MUB (LISU Grossa, Guerra (Herb. Algarve, Sierra la STUDIED) Ant6n, Alfonso Oliva Bryologist J. s.n.); Beira Muela, Muscorum. Bryol. Oliva muticum 0339); Vegetal Almeria, Munoz); Montilla et Sergio Murcia, 704); Torres de (Bryoth. s.n.); Cerro Potes, ale Fuertes & Ros Cordoba, Litoral, Ros 15, de Luole, X, 43, (Herb. Ros Maria, Ros (LISU (Botanica), Badajoz, complex: Murcia, & 745-777. & la Spain. Vol. 12-23. Guerra Baixo Navarra, Mojacar, Novas, y (Bryoth. de Aboal Sim-Sim Munoz Ribera, (Bryoth. (Bryoth. s.n.); MUB Montemayor, 5. Anciao, Sergio Oliva s.n.). la Ochotorena Castillo Utrecht. a Alentejo, Facultad comparison Cehegin, Sierra Sal, (Bryoth. (Bryoth. Casas Murcia, 3777). Munoz (Herb. Puerto (LISU Lopez SPAIN: 1275); MUB MUB MUB et Ros Rio 283 ale de de de et