Tortula

Tortula muralis, photographed by Eric Hunt.

Belongs within: Pottiaceae.

Tortula is a cosmopolitan genus of mosses mostly found growing on soil. Species of Tortula have lax stem leaves, giving the plant a delicate appearance, with low or almost absent papillae and a costa with a rounded or semicircular abaxial stereid band section (Zander & Eckel). The genus is highly variable and has been divided in the past between multiple genera such as Tortula sensu stricto with peristome teeth long and twisted, Pottia with the peristome teeth absent or short, and Phascum with the capsule cleistocarpous (Dixon 1924) but the existence of species with intermediate morphologies has lead to these genera being synonymised.

Characters (from Zander & Eckel): Plants forming cushions, green, occasionally blackish green above, yellow-brown to dark brown below. Stems usually to 2 cm, occasionally branching; rounded-pentagonal in transverse section, hyalodermis generally absent; sclerodermis absent, central strand present; rhizoids often dense; axillary hairs ca. 5-8 cells in length, basal 1-3 cells thicker-walled. Leaves appressed-incurved to lax when dry, weakly to widely spreading when moist; obovate to spathulate, ovate to elliptical, occasionally ligulate, adaxial surface nearly flat to concave, broadly channeled, occasionally grooved along costa, 1-4(-6) mm; base scarcely differentiated to elliptical; distal margins recurved below or rarely plane, entire or occasionally serrulate near apex, margins occasionally with 1-4 cell rows often less papillose and smaller, walls thicker; apex broadly acute to rounded, lamina inserted laterally or to 45° on costa; costa short- to long-excurrent as an awn, occasionally percurrent or subpercurrent, adaxial outgrowths rarely present as an elliptical pad of cells bulging from adaxial costal surface, adaxial cells quadrate, papillose or smooth, in 3-4(-5) rows adaxially, abaxial cells short-rectangular to elliptic, papillose or smooth; transverse section of costa circular to semicircular, adaxial epidermis present, adaxial stereid band absent (occasionally small), guide cells 2(-3) (rarely absent), in 1(-2) layers, hydroid strand present, often large, abaxial stereid band present, rounded, elliptical, or semicircular in sectional shape, abaxial epidermis present, occasionally present only laterally, rare absent; basal cells reaching across leaf or rising higher medially, rectangular, 18-25 µm, 2-5:1, walls thin, hyaline; distal laminal cells rounded-quadrate to hexagonal, occasionally rhomboidal, distal laminal cells ca. 15-19 µm wide, 1:1-2, papillae mostly hollow, simple or 2-fid, 4-6 per lumen, occasionally on conic salients, cell walls thin, seldom evenly thickened, superficially convex. Specialized asexual reproduction by brood bodies on rhizoids (sometimes by gemmae on leaves). Sexual condition dioicous or monoicous (commonly autoicous or synoicous); perichaetia terminal, inner leaves little different or somewhat larger than the cauline. Seta yellowish brown or brown, very short to 2.5 cm, twisted counterclockwise or not twisted. Capsule usually stegocarpic, occasionally cleistocarpic, yellowish brown or dark brown, spheric, ovate, elliptic or cylindric, occasionally inclined, 0.5-3(-7) mm, exothecial cells rectangular, 25-30 µm, ca. 2-3:1, walls thin or evenly thickened; annulus of 1-2 rows of vesiculose cells, persistent (very rarely revoluble), cells twisted counterclockwise; peristome of 32 filaments or 16 long or shortly triangular teeth, cleft to near base, or absent, straight or twisted counterclockwise, articulations usually many, teeth absent, rudimentary, or up to 2000 µm, spiculose, basal membrane absent, low, or up to 1000 µm, spiculose; operculum long-conic, occasionally shortly rostrate, occasionally not differentiated, 0.5-2.5 mm. Calyptra cucullate, 2.5-6 mm, smooth. Spores 13-30(-50) µm, papillose, rarely densely spiculose, light brown. Laminal color reaction to KOH yellow, occasionally red medially or rarely throughout, or negative or orange.

<==Tortula Hedw. 1801 (see below for synonymy) SK02
    |--T. abruptinervis A49
    |--T. acaulon SK02 [=Phascum acaulon D24]
    |    |--T. a. var. acaulon SK02
    |    `--T. a. var. schreberiana [=Phascum cuspidatum var. schreberanum (Dicks.) Brid. 1806] SK02
    |--T. anderssonii Ångström 1872 (see below for synonymy) SK02
    |--T. angustata [=Barbula subulata var. angustata] D24
    |--T. antarctica (Hampe) Wilson 1859 (see below for synonymy) SK02
    |--‘Pottia’ antarctica non Tortula antarctica (Hampe) Wilson 1859 D03
    |--‘Pottia’ asperula D24
    |--T. atrovirens (Smith) Lindb. 1864 SK02 (see below for synonymy)
    |    |--T. a. var. atrovirens D24
    |    `--T. a. var. edentula D24
    |--T. austroruralis (Müll.Hal.) Broth. 1902 [=Barbula austroruralis] SK02
    |--‘Phascum’ beaugleholei Stone 1989 SK02
    |--T. brevisetacea (Hampe ex Muell.) Thér. 1919 (see below for synonymy) SK02
    |--T. brevissima M08
    |--‘Pottia’ bryoides [=Phascum bryoides] D24
    |    |--P. b. var. bryoides D24
    |    |--P. b. var. pilifera D24
    |    `--P. b. var. thornhillii D24
    |--‘Pottia’ caespitosa [=Weisia caespitosa] D24
    |--T. calcicolens M08
    |--‘Pottia’ commutata D24
    |--T. crawfordii (Paris) Watts 1902 (see below for synonymy) SK02
    |--T. crinita M09 [=Pottia crinita D24]
    |--‘Phascum’ curvicolle D24
    |--‘Phascum’ cuspidatum Schreb. ex Hedw. 1801 non Tortula cuspidata Hooker & Wilson 1859 SK02
    |    |--P. c. var. cuspidatum D24
    |    |--P. c. var. curvisetum D24
    |    `--P. c. var. piliferum non Pottia bryoides var. pilifera D24
    |--‘Phascum’ floerkeanum D24
    |    |--P. f. var. floerkeanum D24
    |    `--P. f. var. badium D24
    |--‘Pottia’ globosa Catches. 1980 SK02
    |--T. inermis D24
    |--T. intermedia [=Barbula intermedia, Syntrichia intermedia; incl. T. montana] D24
    |--‘Pottia’ intermedia SK02 (see below for synonymy)
    |    |--P. i. var. intermedia D24
    |    `--P. i. var. littoralis D24
    |--T. laevipila (Brid.) Schwägr. 1823 (see below for synonymy) SK02
    |    |--T. l. var. laevipila D24
    |    `--T. l. var. laevipilaeformis D24
    |--‘Pottia’ lanceolata [=Leersia lanceolata] D24
    |--‘Phascum’ laticostum Stone 1989 SK02
    |--‘Pottia’ latifolia [=Weisia latifolia] D24
    |    |--P. l. var. latifolia D24
    |    `--P. l. var. pilifera (see below for synonymy) D24
    |--‘Pottia’ latzii Catches. 1980 SK02
    |--T. leucostega (Müll.Hal.) Broth. 1902 [=Syntrichia leucostega (Müll.Hal.) Zander 1993] SK02
    |--‘Phascum’ longipilum Stone 1989 SK02
    |--T. marginata SG-C01 [=Barbula marginata D24; incl. Desmatodon meridionalis Luisier 1915 SG-C01]
    |--‘Pottia’ megapoda D03
    |--‘Pottia’ minutula [=Gymnostomum minutulum; incl. P. starkei var. davallii] D24
    |--T. muralis Hedw. 1801 (see below for synonymy) SK02
    |    |--T. m. var. muralis D24
    |    |--T. m. var. aestiva D24
    |    `--T. m. var. rupestris D24
    |--T. murina (Müll.Hal.) Broth. 1902 (n. d.) [=Barbula murina] SK02
    |--T. mutica D24 (see below for synonymy)
    |    |--T. m. var. mutica SK02
    |    `--‘Pottia’ m. var. brachyphylla Warnst. 1916 SK02
    |--‘Phascum’ muticum non Pottia mutica Venturi 1869 SK02 [=Acaulon muticum D24, Sphaerangium muticum D24]
    |    |--P. m. var. muticum D24
    |    `--‘Acaulon’ m. var. minus D24
    |--T. norvegica [incl. Barbula aciphylla] D24
    |--T. obtusifolia M09
    |--T. pagorum (Milde) De Not. 1869 [=Syntrichia pagorum (Milde) Amann 1918; incl. T. baileyi, Barbula baileyi] SK02
    |--T. papillosa Wilson 1845 (see below for synonymy) SK02
    |--T. paramattana Mitt. 1882 (n. d.) [=Barbula paramattana] SK02
    |--T. phaea (Hooker & Wilson) Dixon 1923 (see below for synonymy) SK02
    |--T. princeps De Not. 1838 (see below for synonymy) SK02
    |--T. pygmaea Dusén 1907 [=Syntrichia pygmaea (Dusén) Zander 1993] SK02
    |--‘Phascum’ readeranum Stone 1989 SK02
    |--‘Pottia’ recta [=Phascum rectum] D24
    |--T. recurvata Hooker 1819 (see below for synonymy) SK02
    |--‘Pottia sullivanii var.’ recurvimucronata Warnst. 1916 SK02
    |--T. revolvens M08
    |    |--T. r. var. revolvens M08
    |    `--T. r. var. obtusata M08
    |--T. robusta Hooker & Greville 1824 SK02 (see below for synonymy)
    |--‘Acaulon’ robustum Broth. ex Roth 1913 [=Phascum robustum (Broth. ex Roth) Stone 1989] SK02
    |    |--A. r. var. robustum SK02
    |    `--‘Phascum’ r. var. crassinervium (Müll.Hal.) Stone 1989 [=Acaulon crassinervium Müll.Hal. 1902] SK02
    |--T. rubella Hooker & Wilson 1859 SK02 (see below for synonymy)
    |    |--T. r. var. rubella D24
    |    |--‘Barbula’ r. var. dentata D24
    |    `--‘Barbula’ r. var. ruberrima [incl. B. ferruginascens] D24
    |--T. rubra Mitt. 1867 [=Barbula rubra, Desmatodon ruber, Syntrichia rubra] SK02
    |--‘Pottia’ rufescens M09
    |--T. ruraliformis [=Barbula ruraliformis; incl. T. ruralis var. arenicola] D24
    |--T. ruralis (Hedw.) Gaertn., Mey. & Scherb. 1802 SK02 (see below for synonymy)
    |--‘Pottia’ scabrifolia Bartram 1951 SK02
    |--T. serrata Dixon 1923 [=Syntrichia serrata (Dixon) Zander 1993] SK02
    |--‘Pottia’ serrata non Tortula serrata A49
    |--T. serrulata Hooker & Greville 1824 [=Barbula serrulata, Hennediella serrulata] SK02
    |--‘Pottia’ spegazzinii D03
    |--‘Pottia’ starkeana [=Weisia starkeana] D24
    |--P. stevensii A49
    |--T. subulata [=Bryum subulatum, Barbula subulata] D24
    |    |--T. s. var. subulata D24
    |    `--T. s. var. subinermis D24
    |--‘Pottia’ tasmanica Broth. 1895 SK02
    |--‘Phascum tasmanicum’ Dixon & Rodway 1923 (see below for synonymy) SK02
    |    |--P. t. var. tasmanicum SK02
    |    `--P. t. var. unguiculatum Stone 1989 SK02
    |--‘Phascum’ tetragonum [=Tetrapterum tetragonum] SK02
    |--T. truncata (Hedw.) Mitt. 1870 (see below for synonymy) SK02
    |--‘Bryum’ truncatulum [=Pottia truncatula] D24
    |--T. virescens M08
    |--‘Pottia’ viridifolia D24
    |--T. willisiana Zander 1993 SK02
    |    |--T. w. var. willisiana (see below for synonymy) SK02
    |    `--T. w. var. obscura (Willis) Zander 1993 (see below for synonymy) SK02
    `--‘Pottia’ wilsoni [=Gymnostomum wilsoni] D24

Nomina nuda: Phascum loreum Wilson ex Jaeger 1869 SK02
             Pottia stackhousii Watts & Whitel. 1902 SK02
             Tortula crassinervis var. longiseta Müll.Hal. ex Watts & Whitel. 1902 SK02
             Tortula mollissima Broth. ex Watts & Whitel. 1902 SK02
             Tortula atrovirens f. nervoexcedente Watts & Whitel. 1902 SK02
             Tortula perbrevifolia Müll.Hal. ex Watts & Whitel. 1902 SK02
             Tortula scabrifolium Mitt. ex Watts & Whitel. 1902 non Pottia scabrifolia Bartram 1951 SK02

‘Phascum tasmanicum’ Dixon & Rodway 1923 non Pottia tasmanica Broth. 1895 [=Microbryum tasmanicum (Dixon & Rodway) Zander 1993] SK02

‘Pottia’ intermedia non Tortula intermedia SK02 [=Gymnostomum intermedium D24; incl. P. lanceolata var. gymnostoma D24]

‘Pottia’ latifolia var. pilifera nec Pottia bryoides var. pilifera nec Phascum cuspidatum var. piliferum [=Bryum piliferum] D24

Tortula Hedw. 1801 [incl. Phascum Linnaeus ex Hedw. 1801, Pottia (Ehrh. ex Rchb.) Fürnr. 1829, Syntrichia Brid. 1801] SK02

Tortula anderssonii Ångström 1872 [=Syntrichia anderssonii (Ångström) Zander 1993; incl. T. bealeyensis, T. brunnea Broth. & Watts 1912 (preoc.), T. cuspidata Hooker & Wilson 1859 (preoc.)] SK02

Tortula antarctica (Hampe) Wilson 1859 [=Syntrichia antarctica (Hampe) Zander 1993; incl. Barbula androgyna, T. androgyna (Müll.Hal.) Broth. 1902, T. brachytheca (nom. inv.), Barbula brachytricha, T. brachythricha (Müll.Hal.) Broth. 1902, B. flexomarginata, T. flexomarginata (Müll.Hal. & Hampe) Mitt. 1882, B. latrobeana, T. latrobeana (Müll.Hal.) Mitt. 1882, B. panduraefolia, T. panduraefolia (Müll.Hal. & Hampe) Broth. 1902, B. preissiana, T. preissiana (Müll.Hal.) Broth. 1902, B. propinqua, T. propinqua (Müll.Hal.) Broth. 1902] SK02

Tortula atrovirens (Smith) Lindb. 1864 SK02 [=Grimmia atrovirens D24, Barbula atrovirens SK02; incl. Dendropogon adustus (nom. inv.) SK02, Pottia starkeana var. affinis D24, Anacalypta brachyodus SK02, P. starkeana var. brachyodus D24, SK02, P. sullivanii var. brachyodus (nom. illeg.) SK02, Trichostomum convolutum SK02, Desmatodon convolutus (Brid.) Grout 1939 SK02, Tortula crassinervioides Broth. ex Ramsay 1980 (n. n.) SK02, Barbula crassinervis (preoc.) SK02, T. crassinervis (preoc.) SK02, Desmatodon nervosus SK02, Pottia sullivanii (nom. illeg.) SK02]

Tortula brevisetacea (Hampe ex Muell.) Thér. 1919 [=Barbula brevisetacea, B. breviseta (preoc.), Syntrichia brevisetacea (Hampe ex Muell.) Zander 1993, Tortula breviseta (preoc.); incl. T. brevipila (n. n.), Barbula brevipila (nom. inv.)] SK02

Tortula crawfordii (Paris) Watts 1902 [=Barbula crawfordii; incl. B. asperifolia (nom. illeg.), Tortula asperifolia (nom. illeg.)] SK02

Tortula laevipila (Brid.) Schwägr. 1823 [=Syntrichia laevipila Brid. 1819, Barbula laevipila; incl. Tortula rotundata Geh. 1902 (n. n.)] SK02

Tortula muralis Hedw. 1801 [=Barbula muralis; incl. B. austromuralis, Tortula austromuralis, B. elaphrotricha, T. elaphrotricha, B. melbourneana, T. melbourneana, B. nanosubulata Müll.Hal. 1901 (n. n.), T. nanosubulata Müll.Hal. ex Watts & Whitel. 1902 (n. n.)] SK02

Tortula mutica D24 [=Pottia mutica Venturi 1869 SK02; incl. Microbryum starckeanum var. brachyodes SK02, Barbula latifolia D24]

Tortula papillosa Wilson 1845 [=Barbula papillosa, Syntrichia papillosa (Wilson) Jur. 1882; incl. Tortula acuminata Broth. ex Burges 1932 (n. n.), T. calodictyon Broth. ex Burges 1932, Barbula involucrata Müll.Hal. 1897, B. vesiculosa var. involucrata, Tortula vesiculosa var. involucrata (Müll.Hal.) Paris 1906, T. schizopogoniacea Watts & Whitel. 1902 (n. n.), Barbula streptopogoniacea, T. streptopogoniacea (Müll.Hal.) Broth. 1902, Barbula vesiculosa, T. vesiculosa (Müll.Hal.) Broth. 1902] SK02

Tortula phaea (Hooker & Wilson) Dixon 1923 [=Syntrichia phaea (Hooker & Wilson) Zander 1993; incl. T. subbrunnea Broth. & Watts 1912, T. submutica] SK02

Tortula princeps De Not. 1838 [=Syntrichia princeps (De Not.) Mitt. 1859; incl. T. muelleri, Barbula muelleri (nom. illeg.)] SK02

Tortula recurvata Hooker 1819 [=Barbula recurvata, Desmatodon recurvatus (Hooker) Mitt. 1880; incl. Trichostomum reflexidens, D. reflexidens] SK02

Tortula robusta Hooker & Greville 1824 SK02 [=Barbula recurvifolia var. robusta D24, SK02, Syntrichia robusta (Hooker & Greville) Zander 1993 SK02]

Tortula rubella Hooker & Wilson 1859 SK02 [=Barbula rubella Lindb. 1879 SK02, D24, Bryum rubellum D24, Syntrichia rubella (Hooker & Wilson) Zander 1993 SK02; incl. Mollia laxula D24, B. readeri SK02, Tortula readeri (Müll.Hal.) Broth. 1902 SK02]

Tortula ruralis (Hedw.) Gaertn., Mey. & Scherb. 1802 SK02 [=Bryum rurale D24, Barbula ruralis SK02, Syntrichia ruralis (Hedw.) Weber & Mohr 1803 SK02]

Tortula truncata (Hedw.) Mitt. 1870 [=Pottia truncata (Hedw.) Bruch & Schimp. 1843; incl. P. melbourniana, P. readeri, P. subphyscomitrioides] SK02

Tortula willisiana var. obscura (Willis) Zander 1993 [=Pottia drummondii var. obscura Willis 1954; incl. Phascum molle] SK02

Tortula willisiana Zander 1993 var. willisiana [incl. Phascum disrumpens, Pottia disrumpens, Schizophascum disrumpens (nom. inv.), Phascum drummondii, Pottia drummondii (Wilson) Willis 1954] SK02

*Type species of generic name indicated

REFERENCES

[A49] Allison, K. W. 1949. Description of a new species with notes and localities for some Otago and Southland mosses. Transactions and Proceedings of the Royal Society of New Zealand 77 (2): 278–283.

[D24] Dixon, H. N. 1924. The Student's Handbook of British Mosses 3rd ed. V. V. Sumfield: Eastbourne.

[D03] Dusén, P. 1903. Patagonian and Fuegian mosses. In: Scott, W. B. (ed.) Reports of the Princeton University Expeditions to Patagonia, 1896–1899 vol. 8. Botany pp. 63–126. The University: Princeton (New Jersey).

[M08] Marstaller, R. 2008. Moosgesellschaften am Südrand des Kyffhäusergebirges bei Bad Frankenhausen (Kyffhäuserkreis). 130. Beitrag zur Moosvegetation Thüringens. Mauritiana 20 (2): 289–348.

[M09] Marstaller, R. 2009. Die Moosgesellschaften des geplanten Naturschutzgebietes "Brandenburg und Göringer Stein" bei Göringen (Wartburgkreis, Eisenach). 136. Beitrag zur Moosvegetation Thüringens. Mauritiana 20 (3): 529–557.

[SG-C01] Sérgio, C., & I. Granzow-de la Cerda. 2001. On the identity of Desmatodon meridionalis Luisier. Journal of Bryology 23: 69–70.

[SK02] Streimann, H., & N. Klazenga. 2002. Catalogue of Australian Mosses. Flora of Australia Supplementary Series 17. Australian Biological Resources Study: Canberra.

Last updated: 15 December 2017.