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. 1801SK02 (see below for synonymy) |--T. abruptinervisA49 |--T. anderssonii Ångström 1872 (see below for synonymy)SK02 |--T. antarctica (Hampe) Wilson 1859 (see below for synonymy)SK02 |--T. atrovirens (Smith) Lindb. 1864S04 (see below for synonymy) | |--T. a. var. atrovirensD24 | `--T. a. var. edentulaD24 |--T. austroruralis (Müll.Hal.) Broth. 1902 [=Barbula austroruralis]SK02 |--T. brevisetacea (Hampe ex Muell.) Thér. 1919 (see below for synonymy)SK02 |--T. brevissimaM08 |--T. canescens Mont. 1833S04 [=Barbula canescensD24] |--T. cernua (Huebener) Lindb. 1887S04 [=Desmatodon cernuusS04, Trichostomum cernuumOZB-O03; incl. Anacalypta inclinataOZB-O03] |--T. crawfordii (Paris) Watts 1902 [=Barbula crawfordii]SK02 |--T. cuneifolia (Dicks.) Turner 1804S04 (see below for synonymy) |--T. euryphylla Zander 1993 (see below for synonymy)OZB-O03 |--T. freibergii Dixon & Loeske 1934S04 |--T. inermisD24 |--T. leucostega (Müll.Hal.) Broth. 1902 [=Syntrichia leucostega (Müll.Hal.) Zander 1993]SK02 |--T. leucostoma (R. W. Br.) Hook. & Grev. 1824S04 (see below for synonymy) |--T. marginata (Bruch & Schimp.) Spruce 1845S04 [=Barbula marginataD24; incl. Desmatodon meridionalis Luisier 1915SG-C01] |--T. mucronifolia Schwägr. 1823 [=Barbula mucronifolia, T. subulata ssp. mucronifolia]OZB-O03 |--T. muralis Hedw. 1801S04 (see below for synonymy) | |--T. m. var. muralisS04 | `--T. m. var. aestiva Hedw. 1801S04 [=Barbula muralis var. aestivaOZB-O03] |--T. murina (Müll.Hal.) Broth. 1902 (n. d.) [=Barbula murina]SK02 |--T. obtusifolia (Schwägr.) Mathieu 1853 [=Desmatodon obtusifolius, T. muralis ssp. obtusifolia]OZB-O03 |--T. paramattana Mitt. 1882 (n. d.) [=Barbula paramattana]SK02 |--T. phaea (Hooker & Wilson) Dixon 1923 (see below for synonymy)SK02 |--T. pygmaea Dusén 1907 [=Syntrichia pygmaea (Dusén) Zander 1993]SK02 |--T. randii (Kenn.) Zander 1993 [=Desmatodon randii]OZB-O03 |--T. recurvata Hooker 1819 (see below for synonymy)SK02 |--T. revolvensM08 | |--T. r. var. revolvensM08 | `--T. r. var. obtusataM08 |--T. robusta Hooker & Greville 1824 [=Syntrichia robusta (Hooker & Greville) Zander 1993]SK02 |--‘Acaulon’ robustum Broth. ex Roth 1913 [=Phascum robustum (Broth. ex Roth) Stone 1989]SK02 | |--A. r. var. robustumSK02 | `--‘Phascum’ r. var. crassinervium (Müll.Hal.) Stone 1989 [=Acaulon crassinervium Müll.Hal. 1902]SK02 |--T. rubra Mitt. 1867 [=Barbula rubra, Desmatodon ruber, Syntrichia rubra]SK02 |--T. serrata Dixon 1923 [=Syntrichia serrata (Dixon) Zander 1993]SK02 |--T. serrulata Hooker & Greville 1824 [=Barbula serrulata, Hennediella serrulata]SK02 |--T. solmsii (Schimp.) Limpr. 1888S04 |--P. stevensiiA49 |--T. subulata Hedw. 1801S04 (see below for synonymy) | |--T. s. var. subulataS04 | |--T. s. var. angustata (Schimp.) Limpr. 1888S04 (see below for synonymy) | |--T. s. var. graeffii Warnst. 1904S04 | `--T. s. var. subinermis (Bruch & Schimp.) Wilson 1855S04 |--T. systylia [=Desmatodon systylius]D24 |--T. vahliana (Schultz) Mont. in Gay 1850S04 [=Barbula vahlianaD24] | |--T. v. var. vahlianaD24 | `--T. v. var. subflaccida [incl. T. oblongifolia]D24 `--T. willisiana Zander 1993SK02 |--T. w. var. willisiana (see below for synonymy)SK02 `--T. w. var. obscura (Willis) Zander 1993 [=Pottia drummondii var. obscura Willis 1954; incl. Phascum molle]SK02 Nomina nuda: Desmatodon pachyneuroides Müll.Hal. 1901 [=Trichostomum pachyneuroides Watts & Whitel. 1902]SK02 Desmatodon pachyneurus Müll.Hal. 1901 [=Trichostomum pachyneurus Watts & Whitel. 1902]SK02 Tortula crassinervis var. longiseta Müll.Hal. ex Watts & Whitel. 1902SK02 Tortula mollissima Broth. ex Watts & Whitel. 1902SK02 Tortula atrovirens f. nervoexcedente Watts & Whitel. 1902SK02 Tortula perbrevifolia Müll.Hal. ex Watts & Whitel. 1902SK02 Tortula scabrifolium Mitt. ex Watts & Whitel. 1902 non Pottia scabrifolia Bartram 1951SK02
Tortula Hedw. 1801SK02 [incl. Barbula sect. Cuneifoliae Schimper 1856OZB-O03, Tortula sect. Cuneifoliae (Schimper) Ochyra in Ochyra, Żarnowiec & Bednarek-Ochyra 2003OZB-O03, Desmatodon Brid. 1819SK02]
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. 1864S04 [=Grimmia atrovirensD24, Barbula atrovirensSK02; incl. Dendropogon adustus (nom. inv.)SK02, Pottia starkeana var. affinisD24, Trichostomum convolutumSK02, Desmatodon convolutus (Brid.) Grout 1939SK02, Tortula crassinervioides Broth. ex Ramsay 1980 (n. n.)SK02, Barbula crassinervis (preoc.)SK02, T. crassinervis (preoc.)SK02, Desmatodon nervosusSK02, 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 cuneifolia (Dicks.) Turner 1804S04 [=Bryum cuneifoliumD24, Barbula cuneifoliaD24, Tortula (sect. *Cuneifolia) cuneifoliaOZB-O03]
Tortula euryphylla Zander 1993 [=*Desmatodon latifolius non T. latifolia, Trichostomum latifolium; incl. D. latifolius var. brevicaulis, D. latifolius var. glacialis, D. guentheri]OZB-O03
Tortula leucostoma (R. W. Br.) Hook. & Grev. 1824S04 [=Desmatodon leucostaomaS04; incl. D. obliquusD24, D. suberectusS04, Tortula suberectaS04]
Tortula muralis Hedw. 1801S04 [=Barbula muralisSK02; incl. B. austromuralisSK02, Tortula austromuralisSK02, B. elaphrotrichaSK02, T. elaphrotrichaSK02, T. muralis f. incanaOZB-O03, T. pusilla var. incanaD24, T. muralis ssp. lingulataOZB-O03, Barbula melbourneanaSK02, T. melbourneanaSK02, B. nanosubulata Müll.Hal. 1901 (n. n.)SK02, T. nanosubulata Müll.Hal. ex Watts & Whitel. 1902 (n. n.)SK02, T. muralis var. rupestrisS04]
Tortula phaea (Hooker & Wilson) Dixon 1923 [=Syntrichia phaea (Hooker & Wilson) Zander 1993; incl. T. subbrunnea Broth. & Watts 1912, T. submutica]SK02
Tortula recurvata Hooker 1819 [=Barbula recurvata, Desmatodon recurvatus (Hooker) Mitt. 1880; incl. Trichostomum reflexidens, D. reflexidens]SK02
Tortula subulata Hedw. 1801S04 [=Bryum subulatumD24, Barbula subulataD24, Syntrichia subulataOZB-O03; incl. Tortula subulata f. dentataOZB-O03]
Tortula subulata var. angustata (Schimp.) Limpr. 1888S04 [=Barbula subulata var. angustataD24, Desmatodon subulatus var. angustatusOZB-O03]
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.
[M08] Marstaller, R. 2008. Moosgesellschaften am Südrand des Kyffhäusergebirges bei Bad Frankenhausen (Kyffhäuserkreis). 130. Beitrag zur Moosvegetation Thüringens. Mauritiana 20 (2): 289–348.
[OZB-O03] Ochyra, R., J. Żarnowiec & H. Bednarek-Ochyra. 2003. Census Catalogue of Polish Mosses. Institute of Botany, Polish Academy of Sciences: Cracow.
[SG-C01] Sérgio, C., & I. Granzow-de la Cerda. 2001. On the identity of Desmatodon meridionalis Luisier. Journal of Bryology 23: 69–70.
[S04] Smith, A. J. E. 2004. The Moss Flora of Britain and Ireland 2nd ed. Cambridge University Press.
[SK02] Streimann, H., & N. Klazenga. 2002. Catalogue of Australian Mosses. Flora of Australia Supplementary Series 17. Australian Biological Resources Study: Canberra.