62. TORTULA Plates 84, 85, 86, 87, 88, – 89.

Tortula Hedw., Sp. Musc. 122, 1801, nom. cons. non Roxburgh, 1800. Lectotype: Tortula subulata Hedw.

Beccaria C. Müll., Nuov. Giorn. Bot. Ital. 4: 11, 1872.

Bauriella Warnst., Hedwigia 57: 88, 1915, nom. inval. prov. Type: Tortula polyseta (C. Müll.) Warnst.

Tortula sect.? Piliferae De Not., Mem. R. Acc. Sc. Torino 40: 287, 1838, rank not indicated; inoperative in piority I.C.B.N. Art. 35.2.

Barbula sect. Amphidiopsis C. Müll., Linnaea 42: 332, 1879. Type: Barbula amphidiifolia C. Müll.

Barbula sect. Pilifera Lzaro é Ibiza, Bot. Descr. Comp. Fl. Esp. 1: 586, 1896.

Barbula sect. Orthopodiae Kindb., Eur. N. Amer. Bryin. 2: 245, 1897.

Barbula sect. Catillaria C. Müll., Gen. Musc. Fr. 425, 1900. Type: Barbula pellata Schimp.

Pottia sect. Beccaria (C. Müll.) C. Müll., Gen. Musc. Fr. 389, 1900.

Pottia subsect. Acutae C. Jens., Skand. Bladmfl. 203, 1939, nom. inval. descr. suec.

See sectional synonymy for additional nomenclature.

Plants forming cushions or turfs, green or occasionally blackish green above, yellow-brown to dark brown below. Stems branching occasionally, to 2 cm in length, transverse section rounded-pentagonal, central strand present or very rarely absent, sclerodermis absent, hyalodermis absent; axillary hairs ca. 5–8 cells in length, basal 1–3 cells thicker walled; rhizoids often dense. Leaves appressed-incurved to lax when dry, weakly to widely spreading when moist, usually obovate to spathulate, occasionally ovate to elliptical or ligulate, 1–4(–6) µm in length, upper lamina nearly flat to concave, broadly channeled, occasionally grooved along costa, margins recurved below or rarely plane, entire or occasionally weakly serrulate near apex, marginal 1–4 rows of cells often less papillose and smaller than the medial or walls thicker, occasionally marginal cells elongate, rarely bistratose; apex broadly acute to rounded; base scarcely differentiated in shape to elliptical, rarely weakly auricled; costa short- to long-excurrent as an awn, occasionally percurrent or subpercurrent, costa with lamina inserted laterally or to 45°, superficial cells quadrate or occasionally short-rectangular and papillose or smooth ventrally, dorsally short-rectangular to elongate and papillose or smooth, 3–4(–5) rows of cells across costa ventrally at midleaf, costal transverse section circular to semicircular, ventral stereid band absent or occasionally small and represented by a few cells, dorsally present and round, elliptical or semicircular in shape, epidermis present ventrally and dorsally or occasionally only laterally on the dorsal side, rarely absent dorsally, guide cells 2(–3) in 1(–2) layers or rarely absent, hydroid strand usually present, often large, very rarely absent; rarely an elliptical pad of cells bulging from ventral surface of the costa; upper laminal cells rounded-quadrate to hexagonal, occasionally rhomboidal, ca. 15–19 µm in width, 1–2:1, walls thin or seldom evenly thickened, superficially convex; papillae usually hollow, simple or bifid, 4–6 per lumen, occasionally on a conical salient, rarely absent; basal cells differentiated across leaf or higher medially, rectangular, often rather lax, 18–25 µm in width, 2–5:1, walls thin, hyaline, rarely little differentiated. Propagula absent. Dioicous or monoicous (commonly autoicous or paroicous). Perichaetia terminal, inner leaves little differentiated or somewhat larger than the cauline. Perigonia terminal or as autoicous buds in subperichaetial or lower leaf axils. Seta very short or to 2.5 cm in length, 1 (very rarely 2) per perichaetium, yellowish brown to brown, twisted counterclockwise, clockwise or not twisted; theca stegocarpous or else cleistocarpous, 0.5–3.0(–7.0) µm in length, yellowish brown to dark brown, spherical, ovate, elliptical or cylindrical, occasionally inclined, occasionally macrostomous, exothecial cells rectangular, 25–30 µm, ca. 2–3:1, rarely 4–5:1, walls thin or evenly thickened, stomates present at base of theca, phaneropore, annulus of 1–2 rows of vesiculose cells, persistent or very rarely revoluble, occasionally absent or rarely with up to 8 circumferential weak lines of dehiscence; peristome teeth of 32 filaments or 16 triangular teeth or rudimentary or absent, long or shortly triangular, cleft to near base, spiculose, up to 2000 µm in length, with many articulations, straight to twisted counterclockwise, basal membrane absent or low or up to 1000 µm in height, tessellated and spiculose. Operculum when differentiated long-conic, occasionally shortly rostrate, 0.5–2.5 µm in length, cells twisted counterclockwise. Calyptra cucullate, smooth, 2.5–6.0 µm in length. Spores 13–30(–50) µm in diameter, light brown, papillose, rarely densely spiculose. Laminal KOH color reaction usually yellow, occasionally red medially, occasionally negative, rarely reddish orange. Reported chromosome numbers: Sect. Tortula: n = 13+m, 14, 24, 26, 27, 28, 30, 39, 40, 48, 48+m, 50, 52, 60, 66. Sect. Pottia: n = 12, 13, 15, 20, 21, 24, 25, 26, 26+m, 27, 28+1–2acc, 30, 32, 42, 52. The most often reported number for both sections is n = 26.

Found on most continents in various habitats, mainly soil.

With the segregation of various genera (Zander 1989: Chenia, Dolotortula, Hennediella, Hilpertia, Sagenotortula, Stonea, Syntrichia), Tortula becomes a fairly homogeneous group with very similar gametophytes and a characteristic tendency to reduction in the sporophyte. Thus, the correlations of season of sporophyte maturation (Zander 1979d) with genus (Pottia having spring and winter sporophyte maturation dates and Tortula in the traditional sense having dates mainly in the spring and summer) is probably a reflection of “life strategy” (cf. During 1979) rather than phylogeny, though this needs to be tested. Major characters of Tortula as presented here include presence of stem central strand (rarely absent or present in different stems of same collection, e.g. T. brevissima) and absence of sclerodermis and hyalodermis (Pl. 84, f. 2, 17); leaves usually obovate to spathulate, margins usually narrowly recurved below and entire; costal stereid band usually semicircular to rounded in section, hydroid strand present, dorsal epidermis usually present (Pl. 84, f. 7, 8, 21); upper laminal cells usually rather large and clear (i.e. walls usually relatively unobscured by the papillae, Pl. 84, f. 9); propagula absent; upper laminal KOH reaction usually yellow.

Tortula is distinguishable from Syntrichia by the semicircular to rounded stereid band (not crescent-shaped) and yellow KOH reaction of the upper laminal cells (not red). Unlike Syntrichia, a dorsal costal epidermis is usually differentiated, either completely or occasionally only laterally (Pl. 89, f. 15—as is also the case in Hennediella). The yellow KOH reaction is usually present, but some few taxa have no color reaction (e.g. T. entosthodontacea), or the basal cells may be brick red (e.g. T. raucopapillosa), or the leaves may blush red medially in the upper part of the leaf (e.g. T. atrovirens), or the older leaves may be red and the younger yellow (e.g. T. lingulata), or all leaves may have a reddish orange cast (e.g. T. nevadensis). But the characters of these taxa otherwise are those of Tortula as here emended. Hennediella is distinguished by its red KOH reaction, commonly dentate or serrate and plane upper laminal margins, and superficially flattened upper laminal cells. Papillae may be absent in some Tortula species or variously expressed in different specimens of the same species.

Chamberlain (1978) recognized Pottia caespitosa, but the leaves have plane margins, the upper laminal cells are rather small and thick-walled, and costal sections show two stereid bands. This small-statured species is actually a Trichostomum (as witness the combination Trichostomum caespitosum (Bruch ex Brid.) Jur.), differing somewhat from other species of that genus by the broadly sheathing perichaetial leaves and quite neckless capsule.

Upon examination of the sporophytes of a considerable range of species of Pottieae, there was found to be no sharp difference between traditional “Tortula” peristomes with 32 similar rami (Pl. 84, f. 12–13) and “Desmatodon” peristomes with 16 teeth cleft to near the base or to a basal membrane (i.e., 32 paired rami, Pl. 86, f. 16; 87, f. 9–10). Even if some difference was statistically demonstrable, it would cut across observed (and as taxonomically recognized here) clearly defined generic groupings based on several gametophytic characters. It is simpler to entertain convergence of one character (short, probably reduced peristomes having teeth paired) than convergent evolution of several distinctive gametophyte morphotypes two or more times. The flattened basal portion of “Desmatodon” peristome teeth may be explained as a cleft portion of the basal membrane. The proximally flat teeth of Desmatodon species are also different from the almost terete filaments of Tortula with long, twisted peristomes simply because the long filaments of Tortula have distal regions little wider than their thickness. A cladistic evaluation at the species level may clarify this.

Visotska's (1967) proposal of a subfamily Tortuloideae (no type cited), based on a chromosome number of x = 12 and intended to contain Tortula, Aloina and Crossidium, was criticized on cytological grounds by Nyholm and Wigh (1973) because several species of Tortula have a basic chromosome number of x = 13. Evaluation of chromosome counts given by Fritsch (1982, 1991) gave both 12 and 13 as basic numbers for both Tortula s. str. and Syntrichia as conceived in the present study; however, Newton (1972) found n = 7 for S. robusta and Ramsay (1974) found n = 6+m for S. papillosa.

An electron microscopical study by Lewinsky (1974) of spore ornamentation in 10 European species of Tortula s. lat. showed differences between the spores of the specimens studied (only one or two collections were examined for each species although 20–30 spores from two to five capsules were studied in each species), which probably represent differences between the species, but she found no evidence of differences between traditional sections of the genus.

Mishler's (1986a) cladogram of postulated phylogenetic relationships of several species of Tortula s. lat. recognized Tortula s. str. (as recognized here) as a primitive group (he listed T. subulata, T. mucronifolia and T. muralis) distinct from several other species (all recognized here as Syntrichia) by the upper laminal cells not strongly mammilose.

Corley et al. (1981) gave an up-to-date presentation of the sections of the genus as represented in Europe. Their apprehension of Desmatodon as a rather small assemblage of the type species and closely related taxa presages the present study. They stated that Desmatodon “is not defined by sound technical characters, and there has been much confusion about which species should be assigned to it. As with Didymodon there has been too much emphasis placed on the peristome, which is not a conservative character in Pottiaceae.”

Tortula muralis, T. leucostoma and T. altipes occasionally have a small ventral (sub)stereid band (Pl. 85, f. 3). The costa is, however, rounded in section and generally unlike that of Barbula.

Selected literature on Tortula s. str. (here including Phascum, Desmatodon and Pottia): Arts (1987a,b, 1988), Bachelot (1813), Bennett (1965), Brown (1894b,c), Bryan (1956), Carrión et al. (1990), Chamerlain (1978), Crundwell (1953, 1955, 1956), Dixon (1927b), Favali and Gianni (1973), Guerra et al. (1988, 1991, 1992), Häusler (1984), Hernnstadt and Heyn (1989), Holzinger (1925), Hughes (1969, 1979, 1982), Hughes and Wiggin (1969), Jiménez et al. (1990), Kanda (1981), Lazarenko (1969, 1974), Lazarenko and Lesnyak (1972), Lazarenko et al. (1961), Lewinsky (1974), Lightowlers (1984, 1985a,c, 1986a,b,c), Lobachevskaya et al. (1986), Matteri (1977a,b), Mishler (1985b, 1986a, 1990), Mishler and Newton (1988), Ripetskij (1978, 1979), Ripetskij et al. (1983), Risse (1985), Rumsey (1992), Rungby (1957), Sainsbury (1936), Saito (1973a), Savicz-Ljubitzkaya and Smirnova (1963b, 1965), Sérgio (1972a, 1978), Springer (1935), Steere (1939a, 1940), Stone (1989), Toth (1987), Ulycna (1977), Wareham (1939a), Wareham and Whitney (1939), Warnstorf (1912, 1916), Zander (1989).

Number of accepted species: 163, none remaining in Desmatodon, plus 10 as yet undistributed in Phascum, plus 30 undistributed in Pottia.

Species examined: see below.

New heterotypic synonymy: Didymodon schimperi (Mont.) Broth. = Tortula atrovirens (Sm.) Lindb.

New taxa, combinations, statuses and names:

Tortula altipes (Broth.) Zand., comb. nov. (Desmatodon altipes Broth., Act. Hort. Bot. Ac. Sc. U.R.S.S. 42: 154, 1931).

Tortula argentinica (Broth.) Zand., comb. nov. (Desmatodon argentinicus Broth., Ark. Bot. 15(6): 5, 1918), not seen.

Tortula atherodes Zand., nom. nov. (Phascum cuspidatum Schreb. ex Hedw., Spec. Musc. 22, 1801).

Tortula atherodes var. arcuata (Herrnstadt & Heyn) Zand., comb. nov. (Phascum cuspidatum var. arcuatum Herrnstadt & Heyn, Bryologist 94: 175, 1991), not seen.

Tortula atherodes var. affinis (Nees & Hornsch.) Zand., comb. nov. (Phascum affine Nees & Hornsch., Bryol. Germ. 1: 74, 1823; Phascum cuspidatum var. affine (Nees & Hornsch.) Hampe), not seen.

Tortula atherodes var. curviseta (Dicks.) Zand., comb. nov. (Phascum curvisetum Dicks., Pl. Crypt. Brit. 4: 2, 1801; Phascum cuspidatum var. curvisetum (Dicks.) Nees & Hornsch.), not seen.

Tortula atherodes var. diaphora (Hag.) Zand., comb. nov. (Phascum acaulon var. diaphorum Hag., K. Norsk. Vid. Selsk. Skrift. 1928(3): 19, 1929; Phascum cuspidatum var. diaphorum (Hag.) C. Jens.), not seen.

Tortula atherodes var. elata (Brid.) Zand., comb. nov. (Phascum elatum Brid., J. Bot. (Schrader) 1800(1): 269, 1801; Phascum cuspidatum var. elatum (Brid.) Drumm.), not seen.

Tortula atherodes var. intertexta (Brid.) Zand., comb. nov. (Phascum intertextum Brid., Mant. Musc. 8, 1819; Phascum cuspidatum var. intertextum (Brid.) Brid., not seen.)

Tortula atherodes var. marginata (Hernnstadt & Heyn) Zand., comb. nov. (Phascum cuspidatum var. marginatum Hernnstadt & Heyn, Bryologist 94: 175, 1991), not seen.

Tortula atherodes var. mitraeformis (Limpr.) Zand., comb. nov. (Phascum cuspidatum var. mitraeforme Limpr., Laubm. Deutschl. 1: 187, 1885), not seen.

Tortula atherodes var. papillosa (Lindb.) Zand., comb. nov. (Phascum papillosum Lindb., Oefv. K. Vet. Ak. Foerh. 21: 217, 1864; Phascum cuspidatum var. papillosum (Lindb.) Roth); Phascum cuspidatum ssp. papillosum Guerra & Ros in Guerra, Jiménez, Ros & Carrión), not seen.

Tortula atherodes var. pilifera (Hedw.) Zand., comb. nov. (Phascum piliferum Scherb. ex Hedw., Spec. Musc. 20, 1801; Phascum cuspidatum var. piliferum Scherb. ex Hedw.) Hook. & Tayl.).

Tortula atherodes var. retortifolia (Guerra & Ros in Guerra, Jiménez, Ros & Carrión) Zand., comb. nov. (Phascum cuspidatum var. retortifolium Guerra & Ros in Guerra, Jiménez, Ros & Carrión, Cryptogamie, Bryol. Lichénol. 12: 390, 1991), not seen.

Tortula atherodes var. schreberiana (Dicks.) Zand., comb. nov. (Phascum schreberianum Dicks., Pl. Crypt. Brit. 4: 2, 1801; Phascum cuspidatum var. schreberianum (Dicks.) Brid.), not seen.

Tortula atrovirens var. leucodonta (Corb.) Zand., comb. nov. (Barbula atrovirens var. leucodonta Corb., Mém. Soc. Sc. Nat. Cherbourg 26: 244, 1889; Desmatodon convolutus var. leucodontus (Corb.) Wijk & Marg.), not seen.

Tortula bogosica (C. Müll.) Zand., comb. nov. (Desmatodon bogosicus C. Müll., Nuov. Giorn. Bot. Ital. 4: 12, 1872).

Tortula capillaris (Chen) Zand., comb. nov. (Desmatodon capillaris Chen, Hedwigia 80: 287, 1941), not seen.

Tortula cernua var. xanthopus (Kindb.) Zand., comb. nov. (Desmatodon cernuus var. xanthopus Kindb., Ottawa Natural. 4: 61, 1890), not seen.

Tortula cuneifolia var. blissii Zand., var. nov. (Pl. 85, f. 21–25.) A varietate typica gametophytis subatris, in solutione KOH colorem profundiorem evolventibus, foliis rigide appressis, seta curta, crassa, longitudine 4–5 µm, latitudine 0.2–0.3 µm differt.

Differs from the typical variety by the blackish gametophytes, these more strongly colored in KOH; leaves stiffy appressed; seta short, ca. 4–5 in length, stout, 0.2–0.3 µm in width. Type: Canada, Northwest Territories, Cornwallis Island, Resolute Bay area, L. C. Bliss, 1977, holotype, BUF; isotype, ALTA. The leaves of var. blissii are identical in morphology to those of muticous-leaved forms of the European species T. cuneifolia, but show a very strong color reaction to KOH: bright yellow upper lamina and deep brick-red basal cells, colors which are pale in European specimens at BUF. The short, thick seta is apparently unique to this arctic variety; European specimens have setae 0.7–1.5 µm in length and ca. 0.15 µm in width. Like European specimens, the capsule is variably macro- and microstomous, and the operculum is broadly to narrowly conic. This specimen was incorrectly reported (Vitt & Zander 1978) as a second known station for Crumia deciduidentata (here treated as a species of Tortula near T. cuneifolia), which has a similar short, thick seta and smooth, weakly bordered leaves that are bright yellow in KOH except for brick-red basal cells, but which differs in its capsule about twice as long (2.7–3.3 µm vs. 1.3–1.6 µm) and proportionately thicker; operculum pushed off by the elongating columella; spores larger (ca. 18 µm vs. ca. 13 µm); leaves ovate to clearly spathulate (vs. short- to long-ovate); constricted leaf apex; and much enlarged basal cells. A section across the leaf base of var. blissii shows what appear to be two stereid cell groups of equal size separated by a group of hydroid cells (also seen in sections of the basal portion of the costa of Tortula muralis, and see discussion of Phascopsis).

Tortula entosthodontacea (Card. & Dix.) Zand., comb. nov. (Hyophilopsis entosthodontacea Card. & Dix., J. Bot. 49: 137, 1911.).

Tortula euryphylla Zand., nom. nov. (Dicranum latifolium Hedw., Spec. Musc. 140, 1801; Desmatodon latifolius (Hedw.) Brid.).

Tortula euryphylla ssp. brevifolia (Kindb.) Zand., comb. nov. (Tortula latifolia (Hedw.) Lindb. ssp. brevifolia Kindb., Bih. K. Svensk. Vet. Ak. Handl. 7(9): 135, 1883; Desmatodon latifolius ssp. brevifolius (Kindb.) Kindb.).

Tortula euryphylla var. eucalyptrata (Lindb.) Zand., comb. nov. (Tortula eucalyptrata Lindb., Bot. Not. 1886: 100, 1886; Desmatodon latifolius varr eucalyptratus (Lindb.) Kaur.).

Tortula euryphylla var. flavescens (Brid.) Zand., comb. nov. (Dicranum latifolium var. flavescens Brid., Spec. Musc. 140, 1801; Desmatodon latifolius var. flavescens).

Tortula euryphylla var. spelaea (Amann) Zand., comb. nov. (Desamtodon spelaeus Amann, Bull. Murithienne 40: 46, 1919; Desmatodon latifolius var. spelaeus (Amann) Podp.).

Tortula euryphylla var. subobliqua (Lindb.) Zand., comb. nov. (Desmatodon latifolius var. suboliquus Lindb., Oefv. K. Vet. Ak. Foerh. 23: d553, 1867).

Tortula chungtienia Zand., nom. nov. (Desmatodon yuennanensis Broth., Symb. Sin. 4: 44, 1929).

Tortula deciduidentata (Sharp & Iwats.) Zand., comb. nov. (Crumia deciduidentata Sharp & Iwats., J. Hattori Bot. Lab. 32: 95, 1969).

Tortula kabir-khanii (Broth.) Zand., comb. nov. (Desmatodon kabir-khanii Broth., Mitteil. Inst. Allg. Bot. Hamburg 8: 400, 1931), not seen.

Tortula lanceola Zand., nom. nov. (Encalypta lanceolata Hedw., Spec. Musc. 63, 1801; Pottia lanceolata (Hedw.) C. Müll.; non Tortula lanceolata (Hedw.) P. Beauv.).

Tortula lanceola var. albidens (Corb.) Zand., comb. nov. (Pottia lanceolata var. albidens Corb., Rev. Bryol. 22: 35, 1895), not seen.

Tortula lanceola var. angustata (B.&S. in BSG) Zand., comb. nov. (Anacalypta lanceolata var. angustata B.&S. in BSG, Bryol. Eur. 2: 48, 1843; Pottia lanceolata var. angustata (B.&S. in BSG) C. Müll.), not seen.

Tortula lanceola var. lejolisii (Corb.) Zand., comb. nov. (Pottia lanceolata var. lejolisii Corb., Mem. Soc. Sci. Nat. Cherbourg 26: 238, 1889), not seen.

Tortula lanceola var. leucodonta (Schimp.) Zand., comb. nov. (Pottia lanceolata var. leucodonta Schimp., Syn. ed. 2: 158, 1876), not seen.

Tortula lanceola var. macrophylla (Warnst.) Zand., comb. nov. (Pottia lanceolata var. macrophylla Warnst., Hedwigia 58: 133, 1916), not seen.

Tortula lanceola var. microphylla (Warnst.) Zand., comb. nov. (Pottia lanceolata var. microphylla Warnst., Hedwigia 58: 134, 1916), not seen.

Tortula lanceola var. mucronata (Amann) Zand., comb. nov. (Pottia lanceolata var. mucronata Amann, Bull. Soc. Vaudoise Sci. Nat. 53: 85, 1920), not seen.

Tortula lanceola var. ovalifolia (Warnst.) Zand., comb. nov. (Pottia lanceolata var. ovalifolia Warnst., Hedwigia 58: 131, 1916), not seen.

Tortula lanceola var. papillosa (Corb.) Zand., comb. nov. (Pottia lanceolata var. papillosa Corb., Mém. Soc. Sci. Nat. Cherbourg 26: 237, 1889), not seen.

Tortula lanceola var. rigidior (Schwaegr.) Zand., comb. nov. (Encalypta lanceolata var. rigidior Schwaegr., Spec. Musc. Suppl. 1(1): 61, 1811.)

Tortula laureri var. setschwanica (Broth.) Zand., comb. nov. (Desmatodon setschwanicus Broth., Symb. Sin. 4: 43, 1929; Desmatodon laureri var. setschwanicus (Broth.) Chen), not seen.

Tortula maritima (R. Br. ter) Zand., comb. nov. (Dendia maritima R. Br. ter, Trans. New Zealand Inst. 30: 411, 1898; Pottia maritima (R. Br. ter) Broth.).

Tortula minor (C. Müll.) Zand., comb. nov. (Beccaria minor C. Müll., Nuov. Giorn. Bot. Ital. 4: 11, 1872; Pottia minor (C. Müll.) Wijk & Marg.), not seen.

Tortula minor var. elatior (C. Müll.) Zand., comb. nov. (Beccaria elatior C. Müll., Nuov. Giorn. Bot. Ital. 4: 11, 1872; Pottia minor var. elatior (C. Müll.) Wijk & Marg.).

Tortula modica Zand., nom. nov. (Gymnostomum intermedium Turn., Musc. Hib. 7, 1804; Pottia intermedia (Turn.) Fuernr.).

Tortula modica var. corsa (Fleisch. & Warnst.) Zand., comb. nov. (Pottia intermedia var. corsa Fleisch. & Warnst., Bot. Centralbl. 65: 299, 1896), not seen.

Tortula modica var. gymnandra (Schiffn.) Zand., comb. nov. (Pottia intermedia var. gymnandra Schiffn., Oesterr. Bot. Zeitschr. 47: 55, 1897), not seen.

Tortula modica var. gymnogyna (Schiffn.) Zand., comb. nov. (Pottia intermedia var. gymnogyna Schiffn., Oesterr. Bot. Zeitschr. 48: 389, 1898), not seen.

Tortula modica var. revoluta (Schiffn.) Zand., comb. nov. (Pottia intermedia var. revoluta Schiffn., Oesterr. Bot. Zeitschr. 47: 55, 1897), not seen.

Tortula modica var. stenocarpa (Velen.) Zand., comb. nov. (Pottia intermedia var. stenocarpa Velen., Rozpravy Cesk. Ak. Ved. Tr. 2, 6(6): 148, 1897), not seen.

Tortula modica var. tenuis (Vent.) Zand., comb. nov. (Pottia intermedia var. tenuis Vent., Muscin. Trent. 31, 1899), not seen.

Tortula nevadensis (Card. & Thér.) Zand., comb. nov. (Pottia nevadensis Card. & Thér., Bot. Gaz. 37: 365, 1904).

Tortula pallida (Lindb.) Zand., comb. nov. (Pottia pallida Lindb., Oefv. K. Vet. Ak. Foerh. 21: 220, 1864).

Tortula pallida var. longicuspis (Warnst.) Zand., comb. nov. (Pottia pallida var. longicuspis Warnst., Hedwigia 58: 113, 1916).

Tortula porteri (Jam. in Aust.) Zand., comb. nov. (Desmatodon porteri Jam. in Aust., Musci Appal. 123, 1870).

Tortula protobryoides Zand., nom. nov. (Phascum bryoides Dicks., Pl. Crypt. Brit. 4: 3, 1801; Pottia bryoides (Dicks.) Mitt.).

Tortula protobryoides var. brevifolia (De Not.) Zand., comb. nov. (Phascum bryoides var. brevifolium De Not., Atti Univ. Genova 1: 734, 1869; Pottia bryoides var. brevifolia (De Not.) Wijk & Marg.), not seen.

Tortula protobryoides var. thornhillii (Wils.) Zand., comb. nov. (Phascum bryoides var. thornhillii Wils., Bryol. Brit. 33, 1855; Pottia bryoides var. thornhillii (Wils.) Braithw.), not seen.

Tortula randii (Kenn.) Zand., comb. nov. (Pottia randii Kenn., Rhodora 1: 78, 1899; Desmatodon randii (Kenn.) Laz.).

Tortula raucopapillosa (X.-j. Li) Zand., comb. nov. (Desmatodon raucopapillosus X.-j. Li, Acta Bot. Yunnan. 3: 105, 1981 “raucopapillosum”).

Tortula rhodonia Zand., nom. nov. (Desmatodon wilczekii Meyl., Bull. Soc. Vaudoise Sc. Nat. 52: 383, 1919), not seen.

Tortula sainsburyana Zand., nom. nov. (Pottia stevensii R. Br. ter, Trans. N. Z. Inst. 26: 291, 1894).

Tortula solomensis (Broth.) Zand., comb. nov. (Desmatodon solomensis Broth., Rev. Bryol. n. ser. 2: 2, 1929), not seen.

Tortula splachnoides (Hornsch.) Zand., comb. nov. (Phascum splachnoides Hornsch., Horae Phys. Berol. 57, 1820; Pottia splachnoides (Hornsch.) Broth.).

Tortula thompsonii (C. Müll.) Zand., comb. nov. (Trichostomum thompsonii C. Müll., Bot. Zeit. 22: 359, 1854; Desmatodon thompsonii (C. Müll.) Jaeg.).

Tortula tonkinensis (Besch.) Zand., comb. nov. (Desmatodon tonkinensis Besch., Bull. Soc. Bot. France 41: 80, 1894), not seen, cf. Brotherus (1924: 297).

Tortula truncata var. brevirostris (Lisa) Zand., comb. nov. (Gymnostomum truncatum var. brevirostre Lisa, Elenco Muschi Torino 16, 1837; Pottia truncata var. brevirostris (Lisa) De Not.), not seen.

Tortula truncata var. illyrica (Latz.) Zand., comb. nov. (Pottia illyrica Latz., Beih. Bot. Centralbl. 48(2): 481, 1931; Pottia truncata var. illyrica (Latz.) Podp.), not seen.

Tortula truncata var. littoralis (Mitt.) Zand., comb. nov. (Pottia littoralis Mitt., J. Bot. 9: 4, 1871; Pottia truncata var. littoralis (Mitt.) Warnst.), not seen.

Tortula truncata var. minutissima (Warnst.) Zand., comb. nov. (Pottia truncata var. minutissima Warnst., Hedwigia 58: 117, 1916), not seen.

Tortula ucrainica (Laz.) Zand., comb. nov. (Desmatodon ucrainicus Laz., Bull. Jard. Bot. Kieff 4: 34, 1926).

Tortula willisiana Zand., nom. nov. (Phascum drummondii Wils., London J. Bot. 7: 26, 1848; Pottia drummondii (Wils.) Willis).

Tortula willisiana var. obscura (Willis) Zand., comb. nov. (Pottia drummondii var. obscura Willis, Vict. Nat. 70: 171, 1954), not seen.

Tortula wilsonii (Hook.) Zand., comb. nov. (Gymnostomum wilsonii Hook., Bot. Misc. 1: 143, 41, 1829; Pottia wilsonii (Hook.) BSG).

Tortula wilsonii var. asperula (Mitt.) Zand., comb. nov. (Pottia asperula Mitt., J. Bot. 9: 4, 1871; Pottia wilsonii ssp. asperula (Mitt.) Kindb.), not seen.

Tortula wilsonii var. crinita (Wils. ex B.&S.) Zand., comb. nov. (Pottia crinita Wils. ex B.&S. in BSG, Bryol. Eur. 2: 43, 1843; Pottia wilsonii var. crinita (Wils. ex B.&S.) Warnst.), not seen.

Tortula wilsonii var. mucronifolia (Bruch in F. A. Müll.) Zand., comb. nov. (Entosthymenium mucronifolium Bruch in F. A. Müll., Flora 12: 387, 1829; Pottia wilsonii var. mucronifolia (Bruch in F. A. Müll.) Warnst.), not seen.

Tortula zoddae Zand., nom. nov. (Pottia cuneifolia Solms ex Schimp., Syn. ed 2, 154, 1876).

TORTULA Sect. TORTULA

Tortula sect. Tortula Hedw., Sp. Musc. 122, 1801, nom. cons. non Roxburgh, 1800. Lectotype: Tortula subulata Hedw.

Desmatodon Brid., Mant. Musc. 86, 1819. Lectotype: Desmatodon latifolius (Hedw.) Brid. fide Venturi, Comm. Fauna Fl. Gea Venezia 1: 123, 1868.

Zygotrichia Brid., Bryol. Univ. 1: 520, 1826. Type: Zygotrichia leucostoma (R. Br.) Brid.

Dermatodon Hüb., Musc. Germ. 14: 109, 1833, p.p.

Pachyneurum Amann, F. Mouss. Suisse 2: 112, 1912.

Tortula subg. Desmatodon (Brid.) Lindb., Musci Scand. 20, 1879.

Tortula subg. Zygotrichia (Brid.) Lindb., Musci Scand. 20, 1879.

Tortula subg. Tortula C. Jens., Medd. Groenland 3: 379, 1887, nom. illeg.

Didymodon subg. Desmatodon (Brid.) Kindb., Eur. N. Amer. Bryin. 2: 273, 1897.

Barbula subg. Zygotrichia Kindb., Eur. N. Amer. Bryin. 2: 245, 1897.

Barbula subg. Tortula Kindb., Eur. N. Amer. Bryin. 2: 246, 1897, hom. illeg.

Tortula subg. Pachyneurum (Amann) C. Jens., Skand. Bladmfl. 200, 1939.

Pottia subg. Pseudodesmatodon Medel. in C. Jens., Skand. Bladmfl. 207, 1939, nom. inval. descr. suec.

Tortula subg. Eutortula C. Jens., Skand. Bladmfl. 199, 1939, nom. illeg.

Barbula sect. Tortulae Rebent., Prodr. Fl. Neomarch. 257, 1804.

Tortula sect. Subulatae De Not., Mem. R. Acc. Sc. Torino 40: 287, 1838. Type: Tortula subulata Hedw.

Barbula sect. Subulatae B.&S. in BSG, Bryol. Eur. 2: 98, 1842 (fasc. 13–15 Mon. 36).

Trichostomum sect. Desmatodon (Brid.) C. Müll., Syn. 1: 588, 1849.

Barbula sect. Crassinerves (De Not.) Milde, Bryol. Soles. 112, 1869. Type: Barbula nervosa (BSG) Milde.

Tortula sect. Desmatodon (Brid.) Mitt., J. Linn. Soc. Bot. 12: 145, 164, 1869.

Tortula sect. Zygotrichia (Brid.) Mitt., J. Linn. Soc. Bot. 12: 145, 168, 1869.

Barbula sect. Crassicostatae Schimp., Syn. ed. 2: 194, 1876.

Desmatodon sect. Eudesmatodon Jur., Laubmfl. Öst. Ungarn 129, 1882, nom. illeg.

Desmatodon sect. Crassicostati Jur., Laubmfl. Öst.-Ungarn 135, 1882. Type: Desmatodon atrovirens (Sm.) Jur.

Desmatodon sect. Subulati Jur., Laubmfl. Öst. Ungarn 129, 1882. Lectoype nov.: Desmatodon subulatus (Hedw.) Jur.

Pottia sect. Didyctium C. Müll. in Broth., Hedwigia 34: 124, 1895. Type: Pottia asperula C. Müll. in Broth.

Barbula sect. Crassinervia Lzaro é Ibiza, Bot. Descr. Comp. Fl. Esp. 1: 506, 1896, p.p. Crossidium & p.p. Aloina.

Barbula sect. Muraliformes Kindb., Eur. N. Amer. Bryin. 2: 246, 1897. Type: Barbula muralis (Hedw.) Crome.

Barbula sect. Canescentes Kindb., Eur. N. Amer. Bryin. 2: 245, 1897. Type: Barbula canescens (Mont.) B.&S.

Barbula sect. Limbatae Kindb., Eur. N. Amer. Bryin. 2: 246, 1897. Type: Barbula marginata B.&S.

Barbula sect. Subulataeformes Kindb., Eur. N. Amer. Bryin. 2: 245, 1897. Type: Barbula subulata (Hedw.) P. Beauv.

Barbula sect. Tortula Hérib., Mém. Ac. Sc. Clermont-Ferrand ser. 2, 14: 364, 1899, nom. illeg.

Tortula sect. Tortula Broth., Nat. Pfl. 1(3): 429, 1902, nom. illeg.

Tortula sect. Eutortula Broth., Act. Hort. Gothoburg 1: 191, 1924, nom. illeg.

Syntrichia sect. Zygotrichia (Brid.) Mönk., Laubm. Eur. 306, 1927.

Tortula sect. Crassicostatae (Schimp.) Podp., Consp. Musc. Eur. 242, 1954.

Tortula sect. Crassinerves (Milde) Wijk & Marg., Taxon 8: 75, 1959. Type: Barbula nervosa (De Not.) Milde, hom. illeg.

Tortula subsect. Desmatodon (Brid.) Braithw., Brit. Moss. Fl. 3: 212, 1885.

Upper laminal cells usually densely papillose and relatively small (ca. 10–13 µm in width), ventral costal cells 3–6 in transverse section, usually forming a convex pad and usually densely hollow papillose with simple or bifid papillae, leaves usually with rounded apices and commonly with long hyaline awns; costal sections with guide cells; peristomes usually present.

Lindberg (1864) treated T. subulata and T. mucronifolia as a single taxon. Steere (1940) pointed out that, although the variable (Warnstorf 1912) T. subulata and T. mucronifolia intergrade extensively in Europe, the two species appear to have have distinct ranges and morphology in North America. Chen (1941) commented, significantly as it turns out, that T. mucronifolia (as Syntrichia mucronifolia) has a leaf morphology quite like that of many Pottia species. This is similarly unfortunate for nomenclatural purposes, since the closely related T. subulata is the type of the genus and therefore of the sect. Tortula. In the present study, the variation between T. subulata and T. mucronifolia was found to parallel major differences in laminal structure between sect. Tortula and sect. Pottia. Thus, in spite of an apparent close relationship, these two species are placed in different sections of the genus. This is not unacceptable if the two are considered taxa whose ancestors are inserted near the base of the lineage of Tortula species; this needs to be analysed at the species level.

Tortula sect. Tortula includes species with papillose leaves and small cells (e.g. T. muralis) that are nearly identical with species with larger, smooth laminal cells (e.g. T. californica and T. transcaspica, Pl. 86, f. 23–25), but these last are distinctly awned and placed in sect. Tortula on that account. Perhaps T. subulata, with its short awn, ought to be placed with the species of section Pottia (a name change at the sectional level would be necessary), but T. subulata otherwise has the bulk of characters given above for the section. However, many species of sect. Tortula with the laminal morphology of T. muralis, the species probably most representative of sect. Tortula, also have short awns or lack them altogether. The evidence that sect. Pottia demonstrates far more sporophyte reduction than sect. Tortula may be interpreted either as an additional character supporting sect. Pottia as a distinctive line of evolution (just as Syntrichia characteristically lacks sporophyte reduction) or that smooth cells and reduced sporophytes are correlated at the end of several separate reduction series from sect. Tortula. In any case, this should be further analysed at the species level.

Steere (1939a) felt that Tortula leucostoma (Pl. 87, f. 5–10) and T. guepinii would better be regarded as subspecies of T. euryphylla (all as Desmatodon spp., the last as D. latifolius).

Tortula revolvens (Pl. 89, f. 1–4) has the innermost cells of its revolute upper margins hollow-papillose, the group somewhat differentiated as a photosynthetic organ like that of Hilpertia and species of Pseudocrossidium; T. revolvens is, however, closely related to T. muralis by the costa round in section, with a single strong stereid band, and upper lamina highly papillose, cells superficially flat, and KOH reaction yellow.

Tortula atrovirens (Pl. 85, f. 5–8) is near T. revolvens in short-ovate leaf shape, thickened costa, and upper marginal cells epapillose. Tortula muralis has a similarly thickened costa, likewise often with two apparent stereid bands. Tortula atrovirens is similar to Crossidium by its habit of low-growing, gemmiform plants with spiralling ovate to obovate leaves, the development of specialized ventral costal photosynthetic tissue (the cells of which have lumens vertically elongate in section, as was nicely illustrated by Flowers 1973a), the rounded dorsal stereid band, the medial placement of the simple, hollow papillae on the leaf, upper lamina yellow but with a reddish blush medially on the leaf in KOH, and a general similarity to Crossidium aberrans and C. seriatum of the areolation (upper cells ca. 15–18 µm in width), the thickened dorsal superficial upper median cell walls, and other features. Delgadillo (1975a, p. 282) proposed that Crossidium was derived, along with Aloina, from a Tortula or Desmatodon-like ancestor through elaboration of ventral costal outgrowths and reduction of leaves and sporophytes. The cladograms do not support this otherwise cogent inference, either placing Crossidium in a different tribe (the Hyophileae, Cladograms 14–16) or having the immediate ancestors of Crossidium inserted deeper in the cladogram than those of Tortula (see Cladograms 9, 10 and 13). The costal sections of C. aberrans and T. atrovirens are extraordinarily similar. Tortula atrovirens has sporophyte maturation dates (Zander 1979d) more characteristic of Crossidium than species previously placed in Desmatodon. As in the extension of the Pseudocrossidium evolutionary series (Zander 1979f) to include P. revolutum, which has no specialized ventral photosynthetic tissue, it may be possible to include T. atrovirens in Crossidium to fill it out as a natural genus. On the other hand, Crossidium sect. Pseudocrossidium (no nomenclatural relation to the genus Pseudocrossidium), which includes C. aberrans and other species with similar costal filaments, may be more appropriately included in Tortula, perhaps as a section or recognized as a separate genus in its own right. This needs further evaluation.

Tortula bogosica, because of its rather deep ventral costal groove and lack of dorsal costal epidermal cells, actually may well be Barbula indica or a closely related species lacking the ventral stereid band; this also requires further study. The little-papillose Tortula californica is placed in sect. Tortula largely because of its evident relationship with T. transcaspica, especially through the rounded leaf apex and long, hyaline awn, but has the rather large upper laminal cells of sect. Pottia; the same is true for T. grandiretis, which has somewhat longer leaves than the first two.

Species of sect. Tortula examined: T. altipes (H), T. atrovirens, T. bogosica (NY), T. brevipes (BUF), T. brevissima (US), T. californica (BUF), T. canescens (BUF), T. euryphylla, T. grandiretis (NY), T. guepinii (CANM), T. leucostoma (NY), T. lingulata (NY), T. peruviana (NY), T. marginata (BUF), T. muralis, T. obtusifolius, T. platyphylla (NY), T. plinthobia, T. raucopapillosa (NY), T. revolvens (NY), T. sublimbata (NY), T. subulata (BUF), T. thianschanica (H), T. trachyphylla (H), T. transcaspica (H), T. vahliana (NY), T. wilsonii (BUF, NY).

TORTULA Sect. POTTIA

Tortula sect. Pottia (Ehrh. ex Reichenb.) Kindb., Bih. K. Svensk. Vet. Ak. Handl. 7(9): 131, 1883. Lectotype: Pottia truncata (Hedw.) BSG, see Wareham in Grout, Moss Fl. N. Amer. 1: 197, 1939.

Phascum L. ex Hedw., Spec. Musc. 19, 1801. Lectotype: Phascum cuspidatum Hedw., see Wareham in Grout, Moss Fl. N. Amer. 1: 195, 1939.

Anacalypta Röhl. ex Leman, Dict. Sci. Nat. ed. 2, 2 Suppl. 38, 1816. Type: Anacalypta lanceolata (Hedw.) Nees & Hornsch.

Physedium Brid., Bryol. Univ. 1: 51, 1826. Type: Physedium splachnoides (Hornsch.) Brid.

Pottia (Ehrh. ex Reichenb.) Ehrh. ex Fürnr., Flora 12(2 Erg.): 10, 1829.

Mildeella Limpr., Laubm. Deutschl. 1: 191, 1885, hom. illeg. non Trevisan, 1876. Type: Mildeella bryoides (Dicks.) Limpr.

Mildea Warnst., Krypt. Fl. Brandenburg 2: 82, 1904, hom. illeg. non Grisebach, 1866.

Phascum subg. Phascum Sull. in A. Gray, Man. Bot. N. U.S. ed. 2: 615, 1856.

Pottia subg. Anacalypta (Röhl. ex Leman) Boul., Fl. Crypt. Est Muscin. 509, 1872.

Tortula subg. Pottia (Ehrh. ex Reichenb.) Lindb., Musci Scand. 21, 1879.

Pottia subg. Eupottia Boul., Muscin. France 471, 1884, nom. illeg.

Phascum subg. Euphascum Limpr., Laubm. Deutschl. 7: 185, 1885, nom. illeg.

Phascum subg. Mildeella Kindb., Eur. N. Amer. Bryin. 2: 403, 1897. Type: Phascum bryoides Dicks.

Pottia subg. Pottia (Ehrh. ex Reichenb.) Broth., Nat. Pfl. 1(3): 423, 1902, hom. illeg.

Pottia subg. Mildeella (Kindb.) Broth., Nat. Pfl. 1(3): 423, 1902.

Gymnostomum sect. Pottia Ehrh. ex Reichenb., Consp. Regn. Veg. 1: 33, 1828.

Barbula sect. Cuneifoliae BSG, Bryol. Eur. 2: 93, 1842 (fasc. 13–14 Mon. 31).

Phascum sect. Annua Ångstr. in Fries, Summ. Veg. Scand. 1: 97, 1846.

Pottia sect. Eupottia C. Müll., Syn. 1: 550, 1849, nom. illeg.

Pottia sect. Anacalypta (Röhl. ex Leman) C. Müll., Syn. 1: 547, 1849, nom. illeg. prior. ut gen.

Tortula sect. Cuneifoliae (B.&S. in BSG) Spruce, Ann. Mag. Nat. Hist. ser. 2, 3: 375, 1849. Type: Tortula cuneifolia (With.) Turn.

Trichostomum sect. Anacalypta (Röhl. ex Leman) C. Müll., Linnaea 42: 312, 316, 1879.

Desmatodon sect. Cuneifolii (BSG) Jur., Laubmfl. Oest. Ungarn 129, 1882. Lectotyp. nov.: Desmatodon cuneifolius (With.) Jur.

Barbula sect. Camptopodiae Kindb., Eur. N. Amer. Bryin. 2: 245, 1897. Type: Barbula laureri (Schultz) Kindb.

Pottia sect. Mildeella (Kindb.) Mönk., Laubm. Eur. 324, 1927.

Phascum sect. Euphascum Podp., Consp. Musc. Eur. 222, 1954, nom. illeg. Basionym: Phascum subg. Euphascum Limpr., nom. illeg.

Pottia sect. Pseudodesmatodon Podp., Consp. Musc. Eur. 232, 1954, nom. inval. Type: Pottia randii Kenn.

Pottia sect. Pottia (Ehrh. ex Reichenb.) Nyholm, Ill. Fl. Nordic Mosses 2: 81, 1989, nom. superfl.

Upper laminal cells usually smooth or weakly papillose and large (ca. 15–20 µm in width), ventral costal cells 2–3 in transverse section, commonly bulging individually or arranged in longitudinal rows as low lamellae and usually smooth or simply once- or twice-papillose, leaves usually with acute apices and short yellow-brown awns (rarely hyaline awned); costal section with guide cells; peristomes often rudimentary or absent.

Magill (1981) pointed out that Acaulon and Phascum are not easily distinguishable in South Africa by the traditional characters of size of capsule beak, curvature of leaf margins and the papillosity of the laminal cells. He defined Phascum by its narrower, plane-margined, erect to spreading leaves, and emergent capsule and cucullate calyptra, distinguishing Acaulon by the bulbiform habit, broad, concave leaves and small, mitrate calyptra. Acaulon is recognized here as a distinctive genus. Chamberlain (1978) suggested that Phascum and Pottia are linked through Pottia recta and Pottia bryoides and that it would be taxonomically more satisfactory if the two genera were united. The two genera are here synonymized with Tortula s. str., but Pottia recta belongs with Microbryum.

Holzinger (1925) thought Pottia randii to be a depauperate form of Desmatodon cernuus; this is also reflected in the combination D. randii (Kenn.) Laz. of Lazarenko (1963b). These two (Pl. 85, f. 15–20; 88, f. 25–27) are kept together here in sect. Pottia. These and other species, like T. laureri (Pl. 87, f. 1–4) and T. thompsonii, with bordered, mostly plane and weakly serrulate margined leaves and large, superficially flat upper laminal cells are quite like species of Hennediella, a genus otherwise only distinguished from Tortula by commonly plane and serrate laminal borders and red laminal KOH reaction. These species of sect. Pottia (often placed in Desmatodon but not including the type of that genus) commonly have summer sporophyte maturation dates (Zander 1979d) rather than the more usual winter and spring dates of traditional Tortula species, and, on revision, may prove worthy of a section of their own.

Tortula lanceola (Pl. 86, f. 17–22) is similar to various Crossidium species in the weakly twisted peristome of 16 split, triangular teeth, the short, bulging cells of the ventral surface of the upper costa, the smooth areolation, and the laminal KOH reaction yellow with a red blush in the upper medial portion of the leaf. It differs mainly in the narrow recurvature of the laminal margins and very weak development of the ventral costal bulge, not quite sufficient for one to consider it a specialized photosynthetic organ. Another species similar to Crossidium, T. atrovirens, is here placed with sect. Tortula.

Certain specimens of small-statured species with basal laminal cells commonly reddish in KOH (e.g. Tortula truncata, Pl. 89, f. 11–15) may react red throughout the lamina giving the impression of the genus Microbryum; these are apparently from unusually harsh environments, and usually may be assigned to Tortula by a lack of laminal papillae.

Species of sect. Pottia examined: T. atherodes, T. cernua (NY), T. chungtienia (H, NY), T. cuneifolia (ALTA, BUF, NY), T. deciduidentata (NY), T. modica (BUF), T. lanceola (BUF), T. laureri (BUF, NY), T. minor (NY), T. mucronifolia (BUF), T. nevadensis (US), T. pallida (NY), T. paulsenii (H), T. planifolia (NY), T. protobryoides (BUF), T. randii (H, NY), T. sainsburyana (NY), T. solmsii (BM), T. systylia (NY), T. thompsonii (NY), T. truncata, T. ucrainica (NY), T. websteri (US), T. zoddae (NY).

TORTULA Sect. SCHIZOPHASCUM

Tortula sect. Schizophascum (C. Müll.) Zand., comb. nov.

Phascum sect. Schizophascum C. Müll., Flora 71: 6, 1888. Type: Phascum disrumpens C. Müll.

Dendia R. Br. ter, Trans. New Zealand Inst. 30: 411, 1898. Type: Dendia maritima R. Br. ter

Pottia subg. Schizophascum (C. Müll.) Broth., Nat. Pfl. 1(3): 423, 1901.

Pottia sect. Schizophascum (C. Müll.) Wareham in Grout, Moss Fl. N. Amer. 1(4): 197, 1939.

Costal section lacking guide cells, and stereid band distinctly central in the section, otherwise gametophyte similar to that of sect. Pottia; sporophyte with elongate seta and elliptical capsule, this being cleistocarpous or dehiscing along up to 8 circumferential lines, usually along weakened cell walls at butt ends of longitudinally elongate rectangular cells (ca. 3–5:1) arranged in even rows (or “pallisades”).

Found on soil, often near the sea; South Africa, Australia and New Zealand.

Brotherus (1902–09) sunk the monotypic Dendia (type species Dendia maritima R. Br. ter = Tortula maritima, Pl. 88, f. 1–9) into Pottia subg. Schizophascum, a move approved by Dixon (1923), who regarded the curious tattered capsule dehiscence of the two known collections as a merely “unhealthy, not to say thoroughly rotten condition” associated with “more or less complete immersion of the capsules from time to time in sea-water.” Tortula willisiana, Pl. 89, f. 16–20, from Australia, however, has similar wall weakenings in its exothecial cells. Characters allowing recognition at least at the section level are the elongate, palisade-like exothecial cells that have weak walls along the butt ends allowing dehiscence along up to 8 circumferencial lines in at least two of the three species, and the lack of guide cells in the costa in all species. The hydroid strand is sometimes absent in some leaves of species in this section. Species of other sections of Tortula, if cleistocarpous, dehisce along irregular lines, these not distinctly arranged circumferencially, and have guide cells in their costae.

The type of Tortula sect. Schizophascum, Pottia disrumpens (C. Müll.) Broth., was referred to T. willisiana (as Pottia drummondii) by Willis (1954), a species very similar or the same as T. maritima. Tortula sect. Schizophascum may be apprehended as an end member of a reduction series through sect. Pottia. This may possibly involve ancestors of T. cuneifolia, which has nearly identical gametophytes but does have guide cells, but not ancestors of the cleistocarpous T. atherodes (Pl. 84, f. 16–22) (which, although it has a minute seta and spherical capsule, has more strongly awned, narrower, little concave leaves with a less lax areolation). Involved would be loss of the costal guide cells and elaboration of a unique dehiscence feature in some species. The spores are very large (ca. 35–40 µm diam.), but this is a common condition of other cleistocarpous (e.g. T. atherodes) and eperistomate species (e.g. T. nevadensis). Note, however, that the various cladograms demonstrate that reduced austral taxa are commonly relicts of lineages inserted deeply on the tribal or subfamilial subclade. Thus, sect. Schizophascum may be better recognized as a separate genus. Again, further analysis at the species level is called for.

Tortula maritima (syntype NY! New Zealand, “Godley Heads”), of which only two collections are known, is not operculate, but neither is it exactly cleistocarpous since capsules are cleanly ruptured along the central one or two of five to eight weak annular lines encircling the capsule from near the base to near the apex. In nearly mature but undehisced capsules, these annular lines are easily seen as minute cleavages between the cells in lateral view around the outline of the somewhat flattened capsule on a microscope slide. No exothecial cells are differentiated, however, as smaller, specialized annular cells as in the case with stegocarpous mosses. The exothecial cells are rectangular and have much the same “pallisade” arrangement as those of Aschisma and Tetrapterum, (see Cladograms 14–16, but compare with other cladograms in the study). In Aschisma rectangular exothecial cells (ca. 3–4:1) are considered unusual because the capsule is spherical, and, in taxa with spherical capsules, the exothecial cells are generally short-rhomboidal or short-rectangular (e.g. as in T. atherodes).

A significant character of all species here referred to sect. Schizophascum is the costal section showing a central region of stereid cells, this often including a ventrally or centrally situated hydroid strand, the stereid band surrounded by a ring of substereid cells, then surrounded again by an epidermis of thin-walled and often bulliform cells; thus, the guide cells are either absent or somehow hidden amoung similar cells of different developmental origin. The costal section is somewhat like that of Stegonia in the bulliform ventral cells and flattened dorsal surface, and similar to that of Saitoella, which lacks guide cells and has bulliform ventral epidermal cells, but also differs in lacking a hydroid strand.

Magill (1981, p. 209) suggested that Tortula maritima of New Zealand, T. splachnoides (Pl. 89, f. 10) of South Africa (as Pottia) and T. willisiana of Australia (as Pottia drummondii) may constitute a single, albeit variable circumantarctic taxon (see also Catcheside 1980). He did not, however, synonymize them, but referred South African specimens previously identified as T. maritima to T. splachnoides (as Pottia). An authentic specimen of T. willisiana (NY!) has the characters of T. maritima: concave leaves, upper laminal cells either epapillose or papillose medially, the costal section with centrally located stereid band, and the sporophytes, though immature, show a pallisade arrangement of cells with weak breaks along circumferencial lines. On the other hand, an isotype (NY! one plant) of Tortula splachnoides has exothecial cells ca. 2:1 in dimension, not in a pallisade arrangement, and breaks in the capsule wall are across cells, not along cell walls; the costal section, however, is like that of the other two species in the stereid (substereid) band positioned centrally and lacking guide cells. Tortula splachnoides is included here with other species (pending further study) because of the like costal section, the fact that immature capsules of T. maritima have rather short exothecial cells, and in view of the short, concave, nearly smooth leaves. The non-type specimens identified as T. splachnoides at BM have plane leaves with rather papillose laminae (except for a smooth border), and a distinct guide cell layer with the stereid band not central, but rather situated dorsally, thus there may be more than one taxon referred to T. splachnoides in herbaria; these non-type and as yet unidentified specimens, although cleistocarpous, lack a pallisade arrangement of exothecial cells but have similarly large spores. The number of accepted species in this section is three.

Additional literature: Brown (1898b).

Species of sect. Schizophascum examined: Tortula maritima (NY), T. splachnoides (NY), T. willisiana (NY).

TORTULA Sect. HYOPHILOPSIS

Tortula sect. Hyophilopsis (Card. & Dix.) Zand., comb. et stat. nov.

Hyophilopsis Card. & Dix., J. Bot. 49: 137, 1911. Type: Hyophilopsis entosthodontacea Card. & Dix.

Near Tortula sect. Pottia but differing by the annulus being revoluble, or persistent but strongly developed and bulging-reflexed when well wetted. Additional characters of value in identification are upper laminal cells 2–3:1, usually weakly papillose, costa percurrent, margins narrowly bordered by elongate cells, autoicous or rhizautoicous sexual condition; perigonia terminal on small, loosely foliate stems at base of perichaetiate plants; seta elongate and capsule cylindrical but peristome teeth rudimentary, consisting of several nubbins barely rising above a low basal membrane which itself scarcely exceeds the annulus; operculum conic, cells very weakly twisted counterclockwise.

A single rare taxon found on walls, mortar, and lateritic rocks in India's Western Ghats.

Hyophilopsis was found to be weakly distinguishable from Tortula sect. Pottia, mainly by the peristome reduction unaccompanied by seta and capsule reduction (Pl. 86, f. 1–10). The gametophyte of this monotypic genus (type: India, Western Ghats, Sedgwick 119, isotypes, NY, PC) is much like that of Tortula laueri.

Literature: Dixon (1911a).

Species examined: T. entosthodontacea (NY, PC, US).