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).