74. SYNTRICHIA Plates
105,
106,
107,
108,
109,
110–
111.
Syntrichia Brid., J. Bot. (Schrader) 1(2): 299, 19 April 1801 (vide Sayre
1959). Type: Syntrichia ruralis (Hedw.) Web. & Mohr, lectotype fide
Zander, Phytologia 65: 432, 1989.
Tortula subg. Syntrichia (Brid.) Chev., Fl. Gén. Env. Paris 2: 52,
1827.
Barbula subg. Syntrichia (Brid.) BSG, Bryol. Eur. 2:10, 1851 (fasc.
46–47 Consp. 2: III).
Tortula sect. Syntrichia (Brid.) Lam. & Cand., Syn. 100, 1806.
Tortula sect. Rurales De Not., Mem. R. Acc. Sc. Torino 40: 286, 1838.
Type: Tortula ruralis (Hedw.) Gaetrn., Meyer & Scherb.
Barbula sect. Rurales BSG, Bryol. Eur. 2: 101, 1842 (fasc. 13–15 Mon.
39).
Barbula sect. Syntrichia (Brid.) C. Müll., Syn. 1: 632, 1849.
Barbula sect. Vallidens C. Müll., Linnaea 42: 347, 1879, nom. inval.
Type: Barbula percarnosa C. Müll.
Barbula sect. Syntrichiae Lesq. & James, Man. N. Amer. Moss. 130,
1884, nom. illeg.
Barbula sect. Ruraliformes Kindb., Eur. N. Amer. Bryin. 2: 245, 1897, nom.
illeg. incl. sect. prior. Type: Barbula ruralis Hedw.
Syntrichia sect. Eusyntrichia Moenk., Laubm. Eur. 306, 1927, nom.
illeg.
Plants
usually coarse, forming often
deep turf, green, occasionally reddish or blackish above, reddish brown below.
Stems branching occasionally, 1–4(–12) cm in length, transverse section
rounded-pentagonal, central strand present or absent, sclerodermis absent or
occasionally of 1–2 layers of substereid cells, hyalodermis absent or
present, sometimes collapsed; axillary hairs ca. 3–10 cells in length, basal
1–3 cells thicker walled or all hyaline; rhizoids common, often dense. Leaves
appressed to weakly spreading when dry, spreading to squarrose when moist,
narrowly ligulate to broadly spathulate, occasionally broadly lanceolate, (1.5–)4–7
µm (inclusive of awn) in length, upper lamina broadly channeled to keeled,
often narrowly grooved along costa, margins plane to recurved, rarely
broadly incurved, usually entire, occasionally serrulate to dentate
above, occasionally bordered with thick-walled, less papillose cells, these
occasionally intramarginal or rarely elongate; apex rounded to broadly acute,
occasionally narrowly acute or cucullate; base not differentiated in shape to
elliptical; costa often stout, commonly prominent dorsally, ending
several cells before apex to excurrent as a long, often hyaline, often serrate,
occasionally flattened awn, costa with lamina inserted at 45°,costal superficial cells rounded-quadrate,
papillose, occasionally elongate and smooth near apex ventrally, dorsally
elongate, papillose to denticulate or spinose, occasionally nearly smooth, 2–4
rows of cells across costa ventrally at midleaf, costal transverse section
round, occasionally elliptical or semicircular, stereid bands ventrally
absent or very rarely of a few substereid or stereid cells, dorsally present
and crescent-shaped, occasionally semicircular, ventral epidermis present, the
dorsal usually absent, guide cells 2–4 per layer in 1–4 layers, hydroid
strand absent or present, occasionally multiple; upper laminal cells
rounded-quadrate, ca. 11–15(–20) µm in width, 1(–2):1,
occasionally bistratose entirely or in patches, walls thin or occasionally
collenchymatous and interiorly porose, superficially strongly convex on both
sides; papillae ca. (1–)4–8 per lumen, usually bifid,
occasionally simple, solid or hollow, rarely simple; basal cells differentiated
across base, rising higher medially, often sharply differentiated from the
upper cells, occasionally inflated medially, 18–30 µm in width, 2–6:1, walls
thin, sometimes irregularly porose, 3–6 rows of basal marginal cells usually
narrowly rectangular and thicker walled. Propagula occasionally present,
variously clusters of small, deciduous leaves; rhizoid-borne tubers; very
fragile cauline leaves or leaf tips (rarely deciduous); obovate or clavate or
raspberry-shaped propagula found on ventral surface of costa or medially or
marginally on upper lamina. Dioicous or monoicous (then usually autoicous or
synoicous). Perichaetia terminal, inner leaves usually little differentiated in
size and shape, rarely enlarged and sheathing, lower cells rhomboidal, rarely
throughout leaf. Perigonia gemmate, terminal or lateral as stalked buds,
paraphyses often clavate. Seta elongate, (0.8–)1.0–3.0 cm in
length, 1(–2) per perichaetium, reddish brown, twisted counterclockwise; theca
2–6 µm in length, reddish brown, cylindrical, commonly slightly curved,
exothecial cells rectangular, 20–30 µm in width, ca. 4–6:1, walls thin to
evenly thickened, capsule neck occasionally distinct, stomates phaneropore, at
base of theca, annulus of 2–4 rows of vesiculose cells, persistent; peristome
teeth 32, filamentous, densely spiculose, ca. 1000–2000 µm in length, with many
articulations, twisted counterclockwise ca. once, basal membrane usually
present, ca. 100–800 µm in height, papillose to spiculose. Operculum
conic, rather large, 1.2–2.2 µm in length, cells twisted counterclockwise. Calyptra
cucullate, smooth, rather large, 3.0–4.5 µm in length. Spores 8–15 µm in
diameter, light brown, lightly papillose. Laminal KOH color reaction brick
red. Reported chromosome number n = 6+m, 12, 12+m, 13, 13+m, 24, 24+2m, 26,
28, 32+2m, 48.
Found
on rock, soil and bark on all continents, most commonly in temperate areas but
also characteristic of dry climates.
Syntrichia is segregated from Tortula s. lat.
by the combination of red KOH reaction of the upper laminal cells, lack of
narrowly elongate upper laminal marginal cells, the crescent-shaped transverse
section of the stereid band, and the general lack of differentiated dorsal
costal epidermal cells. This concept is essentially that of Kramer (1980,
1988), who emphasized the exposed dorsal stereid band (Pl. 105, f. 7, 15), not
covered dorsally by parenchymatous or otherwise differentiated epidermal cells.
Recently Ochyra (1992) supported this concept (Zander 1989) as “a natural group
that deserves recognition as a genus of its own” with a series of new
combinations appropriate for the Polish flora. Syntrichia is
distinguished from Hennediella (likewise KOH red) by the leaves ligulate
to spathulate or very seldom lanceolate, margins usually recurved and seldom dentate
or bordered, upper laminal cells smaller and superficially strongly convex,
costa commonly excurrent as an awn, almost always lacking a dorsal costal
epidermis, stereid band semicircular to crescent-shaped, and sporophytes not in
a reduction series within the genus.
Note
that in some few species (e.g. S. brandisii) a dorsal epidermis is very
weakly differentiated (seen as somewhat wider lumens in costal transverse
section), but these species are placed here rather than with Hennediella
because of their recurved, unbordered margins and small upper laminal cells. Syntrichia
costesii (Pl. 107, f. 17–19), like certain species of Hennediella,
has an intramarginal border of differentiated cells, but those cells are
isodiametric and the costa lacks a differentiated dorsal epidermis. The
existence of a weakly differentiated dorsal epidermis in some species may
indicate a derivation of Syntrichia from Bryoerythrophyllum,
which differs, in traditional characters, in the presence of a second stereid
band and usually narrower (broadly lanceolate to ligulate) leaves. Most
cladograms in the phylogenetic analysis show Syntrichia and Bryoerythrophyllum
to be quite distant. A few species, like S. laevipila (Pl. 109, f. 4–5),
may occasionally and perhaps abnormally have one or two stereid or substereid
cells present ventrally between the guide cells and the ventral epidermis as a
tiny second stereid band, while in the type (isotype, NY) of S. rubra
(Pl. 111, f. 5–8) and a second specimen seen (as Tortula rubra var. subantarctica,
Campbell I., Sorensen 1946, isotype, NY), the ventral stereid band is of
several stereid cells, the ventral epidermis is lacking, and superficially, the
ventral surface of the costa is similar to the dorsal surface, being of
superficially elongate, simply papillose cells. Syntrichia percarnosa
(Pl. 109, f. 20–22) has much the same leaf morphology as Trichostomum
crispulum, and the former species may well belong with that genus; the
substereid cells of the costa are, however, arranged more like those of Syntrichia.
Certain
species of Didymodon sect. Vineales (KOH red) may lack a ventral
stereid band, but have lanceolate leaves and narrowly rectangular basal cells. Bryoerythrophyllum
and Mironia species are also KOH red and may have long-ligulate leaves,
but the transverse section of the costa is reniform in section and the stem has
a sclerodermis. Syntrichia geheebiaeopsis (Pl. 108, f. 13–17, cf. discussion
by Lightowlers 1985a as Tortula) has many of the characters of Bryoerythrophyllum
and Mironia, including broadly lanceolate, dentate leaves with a
sheathing leaf base, small upper laminal cells with low, crowded papillae. It
further differs from Syntrichia by the very narrowly rectangular basal
cells. Bryoerythrophyllum and Mironia have semicircular to
reniform costal sections, however, with usually two distinct stereid bands and
a differentiated dorsal epidermis, but S. geheebiaeopsis has the typical
costal morphology of Syntrichia: a nearly circular costal section, with
a single stereid band and no differentiated dorsal epidermal cells. It mayprove
to be a good genus in the Merceyoideae apparently having lost, like Streptopogon,
the ventral stereid band.
Syntrichia
cavallii (Pl. 107, f. 7–12) is
unusual in its strongly differentiated perichaetial leaves that are similar to
those of S. papillosa (sect. Collotortula); it is, however,
apparently related to S. percarnosa, instead. Syntrichia cavallii
differs from Calyptopogon, in which perichaetial leaves are
differentiated, in that propagula are lacking and the upper laminal papillae
are multiplex. Willia also has strongly differentiated perichaetial
leaves, but these are usually somewhat secund, and the sporophyte is
distinctively reduced in length and complexity. Syntrichia cavallii is
here placed in sect. Syntrichia, not sect. Collotortula, because
of the lack of collenchymatous thickenings in the upper laminal cell walls.
Eventually, perhaps, further study may require that a separate section or genus
be devised for S. cavallii (see also discussion under Willia).
Syntrichia sect. Collotortula Zand., sect. nov. Type: Syntrichia
andicola (Mont.) Zand.
A sectione typica cellulis laminalibus in regione
mediana superna plerumque distincte in angulis incrassatis vel tumescentitrigonis,
marginibus folii plerumque recurvis, papillis interdum simplicibus, propagulis
si effectis clavatis vel ellipticis differt.
Differs
from the typical section by the following combination of characters: medial
upper laminal cells usually somewhat thickened at the corners or even trigonous
(i.e. with bulging knots), leaf margins usually recurved, papillae sometimes
simple, and propagula when present clavate or elliptical. Examination of
transverse leaf sections indicates that the interior upper laminal cell walls
may have large, round central pores (staining helps define this) in at least
some species (Syntrichia aculeata, S. amphidiacea, S. andicola
(Pl. 106, f. 19), S. bogotensis, S. gemmascens, S. gromschii
and S. papillosa), while in other species, including those of other
sections of Syntrichia, the pores are absent or what appear to be
interior pores are occasional, irregular and possibly artifacts. Features
correlated with laminal cell collenchymatous thickenings are the upper leaf
margin usually serrate and the stem central strand often lacking, but these are
not unique characters. The section may not be sharply distinct from sect. Syntrichia
because certain species of that section may have weakly thickened cell corners
(appearing as bright triangular points of light under high magnification). The
counterclockwise-twisted seta of Syntrichia papillosa commented upon by
Dixon (1923) as unusual is not taxonomically significant, since long setae in
the Pottiaceae are generally twisted clockwise below and counterclockwise
above, and short setae are untwisted or merely twisted clockwise. The seta of S.
papillosa (Pl. 109, f. 19—from a specimen misidentified as Tortula
panduraefolia, Tasmania, Weymouth 2821, NY) is only about 5 µm in length.
Species (some surely synonyms of S. andicola) belonging to sect. Collotortula
include: S. aculeata, S. alpestris, S. amphidiacea (Pl.
106, f. 1–8), S. andicola (Pl. 106, f. 14–20), S. antarctica
(possibly belongs here, but laminal cell walls are only weakly collenchymatous),
S. bogotensis, S. cainii (Pl. 107, f. 5–6), S. ciliata, S.
conferta, S. fontana (laminal cells flat superficially, as in Hennediella),
S. gemmascens (Pl. 106, f. 18–21), S. goudotii, S. gromschii,
S. mollis, S. papillosa (Pl. 109, f. 14–19), S. rivularis
(Pl. 109, f. 20–23), S. robusta (Pl. 111, f. 1–4), S. rubra (Pl. 111, f. 5–8), and S.
subaristata. Curiously, S. rigescens has propagula (Pl. 110, f.
18–19) borne ventrally on the costa, as in sect. Collotortula, but is
clearly in sect. Syntrichia near S. caninervis. Note also Syntrichia
leucostega, not seen, for which ventral costal propagula are illustrated by
Kramer (1988).
Syntrichia sect. Aesiotortula Zand., sect. nov. Type: S. pagorum
(Milde) Amann.
A sectione typica plantis parvioribus quam
congeneribus, foliis ligulatis vel spathulatis, marginibus omnino planis,
papillis bifidis, propagulis si effectis foliaceis, in termino caulis vel in
apicibus foliorum deciduis portatis, costa supra regione folii mediana
perincrassata et in sectione transversalis semicircularibus differt.
Differs
from the typical section by the following combination of characters: plants
rather small for the genus, leaves ligulate to spathulate, margins plane
throughout, papillae bifid, and propagula when present leaf-like, borne
terminally on the stem or as deciduous leaf apices. The costa is often very
strongly thickened above midleaf and semicircular in transverse section. This
group of species includes S. ammonsiana (Pl. 105, f. 12–16), S.
baileyi, S. bartramii, S. chisosa (Pl. 107, f. 13–16), S.
epilosa (Pl. 108, f. 1–9), S. pagorum (Pl. 109, f. 10–11), S.
phaea (Pl. 109, f. 23–27) and S. pygmaea (Pl. 110, f. 10–16). Syntrichia
papillosa, although here placed with sect. Collotortula because of
the clearly trigonous upper laminal cells and clavate to spherical propagula,
also has plane margins and a much thickened costa.
Section
Aesiotortula (as the “Tortula laevipila-Tortula pagorum-Gruppe”)
was not dealt with by Kramer (1980), who viewed it as a complex requiring special
study. Certain of the infraspecific taxa of S. laevipila may also belong
here, but not var. laevipila (Pl. 109, f. 1–9) itself. Syntrichia
pygmaea is only tentatively placed in this section; it is propaguliferous,
however, by a deciduous leaf apex and has plane margins. On the other hand,
species of similarly small stature in many genera have a tendency to lose or
exhibit less distinctly some important characters, including recurvature of the
lower margins. Syntrichia pygmaea is also unusual for its ventral leaf
surface very narrowly grooved along the costa and ventral costal cells
elongate, which may be an indication of a relationship with the
Bryoerythrophylleae for this species. The upper laminal cells of the most
highly derived of the species of sect. Aesiotortula are relatively large
for the genus, generally 13–15 µm in diameter; species with upper laminal cells
small (ca. 8–10 µm in diameter), such as S. bartramii and S. chisosa,
are less distinctive. The cleistocarpous genus Phascopsis could be seen
as having been derived from this section of Syntrichia, but is
apparently a relict of a more primitive lineage (see Cladograms 13 and 14).
The
New World austral species Syntrichia epilosa is much the same as S.
bartramii of western North America and maybe synonymous; a parallel
southern South America and southwestern North America distribution is that of
the austral Pseudocrossidium crinitum and its Arizona population
previously known as Tortula aurea (see treatment of Pseudocrossidium).
Additional,
selected literature: Allorge (1938), Anderson (1943), Barkman (1963), Bartram
(1924b, 1926b), Bewley (1972, 1973a,b), Bewley et al. (1974), Bizot (1954,
1956), Blomquist (1930), Boudier (1992), Casas de Puig (1975a), Casas de Puig
and Molinas (1975), Catcheside (1980, 1992), Dedkov et al. (1989), Doei et al.
(1985), El-Oqlah et al. (1988), Hedenäs (1989b), Kalenov (1977), Kramer (1978),
Lazarenko (1959), Lightowlers (1985a, 1986a,b,c), Magill et al. (1983), Maya
(1986), Mishler (1984a, 1985a,b, 1986a, 1987a, 1990), Mishler and Newton (1987,
1988), Mishler and Oliver (1991), Mishler and Scheirer (1983), Oliver and
Mishler (1988, 1990), Ovezova (1989), Saito (1973a), Side and Whitehouse
(1974), Steere (1939a, 1940), Stone (1971), Studlar et al. (1984), Toth (1987),
Tuba (1984, 1985), Willis (1964), Zander (1989).
Number
of accepted species: 82.
New
heterotypic synonymy: Tortula abruptinervis Dix. = Syntrichia pygmaea
(Dus.) Zand. Tortula kingii Robins. = Syntrichia percarnosa (C.
Müll.) Zand. Tortula lemniscata Zand. = Syntrichia aculeata
(Wils.) Zand. Tortula nigra Zand. = Syntrichia percarnosa (C.
Müll.) Zand. Tortula tanganyikae Dix. = Syntrichia amphidiaceus
(C. Müll.) Zand.
Species
examined: S. aculeata (BUF, NY), S. alpestris (BM), S.
ammonsiana (BUF), S. amphidiaceus (BM, BUF, TENN), S. amplexa
(CU), S. anderssonii (NY), S. andicola, S. antarctica
(NY), S. baileyi (NY), S. bartramii (BUF), S. bipedicellata
(NY), S. bogotensis (NY), S. bolanderi (BUF), S. brandisii
(NY), S. cainii (BUF, TRTC), S. caninervis (BUF, H), S.
cavallii (NY, US), S. chisosa (BUF), S. ciliata (H), S.
conferta (NY), S. costesii (NY), S. didymodontoides (H), S.
epilosa (COLO), S. filaris (H, US), S. flagellaris (BUF), S.
fontana (S) as Tortula rivularis, S. fragilis, S.
fuscoviridis (NY), S. geheebiaeopsis (NY), S. gemmascens
(NY), S. gromschii (US), S. inermis, S. intermedia (BUF), S.
jaffuelii (NY), S. lacerifolia (NY), S. laevipila (BUF), S.
latifolia, S. limensis (NY), S. linguifolia (L), S.
longimucronata (NY), S. mongolica (NY), S. mollis (US), S.
norvegica, S. obtusissima (BM, BUF, TENN), S. papillosa (BUF,
NY), S. percarnosa (BUF, H, US), S. phaea (BUF), S. pichinchensis
(US), S. princeps, S. prostrata (US), S.
pseudodesertorum (S), S. pseudorobusta (NY), S. pygmaea (BM,
BUF, MICH, NY), S. ramosissima (NY), S. reflexa (NY), S.
rigescens (FU), S. robusta (NY), S. rubella (NY), S. rubra
(NY), S. ruralis, S. saxicola (NY), S. scabrella (NY), S.
scabrinervis (US), S. schnyderi (NY), S. serrata (NY), S.
serripungens (NY), S. sinensis (NY), S. socialis (S), S.
subaristata (NY), S. virescens (BUF), S. viridula (NY).
New combinations and statuses:
Syntrichia aculeata (Wils.) Zand., comb. nov. (Barbula
aculeata Wils., Kew J. Bot. 3: 51, 1851; Tortula aculeata (Wils.)
Mitt.).
Syntrichia alpestris (Dix. in Herz.) Zand., comb. nov. (Tortula
alpestris Dix. in Herz., Rep. Spec. Nov. Reg. Veg. 38: 103, 1935).
Syntrichia amphidiacea (C. Müll.) Zand., comb. nov. (Barbula
amphidiacea C. Müll., Linnaea 42: 332, 1879; Tortula amphidiacea (D.
Müll.) Broth.).
Syntrichia amplexa (Lesq.) Zand., comb. nov. (Barbula
amplexa Lesq., Trans. Am. Phil. Soc. n. ser. 13: 5, 1865; Tortula
amplexa (Lesq.) Steere in Grout).
Syntrichia anderssonii (Ångstr.) Zand., comb. nov. (Tortula
anderssonii Ångstr., Oefv. K. Vet. Ak. Foerh. 29(4): 6, 1872).
Syntrichia anderssonii var. fagicola (C. Müll.) Zand., comb.
nov. (Barbula conotricha var. fagicola C. Müll., Flora 68:
416, 1885), not seen.
Syntrichia antarctica (Hampe in C. Müll.) Zand., comb. nov. (Barbula
antarctica Hampe in C. Müll., Syn. 1: 642, 1849; Tortula antarctica
(Hampe in C. Müll.) Wils. in Hook. f.).
Syntrichia austroafricana (Kramer) Zand., comb. nov. (Tortula
austroafricana Kramer, J. Hattori Bot. Lab. 65: 92, 1988), not seen but
cf. Kramer, J. Hattori Bot. Lab. 65: 84, 1988.
Syntrichia baileyi (Broth.) Zand., comb. nov. (Tortula
baileyi Broth., Oefv. Finsk. Vet. Soc. Foerh. 33: 97, 1891).
Syntrichia bartramii (Steere in Grout) Zand., comb. nov. (Tortula
bartramii Steere in Grout, Moss Fl. N. Amer. 1: 241, 1939).
Syntrichia bipedicellata (Britt.) Zand., comb. nov. (Tortula
bipedicellata Britt., Bull. Torr. Bot. Cl. 23: 431, 1896).
Syntrichia bogotensis (Hampe) Mitt. (Barbula bogotensis Hampe,
Ann. Sc. Nat. Bot. ser. 5, 3: 349, 1865; Tortula bogotensis (Hampe)
Mitt.).
Syntrichia bolanderi (Lesq. & James) Zand., comb. nov. (Barbula
bolanderi Lesq. & James, Trans. Am. Phil. Soc. n. ser. 13: 5, 1865; Tortula
bolanderi (Lesq. & James) Howe).
Syntrichia brachyclada (Card.) Zand., comb. nov. (Tortula
brachyclada Card., Bull. Herb. Boiss. ser. 2, 5: 1002, 1905), not seen but
cf. Kramer, J. Hattori Bot. Lab. 65: 84, 1988.
Syntrichia brandisii (C. Müll.) Zand., comb. nov. (Barbula
brandisii C. Müll., Flora 61: 82, 1878; Tortula brandisii (C. Müll.)
Broth.).
Syntrichia brevisetacea (F. Müll.) Zand., comb. nov. (Barbula
brevisetacea F. Müll., Analyt. Draw. Austral. Moss. Tab. 4, 1864; Tortula
brevisetacea (F. Müll.) Thér.), not seen but cf. Kramer, J. Hattori
Bot. Lab. 65: 84, 1988.
Syntrichia cainii (Crum & Anders.) Zand., comb. nov. (Tortula
cainii Crum & Anders., J. Elisha Mitchell Sci. Soc. 74: 35, 1958).
Syntrichia campestris (Dus.) Zand., comb. nov. (Tortula
campestris Dus., Ark. Bot. 6(8): 25, 1906), not seen but cf. Kramer,
J. Hattori Bot. Lab. 65: 84, 1988.
Syntrichia caninervis var. spuria (Amann) Zand., comb. nov.
(Tortula spuria Amann, Bull. Murithienne 39: 351, 1916; Tortula
caninervis ssp. spuria (Amann) Kramer), not seen.
Syntrichia caninervis var. abranchesii (Luis.) Zand., comb. nov.
(Tortula abranchesii Luis., Broteria ser. Bot. 14: 115, 1916; Tortula
caninervis var. abranchesii (Luis.) Kramer), not seen.
Syntrichia caninervis var. spuria (Amann) Zand., comb. nov.
(Syntrichia spuria Amann, Fl. Mouss. Suisse 2: 119, 384, 1918).
Syntrichia chisosa (Magill, Delg. & Stark) Zand., comb. nov.
(Tortula chisosa Magill, Delg. & Stark, Ann. Missouri Bot. Gard. 70:
200, 1983).
Syntrichia ciliata (Broth.) Zand., comb. nov. (Tortula
ciliata Broth., Bot. Jahrb. 49: 175, 1912).
Syntrichia conferta (Bartr.) Zand., comb. nov. (Tortula
conferta Bartr., Bryologist 60: 140, 1957; Tortula princeps var. conferta
(Bartr.) Lightowlers).
Syntrichia costesii (Thér.) Zand., comb. nov. (Tortula
costesii Thér., Rev. Chil. Hist. Nat. 25: 298, 1921).
Syntrichia didymodontoides (Broth. in Dryg.) Zand., comb. nov. (Tortula
didymodontoides Broth. in Dryg., Deutsch. Südpolar Exp. 8: 86, 1906).
Syntrichia epilosa (Broth. ex Dus.) Zand., comb. nov. (Tortula
epilosa Dus., Ark. Bot. 6(8): 25, 1906.
Syntrichia epilosa var. pilifera (Thér.) Zand., comb.
nov. (Tortula epilosa var. pilifera Thér., Rev. Chil. Hist.
Nat. 30: 343, 1926), not seen.
Syntrichia filaris (C. Müll. in Neum.) Zand., comb. nov. (Barbula
filaris C. Müll. in Neum., Deutsch. Exp. Int. Polarforsch. 2: 309, 1890; Tortula
filaris (C. Müll. in Neum.) Broth.).
Syntrichia flagellaris (Schimp.) Zand., comb. nov. (Barbula
flagellaris Schimp., Ann. Sci. Nat. Bot. ser. 2, 6: 146, 1836; Tortula
flagellaris Mont. in Gay).
Syntrichia flagellaris var. densiretis (Thér.) Zand., comb.
nov. (Tortula flagellaris var. densiretis Thér., Recueil
Publ. Soc. Havraise Etud. Div. 1917: 7, 1917), not seen.
Syntrichia fontana (C. Müll. in Neum.) Zand., comb. nov. (Barbula
fontana C. Müll. in Neum., Deutsch. Exp. Int. Polarforsch. 2: 308, 1890; Tortula
fontana (C. Müll.) Broth.).
Syntrichia fuscoviridis (Card.) Zand., comb. nov. (Tortula
fuscoviridis Card., Bull. Herb. Boiss. ser. 2, 6: 6, 1906).
Syntrichia geheebiaeopsis (C. Müll.) Zand., comb. nov. (Barbula
geheebiaeopsis C. Müll., Bot. Jahrb. 5; 80, 1883; Tortula geheebiaeopsis
C. Müll.) Broth.).
Syntrichia gemmascens (Chen) Zand., comb. nov. (Desmatodon
gemmascens Chen, Hedwigia 80: 297, 1941; Didymodon gemmascens
Broth., Symb. Sin. 4: 38, 1929, hom. illeg.).
Syntrichia gemmascens var. hopeiensis (Chen) Zand., comb.
nov. (Desmatodon gemmascens var. hopeiensis Chen, Hedwigia
80: 229, 1941).
Syntrichia glacialis (Kunze ex C. Müll.) Zand., comb. nov. (Barbula
glacialis Kunze ex C. Müll., Syn. 1: 634, 1849; Tortula glacialis (Kunze
ex C. Müll.) Mont. in Gay).
Syntrichia gromschii (Thér.) Zand., comb. nov. (Tortula
gromschii Thér., Rev. Bryol. Lichénol. 7: 173, 1935).
Syntrichia jaffuelii (Thér.) Zand., comb. nov. (Tortula
jaffuelii Thér., Rev. Chil. Hist. Nat. 27: 9, 1923).
Syntrichia lacerifolia (Williams) Zand., comb. nov. (Tortula
lacerifolia Williams, Bull. Torr. Bot. Cl. 43: 326, 1916).
Syntrichia leucostega (C. Müll.) Zand., comb. nov. (Barbula
leucostega C. Müll., Syn. 1: 641, 1894; Tortula leucostega (C.
Müll.) Broth.), not seen but cf. Kramer, J. Hattori Bot. Lab. 65: 106,
1988.
Syntrichia leucostega var. trachyneura (Dix.) Zand., comb.
nov. (Tortula trachyneura Dix., Trans. R. Soc. S. Africa 8: 195,
1920; Tortula leucostega var. trachyneura Kramer), not seen but
cf. Kramer, J. Hattori Bot. Lab. 65: 107, 1988.
Syntrichia limensis (Williams) Zand., comb. nov. (Tortula
limensis Williams, Bull. Torr. Bot. Cl. 42: 398, 1915).
Syntrichia linguifolia (Herz.) Zand., comb. nov. (Tortula
linguifolia Herz., Biblioth. Bot. 87: 49, 1916).
Syntrichia longimucronata (X.-j. Li) Zand., comb. nov. (Tortula
longimucronata X.-j. Li, Act. Bot. Yunnan. 3: 107–109, 1981).
Syntrichia magellanica (Mont. in Gay) Zand., comb. nov. (Tortula
magellanica Mont. in Gay, Hist. Fis. Polit. Chile Bot. 7: 145, 1850; Tortula
princeps var. magellanica (Mont. in Gay) Lightowlers), not seen but
cf. Kramer, J. Hattori Bot. Lab. 65: 110, 1988.
Syntrichia mollis (B.&S. ex C. Müll.) Zand., comb. nov.
(Barbula mollis B.&S. ex C. Müll., Syn. 1: 637, 1849; Tortula
mollis (B.&S. ex C. Müll.) Broth.).
Syntrichia obtusissima (C. Müll.) Zand., comb. nov. (Barbula
obtusissima C. Müll., Syn. 1: 640, 1849; Tortula obtusissima (C.
Müll.) Mitt.).
Syntrichia papillosa var. chilensis (Thér.) Zand., comb.
nov. (Tortula papillosa var. chilensis Thér., Rev. Chil.
Hist. Nat. 25: 297, 1921), not seen.
Syntrichia percarnosa (C. Müll.) Zand., comb. nov. (Barbula
percarnosa C. Müll., Linnaea 42: 347, 1879; Tortula percarnosa (C.
Müll.) Broth.).
Syntrichia phaea (Hook. f. & Wils.) Zand., comb. nov.
(Trichostomum phaeum Hook. f. & Wils., Fl. Nov. Zel. 2: 72, 1854; Tortula
phaea (Hook. f. & Wils.) Dix.).
Syntrichia pichinchensis (Tayl.) Zand., comb. nov. (Tortula
pichinchensis Tayl., London J. Bot. 6: 333, 1847).
Syntrichia princeps var. brachycarpa (De Not.) Zand., comb.
nov. (Tortula princeps var. brachycarpa De Not., Atti Univ.
Genova 1: 538, 1869).
Syntrichia princeps var. echinata (Shiffn.) Zand., comb.
nov. (Tortula echinata Schiffn., Oesterr. Bot. Zeitschr. 65: 4–5,
1915; Tortula princeps ssp. echinata (Schiffn.) Kramer).
Syntrichia prostrata (Mont.) Zand., comb. nov. (Tortula
prostrata Mont., Ann. Sc. Nat. Bot. ser. 3, 4: 104, 1845).
Syntrichia pseudorobusta (Dus.) Zand., comb. nov. (Tortula
pseudorobusta Dus., Ark. Bot. 6(8): 19, 1906).
Syntrichia pygmaea (Dus.) Zand., comb. nov. (Tortula
pygmaea Dus., Ark. Bot. 6(10): 8, 1907).
Syntrichia ramosissima (Thér.) Zand., comb. nov. (Tortula
ramosissima Thér., Rev. Chil. Nat. 33: 136, 1926).
Syntrichia reflexa Zand., nom. nov. (Tortula reflexa X.-j.
Li, Acta Bot. Yunnan. 3: 109, 1981, hom. illeg. non Brid.).
Syntrichia robusta (Hook. & Grev.) Zand., comb. nov. (Tortula
robusta Hook. & Grev., Edinburgh J. Sc. 1: 299, 1824).
Syntrichia robusta var. recurva (Lightowlers) Zand., comb.
nov. (Tortula robusta var. recurva Lightowlers, Brit. Ant.
Surv. Bull. 64: 64, 1984).
Syntrichia rubella (Hook. f. & Wils.) Zand., comb. nov.
(Tortula rubella Hook. f. & Wils., Fl. Tasman. 2: 176, 1859).
Syntrichia rubra (Mitt. in Hook. f.) Zand., comb. nov. (Tortula
rubra Mitt. in Hook. f., Handb. New Zealand Fl. 419, 1867).
Syntrichia rubra var. subantarctica (Sainsb.) Zand., comb.
nov. (Tortula subantarctica Sainsb., Svensk. Bot. Tidskr. 44: 72,
1950; Syntrichia rubra var. subantarctica (Sainsb.) Lightowlers).
Syntrichia ruralis var. gigantea (Lesq.) Zand., comb.
nov. (Barbula ruralis var. gigantea Lesq., Mem. California
Acad. Sci. 1: 13, 1868; Tortula ruralis var. gigantea (Lesq.) L.
Koch.), not seen.
Syntrichia ruralis var. gracilis (C. Jens.) Zand., comb.
nov. (Tortula ruralis var. gracilis C. Jens., Medd. Groenland
15: 409, 1898), not seen.
Syntrichia ruralis var. spiralis (Herz.) Zand., comb.
nov. (Tortula ruralis var. spiralis Herz., Biblioth. Bot. 87:
49, 1916), not seen.
Syntrichis ruralis var. submamillosa (Kramer) Zand., comb.
nov. (Tortula ruralis var. submamillosa Kramer, Bryophyt.
Biblioth. 21: 127, 1980), not seen.
Syntrichia ruralis var. subpapillosissima (Biz. &
Pierr.) Zand., comb. nov. (Tortula ruralis var. subpapillosissima
Biz. & Pierr., Acta Bot. Acad. Sci. Hung. 18: 10, 1973; Tortula
ruraliformis var. subpapillosissima (Biz. & Pierr.) Kramer), not
seen.
Syntrichia saxicola (Card.) Zand., comb. nov. (Tortula
saxicola Card., Bull. Herb. Boiss. ser. 2, 5: 1002, 1905).
Syntrichia scabrella (Dus.) Zand., comb. nov. (Tortula
scabrella Dus., Ark. Bot. 6(10): 4, 1907).
Syntrichia scabrinervis (C. Müll.) Zand., comb. nov. (Barbula
scabrinervis C. Müll., Syn. 1: 634, 1849; Tortula scabrinervis (C.
Müll.) Mitt.).
Syntrichia schnyderi (C. Müll.) Zand., comb. nov. (Barbula
schnyderi C. Müll., Linnaea 43: 434, 1882; Tortula schnyderi (C.
Müll.) Broth.).
Syntrichia serrata Zand., comb. nov. (Tortula serrata
Dix., New Zealand Inst. Bull. 3(3): 146, 1923, nom. legit. contrary to
Index Muscorum, also cf. Lightowlers, J. Bryol. 13: 373, 1985).
Syntrichia serripungens (Lor. & C. Müll.) Zand., comb. nov. (Barbula
serripungens Lor. & C. Müll., Linnaea 42: 360, 1879; Tortula
serripungens (Lor. & C. Müll.) Broth.).
Syntrichia serripungens var. excesa (C. Müll.) Zand., comb. nov.
(Barbula serripungens var. excesa C. Müll., Linnaea 42: 351,
1879.
Syntrichia socialis (Dus.) Zand., comb. nov. (Tortula
socialis Dus., Ark. Bot. 6(10): 6, 1907).
Syntrichia subaristata (B.&S. ex C. Müll.) Zand., comb. nov.
(Barbula subaristata B.&S. ex C. Müll., Syn. 1: 644, 1849; Tortula
subaristata (B.&S. ex C. Müll.) Broth.).
Syntrichia virescens var. bizotiana (Kramer) Zand., comb.
nov. (Tortula virescens ssp. bizotiana Kramer, J. Hattori
Bot. Lab. 65: 123, 1988, nom. nov. for Tortula virescens ssp. bizotii
(Laz.) Kramer, Bryophyt. Biblioth. 21: 102, 1980, comb. inval. basion.
inval.; Tortula bizotii Laz., Vopr. Evol. Biogeogr. Genet. Sel. 145,
1960, nom. inval.), not seen but cf. Kramer, J. Hattori Bot. Lab.
65: 84, 1988.
Syntrichia virescens var. iranica (Kramer) Zand., comb.
nov. (Tortula virescens var. iranica Kramer, Bryophyt.
Biblioth. 21: 101, 1980), not seen.
Syntrichia viridula (C. Müll.) Zand., comb. nov. (Barbula
viridula C. Müll., Nuov. Giorn. Bot. Ital. n. ser. 4: 114, 1897; (Tortula
viridula (C. Müll.) Broth.; the isotype at NY is probably S. fragilis).
