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BFNA Title: Sarmentypnum |
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CALLIERGONACEAE-- Sarmentypnum xxx. SARMENTYPNUM
Tuomikoski & T. J. Koponen, Ann. Bot. Fennici 16: 223. 1979 * Plants medium-sized
to rather large, green, yellow-green, brownish, red-brown or with clear red
colors. Stem sparsely to richly
radially branched, with central strand and without or sometimes with a partial
outer hyalodermis, cells within cortex thin-walled; outer pseudoparaphyllia
broad, triangular to broader than long and often irregular in outline;
rhizoids or rhizoid initials at various points on the leaves, or on scattered
points or in rows on the stem; axillary hairs with 1--7 distal hyaline or
early brown cells. Stem leaves
straight or falcate, triangular to ovate or narrowly ovate, either gradually
narrowed to a long-acuminate, acuminate or obtuse apex, or suddenly narrowed
to a shortly acuminate to broadly rounded and usually apiculate apex, not or
indistinctly plicate, concave or strongly concave; margin entire, sinuous or
denticulate; costa single, ending ca. 60% way up leaf to long-excurrent;
median lamina cells incrassate or thin-walled, eporose to porose; alar cells
quadrate or shortly to long-rectangular, hyaline and thin-walled, or,
especially near costa or when old, red or brown and incrassate, inflated or
strongly inflated, in transversely triangular group, distinctly delimited
from surrounding cells, not or hardly decurrent, or in S. tundrae long-and broadly decurrent. Sexual condition dioicous. Inner
perichaetial leaves not plicate; vaginula naked. Capsule curved and ± horizontal; lacking a separating annulus;
exostome reticulate on proximal exterior, margin dentate distally. Spores 11--24 \um. Species 7 (5
in the flora): widespread. The leaves of Sarmentypnum species are either
moderately curved to straight and gradually narrowed to the apex, or straight
and suddenly narrowed to a usually apiculate apex. Red colors are frequent in
several species, and the leaves are non-decurrent in all except S. tundrae. The alar groups are
transversely triangular and consist of inflated, and (at least when young)
usually hyaline cells. Except for S. tundrae,
Sarmentypnum species typically
occur in intermediately mineral-rich fens. Most species are found in
nutrient-poor habitats, but S. tundrae grows
in somewhat nutrient-rich places. Sarmentypnum
and Warnstorfia are well separated
based on molecular evidence, and morphological features differentiating the
two are mentioned under the latter genus. Sarmentypnum
collections with falcate leaves and lacking sporophytes or red pigments
are frequently confused with falcate-leaved species with transversely triangular
alar groups in the Drepanocladus
aduncus complex. The frequently present rhizoid initials occurring close
to leaf apices in all Sarmentypnum species except S. trichophyllum (in which they are
rare) are very helpful in differentiating these from Drepanocladus species, which always lack such initials. In
addition, Sarmentypnum species are
usually radially branched, the leaves generally give an "ordered"
impression when viewing entire plants, and the distal portion of the axillary
hairs frequently consists of more than two cells. Drepanocladus species are distichously branched, their leaves
give a more unordered impression, and their axillary hairs have almost
invariably only one or two distal cells, except D. polygamus, which has 1--3 differentiated hyaline distal cells. When sporophytes are present, the
numerous sporophytic and perichaetial characters mentioned in the
introduction to the family make the separation of these taxa easy. SELECTED
REFERENCES Hedenäs, L. 1993. A generic revision of the Warnstorfia-Calliergon
group. J. Bryol. 17: 447--479. Hedenäs, L. 2006. Additional insights into the phylogeny of Calliergon,
Loeskypnum, Straminergon, and Warnstorfia (Bryophyta:
Calliergonaceae). J. Hattori Bot. Lab. 100: 125--134. Janssens,
J. A. 1983. Past and
extant distribution of Drepanocladus in
1. Stem leaves long- and broadly
decurrent; plants never red........ 1.
Sarmentypnum tundrae 1. Stem leaves not or hardly decurrent;
plant color variable, often with red colors present. 2. Stem and branch leaves with short-
or long-excurrent costa; branch apices (and often shoot apex) of tightly
convolute leaves ; axillary hairs 2--7-celled, abundant, early becoming
brown. 2. Leaf costa not excurrent; branch and
shoot apices not pencil-like; axillary hairs 1--4(--5)-celled, abundant or
not, hyaline when young. 3. Stem leaves oblong or ovate, at apex
rounded, or ± suddenly narrowed to acute or very slightly acuminate apex,
with short apiculus (sometimes lost in old leaves). 3. Stem leaves gradually narrowed to
acuminate apex from an ovate, ovate-lanceolate or ovate-triangular; apiculus
absent. 4. Stem leaves falcate or straight; leaf
margin distinctly denticulate distally or proximally or both; marginal basal
cells usually differentiated from the median cells (least distinct in
submerged plants), often rectangular or distinctly widened and forming
distinct border; leaves concave and acumen sometimes furrowed but not
strongly so . 4. Stem leaves straight or almost so,
erect; leaf margin entire or at most
very finely, obtusely denticulate; marginal cells at widest part of
leaf usually similar to the cells farther in, rarely slightly differentiated.
Leaves strongly concave, acumen deeply furrowed (rare Arctic species). 1. Sarmentypnum
tundrae (Arnell)
Hedenäs, J. Hattori Bot. Lab. 100: 133.
2006 Amblystegium
tundrae Arnell in Lindberg
and Arnell, K. Svensk. Vet. Ak. Handl. 23(10): 128. 1890; Drepanocladus exannulatus var. tundrae (Arnell) Warnstorf; D. tundrae (Arnell) Loeske; Warnstorfia tundrae (Arnell) Loeske Plants medium-sized, green, yellowish or
brownish, never red; branch and shoot apices not of convolute leaves; cells
of stem epidermis not differentiated; axillary hairs with 1--2-celled distal
portion, hyaline when young. Stem
leaves slightly falcate or
straight, from ovate or ovate-triangular base gradually narrowed
to a broadly acuminate and often
hooked apex , slightly concave; margins denticulate proximally, entire or
sparsely denticulate distally; costa ending 70--85% of the way up the leaf;
alar cells in distinctly delimited, transversely triangular groups almost reaching the costa, longly and
broadly decurrent. Mineral and
somewhat nutrient-rich fens and on shores, occasionally submerged in lakes;
0--1560 m; Greenland; Alta., B.C., Man., Nfld., N.W.T., Nunavut, Ont., Que., Yukon; Alaska, Maine,
Wyo.; n Eurasia; incorrectly reported from Australia. Sarmentypnum tundrae shares no morphological apomorphies
with the other species in Sarmentypnum
or with those in Calliergon, which
are the two possible generic choices for these species based on morphology.
It is here included in Sarmentypnum from
molecular evidence. Sarmentypnum tundrae differs from all
other members of the genus by its longly and more or less broadly decurrent
stem leaves. Additional features that aid in separating this species from S. exannulatum and S. procerum are the rather weakly
falcate stem leaves and the total lack of red colors. 2.
Sarmentypnum exannulatum (Schimper) Hedenäs, J. Hattori
Bot. Lab. 100: 132. 2006 Hypnum exannulatum Schimper,
Bryol. Eur. 6: 110. fig. 603. (fasc. 57--61, Mon. 34. 23) 1854; Drepanocladus exannulatus (Schimper) Warnstorf; D. exannulatus var. purpurascens (Schimper) Herzog; D. exannulatus var. rotae (De Notaris) Loeske; Warnstorfia exannulata (Schimper)
Loeske; W. exannulata var. purpurascens (Schimper) Tuomikoski
& T. J. Koponen Plants medium-sized, green, yellowish, or
with red secondary coloration; branch and shoot apices not pencil-like; cells
of stem epidermis usually widened to form a partial hyalodermis; axillary
hairs with 1--4-celled distal portion, hyaline when young. Stem leaves falcate or strongly so,
more rarely straight, gradually narrowed to acuminate apex from an ovate or
ovate-triangular base, concave; margins distinctly denticulate distally or proximally or both; costa ending
60--95% of the way up the leaf; alar cells in a distinctly delimited,
transversely triangular group that reaches costa or almost so, not decurrent. Intermediately
mineral-rich fens, often around springs or in late snow-beds, sometimes
submerged in lakes; 0--4150 m; Greenland;
Alta., B.C., Man., N.B., Nfld., N.W.T., N.S., Nunavut, Ont., Que., Sask., Yukon; Alaska, Calif., Colo., Idaho, Maine,
Md., Mass., Mich., Minn, Mont., Nev., N.H., N.Y., Oreg., Pa., Utah, Wash.,
Wyo.; Mexico; South America; Eurasia; Africa; Australia; Pacific Islands (New
Zealand). Among the
wetland species with leaves that narrow gradually to the leaf apex and are
gradually curved throughout, Sarmentypnum exannulatum is one of the most frequent. In S.
exannulatum, the
marginal lamina cells in the widest part of the stem leaves are often
distinctly rectangular or distinctly widened or both. This is never as
pronounced in other Sarmentypnum species as it can be in S. exannulatum. However, because this feature is
rather variable it should be used with caution when identifying material. Warnstorfia fluitans and W. pseudostraminea differ from S.
exannulatum in
being autoicous, having mainly narrowly triangular to lanceolate
pseudoparaphyllia, hardly ever with pure red colors, and with more weakly
differentiated alar groups. The alar groups in W. fluitans are usually narrower, i.e., reaching less far
distally in the leaf than in is the case with S. exannulatum, whereas those in W. pseudostraminea often form ovate groups together with the supra-alar
cells. Certain phenotypes of S. exannulatum from springs, with inflated alar cells
more or less in only one row along the leaf base, have been called W. exannulata var.
purpurascens. However, transitions
between these phenotypes and those from other habitats exist, and if the
different phenotypes are cultivated together the resulting plants cannot be
separated from each other (based on north European material). Modifications
with less well-delimited alar cells do occur. The only difference mentioned
in the literature between the most common expressions of Sarmentypnum exannulatum and straight-leafed forms, which have
been called “orthophyllus Mönkemeyer,”
is in the leaf curvature. Although the type of the latter was not seen, a
separate taxon cannot be recognized for straight-leaved plants. 3. Sarmentypnum
trichophyllum
(Warnstorf) Hedenäs, J.
Hattori Bot. Lab. 100: 133. 2006 Drepanocladus rotae var. trichophyllus Warnstorf, Krypt. Fl. Plants medium-sized or rather large, green,
brown, or when emergent above water surface with red secondary coloration;
branch apices (and often shoot apex) ± aciculate by long-convolute leaves;
cells of stem epidermis not widened, or occasionally in small parts (less
than 10% of circumference) slightly widened; axillary hairs with 2--7-celled
distal portion, early brown. Stem
leaves straight or more rarely falcate, gradually narrowed to a long- or
usually very long- acuminate apex from an ovate to triangular base, concave;
margins denticulate; costa strong, short- or long- excurrent (excurrent
portion up to 1/3 leaf length); alar cells in distinctly delimited,
transversely triangular group that reaches costa, not decurrent. Normally
submerged or floating in mineral-rich water, but with a relatively high pH
(5.1--6.7 in Europe), mostly in small lakes, water-filled kettle holes,
ox-bows and other small aquatic habitats; 0--250 m; Man., Nunavut; w Alaska; South America (Colombia), n Eurasia. Sarmentypnum
trichophyllum is
occasionally reminiscent of S.
exannulatum, but its distinctly acicular branch and shoot apices,
excurrent costa and long, early brown axillary hairs (shorter and normally
hyaline in S. exannulatum) separate
them. For the differences between Sarmentypnum trichophyllum and Drepanocladus longifolius, see the
note after the latter. Sarmentypnum
trichophyllum has been reported from many areas of 4. Sarmentypnum
pseudosarmentosum
(Cardot & Thériot) Hedenäs, J.
Hattori Bot. Lab. 100: 133. 2006 Hypnum pseudosarmentosum Cardot & Thériot, Univ. California
Publ. Bot. 2: 305, plate 27, fig.
2. 1906; Calliergon pseudosarmentosum (Cardot
& Thériot) Brotherus; Drepanocladus
pseudosarmentosus (Cardot & Thériot) Persson; Warnstorfia pseudosarmentosa (Cardot & Thériot) Tuomikoski
& T. J. Koponen Plants medium-sized, with red secondary
coloration (entirely green plants not seen); branch and shoot apices not
acicular; cells of stem epidermis widened in part of stem circumference and
forming a partial hyalodermis; axillary hairs with 1--3-celled distal
portion, hyaline when young. Stem
leaves straight and erect or weakly homomallous, gradually narrowed to an
acuminate or sometimes blunt apex from
an ovate or triangular-ovate base, apex sometimes hooked, concave, channeled to almost tubular near apex;
margins entire or at most in part minutely, obtusely denticulate; costa
ending shortly below leaf apex; alar cells in distinctly delimited,
transversely triangular group that reaches costa or almost so, not decurrent. In apparently
spring-influenced habitats or along the shores of brooks; 0--1700 m; Sarmentypnum psuedosarmentosum is superficially similar to Sarmentypnum
sarmentosum in habit, with rather thick shoots and short and often curved
branches with obtuse endings due to densely inserted and relatively
short-acuminate leaves. The leaves, however, are more gradually narrowed than
in S. sarmentosum, and the usually
acuminate or short-acuminate leaf apices in the first of these species make
the separation easy. Sarmentypnum pseudosarmentosum is close to S. exannulatum, from which it differs
mainly in the more strongly concave leaves that are mostly deeply furrowed to
almost tubular distally, and in its entire leaf margins. Sometimes the stem
leaf lamina is weakly plicate (when moist) in S. pseudosarmentosum,
which does not occur in S. exannulatum.
Few specimens of S. pseudosarmentosum were available for study and it
is therefore difficult to judge its status. It may be a good species, or it
may represent an extreme, Arctic modification of S. exannulatum. 5. Sarmentypnum
sarmentosum
(Wahlenberg) Tuomikoski & T.J. Koponen, Ann. Bot. Fennici
16: 223. 1979 Hypnum sarmentosum
Wahlenberg, Fl. Lapp. 380. 1812; Calliergon
sarmentosum (Wahlenberg) Kindberg; C.
sarmentosum var. beringianum (Cardot
& Thériot) Grout; C. sarmentosum var.
fallaciosum (Milde) G. Roth; C. sarmentosum var. fontinaloides (Berggren) G. Roth; Warnstorfia sarmentosa (Wahlenberg)
Hedenäs Plants medium-sized, with red or dark red
secondary coloration, sometimes green; branch and shoot apices not
pencil-like; cells of stem epidermis not widened; axillary hairs with 1--4(--5)-celled
distal portion, hyaline when young.
Stem leaves straight, loosely imbricate or erect-spreading, oblong, ovate
or narrowly ovate, distally ± suddenly narrowed to a rounded- or
acute-apiculate apex, the apiculus (rarely absent) often bent inwards over
leaf, concave; margins entire or nearly so; costa ending 80--95% of way up
leaf; alar cells in a distinctly delimited, transversely triangular group
that gradually passes into strongly incrassate cells near costa, not or
hardly decurrent. Intermediately
mineral-rich fens, often around springs or in late snow-beds, sometimes
submerged in lakes; 0--3960 m;
Greenland; Alta., B.C., Man., Nfld., N.W.T., Nunavut, Ont.,
Que., Yukon; Alaska, Colo., Mont., N.H., N.Y., Wash., Wyo.; South America,
Eurasia (including Papua New Guinea), e Africa, Australia, Pacific Islands
(New Zealand), Antarctica. Sarmentypnum sarmentosum is easily separated from other North
American species of the genus by its oblong or ovate stem leaves, suddenly
narrowed to a rounded- or acute-apiculate
apex. The apiculus is most distinct in young leaves and is rarely
completely lacking, but may be eroded in older leaves. In late snow beds or
in cold springs in higher mountains and in the Sarmentypnum sarmentosum was earlier placed in the genus Calliergon because of general
similarities in leaf shape and orientation. However, in the frequent presence
of red pigments, appearance of the alar groups, nondecurrent leaves, general
size of the plants, and in its habitat preferences it is more similar to Sarmentypnum species, such as S. exannulatum. In addition, the differences between S.
sarmentosum and
S. exannulatum are bridged by the
South American Sarmentypnum luipichense (R. S. Williams) Hedenäs, which is
closely related to S. sarmentosum but has leaves more gradually narrowed to the apex and
more or less denticulate leaf margins, and S. pseudosarmentosum, which, despite a leaf shape that is similar
to that in straight-leaved specimens of S.
exannulatum is reminiscent of S. sarmentosum in its habit and entire leaf margins.
Morphology does not justify an exclusion of S. sarmentosum from the genus, and recent molecular
results also support the present treatment of the species. Sarmentypnum procerum (Renauld & Arnell) Hedenäs (Drepanocladus procerus (Renauld & Arnell)
Warnstorf), is widespread and rather common in the Boreal Zone of northern
Europe, and, though unknown in OTHER
REFERENCES Mogensen, G. 1995. Warnstorfia trichophylla (Warnst.) Tuom. et T. Kop.,
a bryophyte new to Greenland (Musci, Amblystegiaceae). Lindbergia 20: 3--4. |
