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BFNA Title: Calliergonaceae |
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XX.
CALLIERGONACEAE (Kanda) Vanderpoorten et al. Lars Hedenäs Plants medium-sized to large; greenish,
yellowish, brownish, or sometimes red. Stem
distichously or radially branched, sometimes nearly unbranched; cortex of
small and incrassate cells; hyalodermis present or not; pseudoparaphyllia
foliose; paraphyllia absent; rhizoids red-brown, smooth, slightly branched;
rhizoids or rhizoid initials on stem at or just before leaf insertions or
from various points on stems and leaves; axillary hairs mostly well
developed. Stem leaves straight or
falcate (gradually curved) or from straight base suddenly curved, plicate or
smooth; costa single and mostly long, or double and usually short; median
laminal cells linear or short-linear, 1-stratose, smooth; alar cells
differentiated or not, often inflated. Inner
perichaetial leaves straight and erect, lanceolate or ovate or oblong or
slightly obovate, plicate or smooth; costa usually well developed, single;
vaginula with paraphyses or naked. Seta
long, smooth. Capsule cylindric,
curved and horizontal, stomata long-pored; annulus separating or not;
operculum conic; peristome perfect, exostome yellow-brown or brownish, more
or less reticulate on outer surface
proximally, rarely entirely cross-striolate, papillose distally, with dentate
or slightly dentate margin and an exostome border ± widened at zone of
transition in outer peristomial layer pattern, endostome with high basal
membrane, segments long and not or narrow-perforate, cilia nodose. Calyptra cucullate, smooth. Genera 7,
species ca. 22 (7 genera, 19 species in the flora): temperate to sub-polar
regions worldwide, tropical mountains. Genera of the
Calliergonaceae are fairly well circumscribed based on both morphology and
molecular data, and are well differentiated from the Amblystegiaceae. All
species, except sometimes Scorpidium
revolvens and S. scorpioides,
have a reticulate outer surface of the proximal portion of the exostome,
whereas all members here treated in the Amblystegiaceae, except Tomentypnum and Conardia, are cross-striolate on the outer basal exostome. Many
Calliergonaceae species become a translucent red when growing in habitats
exposed to sunlight, this never being the case among the Amblystegiaceae. The genera Calliergon, Loeskypnum, Sarmentypnum,
Straminergon, and Warnstorfia are further characterized
by many morphological features. When shoots grow erect, these are radially
rather than distichously branched, except in species that are sparsely
branched. Rhizoid initials, and sometimes also rhizoids, are frequently found
in various parts of the leaf lamina, especially close to the leaf apex.
Leaf-borne rhizoid initial cells are easily recognized in being slightly
wider than the surrounding lamina cells and in lacking the pigments of the
latter. Among the Amblystegiaceae genera, rhizoid initials and rhizoids are
found in leaves only in Conardia,
in which they often occur both close to leaf apices and on the costa
abaxially, and in Tomentypnum, in
which they are found only on the abaxial surface of the costa. The inner
perichaetial leaves are smooth, the vaginula naked or bearing at most a few
paraphyses, and annulus is not separating. The two genera Scorpidium and Hamatocaulis, which lack the character states enumerated for the
other five genera, are characterized by their small alar groups or by lacking
differentiated alar cells, and a unique form of leaf curvature. Curved leaves
in members of these two genera have an almost straight base and are abruptly
curved above, whereas curved leaves in other members of this family and in
the Amblystegiaceae are gradually curved along their entire length. SELECTED
REFERENCES Hedenäs, L.
1993. A generic revision of the Warnstorfia-Calliergon
group. J. Bryol. 17: 447--479. Hedenäs, L. 2003. Amblystegiaceae (Musci).
Flora Neotropica, Monograph 89:
1--107. Hedenäs, L. 2006. Additional
insights into the phylogeny of Calliergon, Loeskypnum, Straminergon,
and Warnstorfia (Bryophyta: Calliergonaceae). J. Hattori Bot. Lab. 100: 125--134. Hedenäs, L., G. Oliván and
P. Eldenäs. 2005. Phylogeny of the Calliergonaceae (Bryophyta) based on
molecular and morphological data. Plant
Syst. Evol. 252: 49--61. Hedenäs, L. and A. Vanderpoorten. 2007. The
Amblystegiaceae and Calliergonaceae. In: A. E. Newton and R. Tangney (eds.).
Pleurocarpous Mosses: Systematics and Evolution. The Systematics Association
Special Volume Series 71. Pp. 163--176. Vanderpoorten, A., L. Hedenäs, C. J.
Cox and A. J. Shaw. 2002. Phylogeny and morphological evolution of the
Amblystegiaceae (Bryopsida). Molec. Phylog. Evol. 23: 1--21. Vanderpoorten,
A., L. Hedenäs, C. J. Cox and A. J. Shaw. 2002. Circumscription,
classification, and taxonomy of Amblystegiaceae (Bryopsida) inferred from
nuclear and chloroplast DNA sequence data and morphology. Taxon 51: 115--122. 1. Stem with full or partial hyalodermis. 2. Inflated alar cells numerous and forming a large
transverse-triangular group that reaches from leaf margin ± to costa; rhizoid
initials frequently present in leaves, especially near leaf apex. 7. Sarmentypnum
(in part), p. xx. 2. Inflated alar cells few, in small transverse-triangular
or isodiametric group that reaches at most about halfway from leaf margin to
costa; rhizoid initials never present in leaves. 2. Scorpidium, p. xx 1. Stem without an hyalodermis. 3. Stem lacking central strand; stem leaves usually
plicate; alar cells of stem leaves not differentiated from other basal
cells...........................................................................
1. Hamatocaulis, p. xx. 3. Stem with central strand; stem leaves not or hardly
plicate; alar cells of stem leaves differentiated, small or inflated, sharply
or gradually delimited from surrounding cells. 4. Alar cells of stem leaves slightly inflated, incrassate
to strongly so, indistinctly delimited from surrounding cells; axillary hairs
well developed, abundant, with early yellowish or brownish distal cells;
plant often yellow-brown or copper-brown, never red.... 4. Loeskypnum,
p. xx. 4. Alar cells of stem leaves inflated or strongly so,
thin-walled or sometimes incrassate, distinctly delimited from surrounding
cells, except in Calliergon cordifolium and Warnstorfia fluitans;
axillary hairs well developed and abundant or small and rare, hyaline except
in Sarmentypnum trichophyllum (brownish); plant color various,
sometimes red. 5. Stem leaves straight or falcate, from (oblong-ovate or)
ovate or ± triangular or cordate basal part gradually narrowing to acuminate
to acute or obtuse leaf apex. 6. Dioicous; translucent red color often present;
pseudoparaphyllia broad; alar cells forming a large transversely triangular
group that is distinctly delimited from the surrounding cells and ± reaches
the costa (in late snow beds or in cold springs in mountainous areas,
modifications with less well delimited alar groups occur). 7. Sarmentypnum
(in part) , p. xx. 6. Autoicous; translucent red color extremely rare; red
colors when present usually overlain by brown pigments, giving a reddish
brown color; at least some outer pseudoparaphyllia narrow, often lanceolate;
alar cells forming a rather narrowly transversely triangular group that may
reach the costa, or an isodiametric group which, together with the supra-alar
cells, often form an ovate group along basal margin, alar groups somewhat
indistinctly delimited from the surrounding cells. 5. Warnstorfia, p.
xx. 5. Stem leaves ± straight and ovate, broadly ovate, or
rounded-triangular, in distal part ± abruptly narrowing to rounded,
rounded-obtuse or rounded-apiculate apex. 7. Stem leaves broadly ovate to broadly rounded-triangular,
with broadly obtuse or rounded apex; axillary hairs 2--8(--10)-celled, large
and abundant. 3. Calliergon, p. xx. 7. Stem leaves ovate to rounded-triangular, with shortly
rounded-apiculate or rounded apex; axillary hairs 1--4(--5)-celled, weak and
usually sparse. 8. Stem leaf apex rounded; alar groups ovate or broadly
ovate, along basal margin of leaf; plants usually sparsely branched; plants
pale or whitish green to yellow-green.............................. 6. Straminergon,
p. xx. 8. Stem leaf apex usually rounded-apiculate, rarely
rounded; alar groups ± transverse-triangular; plants sparsely to more richly
branched; plants deep green to clear red................. 7. Sarmentypnum (in
part) , p. xx. |