A Revision of the Infrageneric Taxa of Fissidens

Author(s): Ronald A. Pursell and Maria A. Bruggeman-Nannenga

Source: The Bryologist, 107(1):1-20. 2004.
Published By: The American Bryological and Lichenological Society, Inc.
DOI: http://dx.doi.org/10.1639/0007-2745(2004)107[1:AROTIT]2.0.CO;2
URL: http://www.bioone.org/doi/full/10.1639/0007-2745%282004%29107%5B1%3AAROTIT
%5D2.0.CO%3B2

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 THE BRYOLOGIST
A JOURNAL OF BRYOLOGY AND LICHENOLOGY

VOLUME 107 SPRING 2004 NUMBER 1

The Bryologist 107(1), pp. 1 20

Copyright q 2004 by the American Bryological and Lichenological Society, Inc.

A Revision of the Infrageneric Taxa of Fissidens

RONALD A. PURSELL
208 Mueller Laboratory, Department of Biology, The Pennsylvania State University, University Park, PA 16802,
U.S.A. e-mail: rap10@psu.edu

MARIA A. BRUGGEMAN-NANNENGA
University of Utrecht, Department of Plant Ecology and Evolutionary Biology, Herbarium Division, W. D. Unnik-
gebouw, P. O. Box 80102, NL-3508 TC Utrecht, The Netherlands

Abstract. The Fissidentaceae are recognized to include a single genus, Fissidens, which is
divided into four subgenera, Aloma, Fissidens, Octodiceras, and Pachyfissidens, on the basis of
new taxonomically useful characters i.e., peristome type, costa type, and number of files of exo-
thecial cells. Subgenera Aloma and Octodiceras are not subdivided. Subgenus Fissidens consists
of sections Fissidens and Sarawakia, comb. nov. Subgenus Pachyfissidens is divided into three
sections: Amblyothallia, comb. nov., Crispidium comb. nov., and Pachyfissidens. The most prim-
itive species of the genus are found in section Amblyothallia, while the most advanced species
are in subgenera Aloma and Octodiceras. A key to the subgenera and sections is provided.

Keywords. Bryophytes, classification, Fissidens, Fissidentaceae, moss.

The most widely used classification of the Fis-
sidentaceae was first published by Brotherus
(1909), modeled essentially on an earlier system by
Müller (1901). Later, Brotherus (1924) revised and
amplified this system. The artificiality of several as-
pects of this classification, based primarily on ga-
metophytic characters, has been recognized for
some time by a number of workers (Crum & Steere
1957; Florschütz 1964; Bruggeman-Nannenga
1974; Bruggeman-Nannenga & Berendsen 1990;
Iwatsuki & Suzuki 1982; Iwatsuki 1985; Pursell
1988). For example, Bruggeman-Nannenga (1974)
pointed out that pluristratose laminal cells occur in
different infrageneric taxa of Fissidens, although
the subgenus Pachyfissidens is defined on this char-
acter. Sections Bryoidium [5 section Fissidens] and
Pachylomidium are distinguished by the thickness
of the limbidium and habitat, the former section
having a limbidium of unistratose cells in primarily
terrestrial species and the latter section having a
limbidium of pluristratose cells in aquatic species.
However, a limbidium of uni- and pluristratose cells
is often found in the same species (Demaret 1959;
Pursell & Allen 1996). Grout (1943) and Norkett
(Gangulee 1971) placed species that otherwise con-
form to section Fissidens in section Semilimbidium
because the limbidium is more or less restricted to
the vaginant laminae or absent (Bruggeman-Nan-
nenga 1978; Pursell 1997; Pursell & Allen 1996).
Brotherus considered the absence of a central
strand diagnostic of subgenera Octodiceras and Pa-
chyfissidens, and section Polypodiopsis, but species
throughout the family lack this structure.
New taxonomically useful characters have been
introduced in the past few years, leading to pro-
found changes in the understanding of relationships
in the Fissidentaceae. These characters are dis-
cussed in the following paragraphs.
Peristome. Hedwig (1801) based the genus Fis-
sidens in part on its single ring of sixteen, broad,
divided peristome teeth. Later, Bridel (1806) used
the peristome to segregate Octodiceras, a genus
based on the misinterpretation of a illustration of
F. semicompletus by Hedwig (1791–1792). Müller

0007-2745/04/$2.15/0
2 THE BRYOLOGIST
(1886), however, was the first to segregate a genus,
Moenkemeyera, on the basis of undivided peri-
stome teeth, and Brotherus (1909, 1924) and
Fleischer (1900–1902) were the first to note vari-
ations in the filaments of divided teeth, describing
them as ‘spirally-thickened’ (‘Peristomschenkel
spiralig verdickt’ and ‘Schenkel der Peristomzähne
spiralig verdickt, glatt oder papillös’) and ‘nodose’
[‘Peristomschenkel dicht knotig verdickt,’ ‘Peris-
tomschenkel selten schwach knotig verdickt’ and
‘Schenkel der Peristomzähne ringförmig (knotig)
verdickt’]. An SEM study by Mueller (1973) re-
vealed the structure of the ‘spiral’ filaments. Iwat-
suki and Suzuki (1982) described these differences
as ‘filaments with spiral thickenings’ and ‘articu-
lated filaments,’ and considered them important in
classification. Later, Ishihara and Iwatsuki (1992)
noted that ‘spirally thickened filaments’ and ‘cris-
tately thickened filaments’ are strongly hygroscopic
whereas ‘nodose filaments’ are not hygroscopic,
further enhancing the taxonomic value of these
structures.
The SEM studies of the Fissidens peristome be-
gun by Allen (1980) and continued by Bruggeman-
Nannenga and Berendsen (1990) firmly established
the usefulness of variations in the peristome in es-
tablishing natural relationships in the Fissidenta-
ceae. The most important of these variations are the
bryoides-, scariosus-, similiretis-, taxifolius-, and
zippelianus-types.
The bryoides-, zippelianus-, and taxifolius-types
of peristome are characterized by trabeculae on the
outer (dorsal) surface of the undivided proximal
(basal) parts that are clearly distinct from and usu-
ally higher than the lamellae (Figs. 1–2, 10–11, 22).
The difference in height can be slight as in the tax-
ifolius-type, or considerable as in the bryoides- and
zippelianus-types. In the scariosus- and similiretis-
types, the trabeculae are indistinct or only slightly
distinct from the equally high to higher lamellae
(Figs. 6–7, 17–18).
Additional differences are found in the changes
that occur in the area of the bifurcation. In the tax-
ifolius-type the ornamentation of the dorsal (outer)
surface of the undivided parts changes gradually
(Fig. 24), whereas in the other types this change is
sudden (Figs. 3, 8, 12, 19). Furthermore, the types
differ in the median and distal ends of the filaments.
Trabeculae are prominent, although not always pro-
truding (‘nodose’) throughout the taxifolius-type
(Fig. 25) and in the proximal and median parts of
the filaments in the zippelianus-type (Figs. 13–16).
In the median and distal parts of the filaments of
the similiretis-, bryoides-, and scariosus-types tra-
beculae are inconspicuous and/or indistinguishable
from the lamellar ornamentation (Fig. 20). The dis-
tal ends of the filaments of the zippelianus-, scario-
[VOL. 107
sus-, and bryoides-types are ‘spiral’ (Figs. 4–5),
those of the similiretis-type are squamose-like (Fig.
21), but the distal ends of the filaments of the tax-
ifolius-type have protruding trabeculae as well as a
spiral or vertical lamellar ornamentation (Figs. 25–
26). Infrequently, the distal ends of the filaments
can be papillose. Fimbriae are often found on the
inner (ventral) trabeculae of the scariosus-type
(Fig. 9).
There are species of Fissidens, however, in
which the peristome differs from the types de-
scribed above. These anomalous peristomes, de-
scribed below, are found in Pachyfissidens sections
Amblyothallia and Pachyfissidens, Fissidens section
Fissidens, and Aloma. Anomalous peristomes are
fairly common in corticolous species. Moreover,
peristomes of aquatic species are frequently re-
duced.
Costa. The structure of the costa in Fissidens
has been noted and figured in a number of publi-
cations (Bruggeman-Nannenga 1974; Iwatsuki &
Suzuki 1982; Pursell 1966, 1987, 1988, 1989; Pur-
sell & Reese 1985; Pursell et. al. 1988; Stone
1987). Kawai (1968), however, made the first sys-
tematic study of the costa throughout the mosses,
and six types, designated A through F, were rec-
ognized in 109 species belonging to 33 families.
Five species of Fissidens were studied and all had
a type E costa in which there are both adaxial and
abaxial sclereid bands. However, if the vaginant
laminae represent the ‘true’ leaf, these species pos-
sess a type D costa, characterized by only an ab-
axial band of sclereids. Subsequent to Kawai’s
work the next systematic studies on the Fissidens
costa were made by Bruggeman-Nannenga (1990)
and Stone (1990). Both workers found that the var-
iations observed correlated well with peristome
types. Although Stone did not designate any types,
Bruggeman-Nannenga recognized three types: the
bryoides-type (Figs. 36–38), characterized in the re-
gion of the vaginant laminae by two peripheral
guide cells and two lateral stereid bands separated
by one (sometimes more) large central cell; the tax-
ifolius-type (Figs. 31–35), characterized in the re-
gion of the vaginant laminae by four or more pe-
ripheral guide cells and two lateral stereid bands
separated usually by 1–5 large central cells; and,
the oblongifolius-type (Figs. 27–30), characterized
in the region of the vaginant laminae by three bands
of stereid cells (two lateral and one ventral) and as
many as 16 guide cells, arranged in the form of a
U or V, and as many as five large central cells. In
some species the lateral stereid bands may not be
separated (Fig. 30). The species studied by Kawai
have either the bryoides- or taxifolius-type of costa.
These two types are very close, the former probably
derived from the latter, the only distinction between
2004] PURSELL & BRUGGEMAN-NANNENGA: FISSIDENS 3

FIGURES 1–26. Sketches of peristome types in Fissidens. — 1–5. Bryoides-type. — 1. Lateral view of base of tooth.
— 2. Dorsal view of base of tooth. — 3. Dorsal view at bifurcation. — 4. Lateral view of median part of filament. —
5. Upper end of filament. — 6–9. Scariosus-type. — 6. Lateral view of base of tooth. — 7. Dorsal view of base of
tooth. — 8. Dorsal view of bifurcation. — 9. Ventral view of ventral fimbriate trabeculae. — 10–16.Zippelianus-type.
— 10. Lateral view of base of tooth. — 11. Dorsal view of base of tooth. — 12. Dorsal view of bifurcation. — 13.
Dorsal view of filament above bifurcation. — 14. Lateral view of filament above bifurcation. — 15. Dorsal view of
median part of filament. — 16. Upper end of filament. — 17–21. Similiretis-type. — 17. Lateral view of base of tooth.
— 18. Dorsal view of base of tooth. — 19. Dorsal view of bifurcation. — 20. Dorsal view of median part of filament.
— 21. Upper end of filament. — 22–26. Taxifolius-type. — 22. Lateral view of base of tooth. — 23. Dorsal view of
base of tooth. — 24. Dorsal view of bifurcation. — 25. Dorsal view of median part of filament. — 26. Upper end of
filament. [All figures, except no. 12, were published previously in Bruggeman-Nannenga & Berendsen, 1990, pp. 198–
199.]
4 THE BRYOLOGIST [VOL. 107

FIGURES 27–40. Costa types in Fissidens. — 27–30. Oblongifolius-type of costa. — 27, 29–30. T.s. (5 Transverse
section) through lower part of leaf. — 28. T.s. of upper part of leaf. — 31–35. Taxifolius-type of costa. — 31–34. T.s.
through lower part of leaf. — 35. T.s. through upper part of leaf. — 36–38. Bryoides-type of costa. — 36–37. T.s.
through lower part of leaf. — 38. T.s. through upper part of leaf. — 39–40. Anomalous costa of F. schusteri. [All
figures, except 39 and 40, appeared in Bruggeman-Nannenga, 1990. Trop. Bryol. 2: 38. figs. 27–28 from F. fasciculatus,
Esterhuysen 25464, U; fig. 29 from F. oblongifolius, De Sloover 20992, U; fig. 30 from F. maschalanthus, Crosby
12041, PAC; fig. 31 from F. dubius, Nannenga-Bremekamp 5, hb. Bruggeman-Nannenga; fig. 32 from F. polypodioides,
Long 10577, E; fig. 33 from F. taxifolius, Bruggeman-Nannenga s.n., hb. Bruggeman-Nannenga; figs. 34–35 from F.
zippelianus, Long 7746, E; 36 from F. rigidulus, Sainsbury s.n., BM; fig. 37 from F. pellucidus var. pellucidus, Florschütz
2057, U; fig. 38 from F. diversifolius, Sedgwick s.n., L; figs. 39–40 from Chen 5156, holotype of F. schusteri, HIRO.]
Upper scale bar 5 50 mm (figs. 39–40); lower scale bar 5 60 mm (figs. 27–30, 34, 36, 38) 5 50 mm (figs. 31–33,
37).
2004] PURSELL & BRUGGEMAN-NANNENGA: FISSIDENS
the two being the number of large cells in the me-
dian and distal parts of the vaginant laminae. As a
note of caution, when determining the type of costa,
for best results one should neither section too close
to the leaf insertion nor too close to the leaf apex.
Also, it is important that perichaetial leaves not be
examined to determine the costa type (see below
under subgenus Fissidens section Fissidens and
subgenus Aloma). Anomalous costae are known in
subgenus Fissidens section Fissidens and subgenus
Aloma.
Number of files of exothecial cells. Bruggeman-
Nannenga (1990) noted that the number of files of
exothecial cells in the periphery of the theca cor-
relate well with the type of peristome and the type
of costa. Thecae with more than 40 files of exothe-
cial cells are characteristic of those taxa with a
bryoides-type peristome/bryoides-type costa, a zip-
pelianus-type peristome/taxifolius-type costa, a tax-
ifolius-type peristome/taxifolius-type costa, a simi-
liretis-type peristome/oblongifolius-type costa, and
a taxifolius-type peristome/oblongifolius-type costa.
Thecae with about 32 files, but generally fewer than
40 files, of exothecial cells are characteristic of taxa
with a scariosus-type peristome/bryoides-type cos-
ta.
In addition to these taxonomically useful char-
acters, there are other characters that at this time
do not appear to be useful in the classification of
the subgenera and sections of Fissidens.
Spores. Boros et al. (1993) and Yu et al. (2002)
in LM and SEM studies of the surface features of
spores of eleven species and one variety of Fissi-
dens found only minor differences.
Protonemata. Fiala (1968) studied protonemata
of nine species of Fissidens. Although her results
are interesting, they are difficult to interpret. Per-
sistent protonemata, as detailed below, have been
reported in a few species.
Chromosome number. Iwatsuki (1985), the first
to use chromosome numbers as a character in the
classification of Fissidens, proposed a number of
changes aimed primarily at the subgeneric level. He
suggested that dioicous species with n 5 12 (sect.
Serridium 5 subgenus Pachyfissidens section Pa-
chyfissidens, below) are primitive and that auto-
icous species (subgenera Aneuron and Fissidens 5
subgenus Aloma and subgenus Fissidens section
Fissidens, respectively, below) are derived. How-
ever, F. oblongifolius Hook. f. & Wilson (section
Amblyothallia 5 subgenus Pachyfissidens section
Amblyothallia, below) is autoicous with n 5 12,
while F. pallidus Hook. f. & Wilson and F. asplen-
ioides Hedw. (also section Amblyothallia) are di-
oicous with n 5 12.
Molecular studies. Few molecular data are cur-
rently available. None can be used to support or
5
reject any of the infrageneric alignments presented
by previous workers or those presented herein.
However, the few molecular data that are available
support earlier conclusions of the placement of the
family as determined from morphological data.
Goffinet and Cox (2000), on the basis of data from
F. subbasilaris Hedw., place the genus in the Hap-
lolepideae, while La Farge et al. (2000) place the
family in the Dicranales, based on data from F.
dubius P. Beauv., F. philonotulus Besch., and F.
subbasilaris.
CLASSIFICATION OF THE FISSIDENTACEAE
Ideally, a classification should be based on a
minimum of two unambiguous differentiating char-
acters. Unfortunately, this ideal was not realized in
any of the previous schemes of classification of the
Fissidentaceae. To be sure, Fissidens is one of the
easiest of moss genera to recognize, but arranging
the species in infrageneric taxa was a challenge to
us just as it must have been to previous workers.
Although general groupings can be determined
there are exceptions, as enumerated below.
Fissidens, the only genus recognized by us in the
family, is one of the most successful genera of
mosses. According to the latest survey (Crosby et
al. 2000), there are somewhat fewer than 450 rec-
ognized species, an amazingly large number since
the majority of these species are based on variations
found in the basic leaf structure.
The classification of the infrageneric taxa of Fis-
sidens presented herein is basically a refinement of
that proposed by Brotherus (1924) in which many
of the characters he used, particularly the distribu-
tion and extension of the limbidium and cell wall
ornamentation, are retained. However, because of
the great degree of intergradation among the sub-
genera and particularly the sections as established
by Müller and followed by Brotherus, we have
adopted a broad view of the infrageneric taxa of
Fissidens. The changes proposed are based primar-
ily on variations in the peristome, the structure of
the costa, and the number of files of exothecial
cells. Other gametophytic and sporophytic varia-
tions are useful secondarily. An overview of this
classification is given in Figure 41.
As recognized herein, subgenus Pachyfissidens
section Amblyothallia includes those species that
are considered to be the primitive species of the
genus since F. asplenioides, F. fasciculatus
Hornsch., F. maschalanthus Mont., and F. plumo-
sus Hornsch. have peristomes that are a form of the
taxifolius-type that is similar to peristomes found in
some species of Dicranaceae (Bruggeman-Nannen-
ga & Berendsen 1990). Moreover, these four spe-
cies, along with all other species in the section,
6 THE BRYOLOGIST [VOL. 107

FIGURE 41. Diagnostic characters for the subgenera and sections. Other characters or combinations of characters
that are diagnostic for a subgenus or section: Aquatic, weak, floating plants with long and narrow leaves are exclusively
found in subgenus Octodiceras. Aquatic species with pleurocarpous appearance are found only in subgenus Fissidens
section Sarawakia. Leaf cells longer than 30 mm, guttulate leaf cells and ecostate leaves are exclusively found in
subgenus Aloma, whereas papillose leaf cells in combination with partly or completely limbate leaves do not completely
exclude other possibilities, but are nevertheless a strong indication that a species belongs in subgenus Aloma.

have an oblongifolius-type costa. Therefore, the
combination of these two characters is assumed to
be the primitive condition.
The ecostate or nearly ecostate species in sub-
genus Aloma, with large, mostly thin-walled lami-
nal cells and a scariosus-type peristome, and the
species of subgenus Octodiceras that have a strong
adaptation to an aquatic habitat, are herein consid-
ered to be the most advanced species of the genus.
Other derived species are those of subgenus Fissi-
dens section Fissidens that have lost or nearly lost
the dorsal and ventral laminae (i.e., the segregate
Nanobryum).
Of the four genera of the Fissidentaceae recog-
2004] PURSELL & BRUGGEMAN-NANNENGA: FISSIDENS
nized by Brotherus only Fissidens is maintained in
the present classification (Bruggeman-Nannenga
1997; Bruggeman-Nannenga & Berendsen 1990;
Pursell 1982; Pursell & Reese 1980b). Species of
Moenkemeyera Müll. Hal. are subsumed under sub-
genera Aloma, Fissidens section Fissidens, and Pa-
chyfissidens section Amblyothallia. Fissidentella
Cardot is subsumed under subgenus Aloma. Sim-
plicidens Herzog is included in section Fissidens of
subgenus Fissidens. A fifth genus, Nanobryum Dix-
on, which Brotherus (1925) placed in the Campy-
lopodioideae of the Dicranaceae between the gen-
era Dicranella (Müll. Hal.) Schimp. and Microcam-
pylopus (Müll. Hal.) M. Fleisch., is also subsumed
under section Fissidens of subgenus Fissidens.
Sainsburia Dixon, a sixth genus, unknown to
Brotherus, is subsumed under Fissidens section Fis-
sidens.
Brotherus divided the genus Fissidens into four
subgenera, Polypodiopsis, Eufissidens (5 Fissi-
dens), Pachyfissidens, and Octodiceras. All of these
are maintained as subgenera with the exception of
Polypodiopsis that is subsumed under subgenus
Aloma.
Brotherus recognized 12 sections in subgenus
Eufissidens (5 Fissidens). Three of these, Ambly-
othallia, Crispidium, and Serridium (this last one is
subsumed under section Pachyfissidens), are main-
tained in subgenus Pachyfissidens. Sections Aloma,
Crenularia, Semilimbidium, Pycnothallia, Reticu-
laria, and Weberiopsis are subsumed under subge-
nus Aloma, whereas sections Bryoidium, Pachylom-
idium, and Heterocaulon are united in section Fis-
sidens of subgenus Fissidens.
Subgenus Aloma and section Fissidens of sub-
genus Fissidens contain the grestest number of spe-
cies. These two taxa are followed by section Pa-
chyfissidens of subgenus Pachyfissidens.
FISSIDENTACEAE Schimp., Coroll. Bryol. Eur. 96.
1856. TYPE SPECIES: Fissidens bryoides Hedw.
Skitophylleae Mitt., J. Linn. Soc., Bot. 12: 11, 23. 1869.
LECTOTYPE SPECIES (here designated): Fissidens semi-
completus Hedw.
Schistophyllaceae Lindb., Utkast Eur. Bladmoss. 16. 1878,
nom. inval. incl. fam. prior.
Archifissidentaceae Dixon & P. de la Varde in P. de la
Varde, Arch. Bot. Mém. 1 (3): 23. 1927, nom. inval.
TYPE SPECIES: Nanobryum dummeri Dixon
Nanobryaceae W. Schultze-Motel, Willdenowia 5: 386.
1969.
Stem, except for initial stages, growing from 2-
sided apical cell. Leaves distichous, equitant, each
consisting of two vaginant laminae (lamina vera)
that clasp stem, a ventral lamina, located above va-
ginant laminae, and a dorsal lamina that occupies
entire length of leaf opposite vaginant and ventral
laminae, infrequently dorsal and ventral laminae
7
lacking or nearly so; costa single, usually well de-
veloped, sometimes reduced, or lacking. Peristome
single, haplolepidous, endostomate, rarely absent,
consisting of 16 teeth, usually divided 1/2–2/3 their
length, infrequently undivided or irregularly divid-
ed.
FISSIDENS Hedw., Sp. Musc. Frond. 152. 1801.
LECTOTYPE SPECIES (designated by E. Britton in
Britton, Fl. Bermuda 435. 1918): Fissidens bryoi-
des Hedw. The name Fissidens is derived from
Latin roots (fissum 5 cleft 1 dens 5 tooth), in
reference to the peristome teeth that are divided
for much of their length.
Skitophyllum Bach. Pyl., J. Bot. Agric. 4: 133. 1814, nom.
illeg. incl. gen. prior.
Schistophyllum Brid., Bryol. Univ. 2: 679. 1827, nom.
nud. in synon.
Conomitrium Mont., Ann. Sci. Nat., Bot., sér. 2, 8: 245.
1837, nom. illeg. incl. gen. prior.
Schistophyllum Lindb., Utkast Eur. Bladmoss. 16. 1878,
nom. illeg. incl. gen. prior.
Orthodens A. Jaeger, Ber. Thätigk. St. Gallischen Natur-
wiss. Ges. 1877–78: 394. 1879, nom. nud. pro Con-
omitrium.
Plants scattered to forming dense mats, terrestri-
al, epilithic, corticolous, epixylic, or aquatic, from
less than one millimeter to several centimeters in
length, pale to dark green, sometimes brownish to
blackish, erect but most often becoming decum-
bent, unbranched or branched, branches most fre-
quently with basal rhizoids; persistent protonemata
infrequent. Stem with proximal and axillary,
smooth, infrequently papillose, tan to reddish rhi-
zoids; axillary hyaline nodules differentiated or not;
central strand differentiated or not; axillary hairs
uniseriate, filiform; axillary or epiphyllous, stalked,
multicellular, filiform or clavate gemmae infre-
quent; multicellular, irregularly globose, subterra-
nean, rhizoidal gemmae infrequent. Leaves disti-
chous, equitant, pinnately arranged on elongate
stems or frondiform on short stems, infrequently
caducous; perichaetial leaves typically differentiat-
ed from cauline leaves, often longest; cauline
leaves of non-fertile stems often smaller than those
on perichaetial stems; proximal cauline leaves
smaller, often lacking dorsal and/or ventral laminae;
costa usually well developed, ending well below
leaf apex to excurrent, sometimes reduced, infre-
quently represented only by proximal vestige; mar-
gin entire to serrate or dentate, marginal cells dif-
fering little or not at all from inner laminal cells,
infrequently conspicuously smaller, or often differ-
entiated into limbidium to varying degrees on all
laminae or only on vaginant laminae, limbidium
consisting of uni- to pluristratose, often pigmented,
prosenchymatous, stereid cells; laminal cells mostly
unistratose, sometimes regularly to irregularly bi-
8 THE BRYOLOGIST
to pluristratose, small, quadrate, rounded to irreg-
ularly hexagonal, firm-walled, changing little upon
drying, to large, oblong to hexagonal or fusiform,
often thin-walled, usually collapsed when dry, outer
walls smooth, flat to bulging, mammillose (conic),
or unipapillose (sharply pointed), or pluripapillose,
infrequently prorate, sometimes lenticularly (con-
vexly) thickened, eguttulate or guttulate. Monoi-
cous (rhizautoicous, gonioautoicous, cladautoicous,
synoicous, polyoicous) or infrequently dioicous;
perigonia terminal on stems and branches as long
or nearly as long as perichaetial stems, or gemmi-
form and axillary or basal to larger stems, dwarf
male plants infrequently produced on leaves; peri-
chaetia usually terminal on short or elongate stems
but also on shorter or longer axillary branches; an-
theridia and archegonia sometimes naked in leaf
axils. Sporophytes usually 1(–2) per perichaetium,
infrequently of greater number, yellow-green when
young, darkening with age, or reddish, sometimes
caducous in aquatic species; seta short to elongate,
smooth or infrequently papillose; theca mostly ex-
serted, infrequently immersed, erect, radially sym-
metric, to inclined, 6 arcuate, bilaterally symmet-
ric, stomatose proximally or infrequently estoma-
tose, stomata phaneroporous, round-pored, each
surrounded by two lunate guard cells, exannulate,
but differentiated abscission zone mostly present,
exothecial cells quadrate to oblong to hexagonal,
vertical walls usually thicker than horizontal walls,
often collenchymatous; operculum conic-rostrate,
rostrum short or as long as theca, straight or curved.
Spores finely papillose to smooth. Calyptra cucul-
late or mitrate, smooth or prorate, covering the
operculum or only the rostrum. n 5 4, 5, 6, 8, 9,
10, 12, 13, 14, 15, 16, 19, 21, 24 (Fritsch 1991).
FISSIDENS subgenus PACHYFISSIDENS
Most species of subgenus Pachyfissidens can be
determined by the characters outlined in Figure 41.
Without sectioning the leaves, however, infertile
specimens might be confused with elimbate species
of subgenus Aloma that differ in the bryoides-type
costa, the usually smaller size, and frequently gut-
tulate laminal cells. Whereas subgenera Octodicer-
as, Fissidens, and Aloma have either smooth or pa-
pillose laminal cell walls, these structures in sub-
genus Pachyfissidens are variable, especially in sec-
tions Pachyfissidens and Crispidium, which seems
to indicate that in subgenus Pachyfissidens papil-
losity has developed more than once.
The subgenus is divided into three sections.
FISSIDENS subgenus PACHYFISSIDENS section AM-
BLYOTHALLIA (Müll. Hal.) comb. nov.
Fissidens section Amblyothallia Müll. Hal., Gen. Musc.
Frond. 63. 1901 [1900]. LECTOTYPE SPECIES (designated
[VOL. 107
by Bruggeman-Nannenga et al. 1994): Fissidens as-
plenioides Hedw.
Fissidens section Orthothallia Müll. Hal., Gen. Musc.
Frond. 66. 1901 [1900]. LECTOTYPE SPECIES (here des-
ignated): Fissidens petrophilus Sull.
Fissidens subgenus Serridium (Müll. Hal.) Z. Iwats. in Z.
Iwats. & S. Inoue section Amblyothallia emend. Brugg.-
Nann., Pursell & Z. Iwats., J. Hattori Bot. Lab. 77: 257.
1994.
Plants small to large, often robust, light to dark
green, unbranched to branched; branches with basal
rhizoids. Stem with basal and axillary, smooth, red-
dish rhizoids; subterranean, multicellular, irregular-
ly globose, rhizoidal gemmae infrequent; axillary
hyaline nodules usually poorly developed when
present; epidermis and outer 1–3 tiers of cortical
cells small, thick-walled, often pigmented; inner
cortical cells larger, hyaline; central strand differ-
entiated or not. Leaves imbricate, pinnately ar-
ranged, mostly lanceolate, infrequently caducous,
sometimes with darker or lighter margins, elimbate
or, rarely with weak limbidium on proximal parts
of vaginant laminae; costa mostly ending below
apex, infrequently percurrent or excurrent, oblon-
gifolius-type; dorsal lamina variable, reaching in-
sertion to ending well above insertion; vaginant
laminae mostly slightly unequal, minor lamina
mostly acute, infrequently rounded and ending on
or near costa; laminal cells mostly unistratose, in-
frequently irregularly bistratose to regularly bi- or
tristratose, smooth, often lenticularly thickened and
bulging or flat in dorsal and ventral laminae, mostly
evenly thickened and flat in vaginant laminae, egut-
tulate. Dioicous or monoicous. perigonia terminal
or axillary; perichaetia terminal on main stems or
branches. Sporophytes 1(–2) per perichaetium, red-
dish or yellow-green to yellow, darkening with age;
seta elongate; theca inclined, arcuate, bilaterally
symmetric or erect, radially symmetric, stomatose,
exothecial cells usually oblong, vertical walls usu-
ally thicker than horizontal walls; peristome simi-
liretis- or, infrequently, taxifolius-type, rarely
anomalous; operculum conic, long-rostrate. Spores
finely papillose to smooth. Calyptra smooth, cu-
cullate. n 5 12, 14, 16 (Fritsch 1991).
Bruggeman-Nannenga et al. (1994) recognized
19 species in this section. However, after careful
consideration, F. grandifrons is now thought to be
best placed in section Pachyfissidens, and F. for-
mosanus Nog. has been subsumed under F. oblon-
gifolius (Beever & Stone 1998). In addition, a spe-
cies recently discovered in New Caledonia by
Frank Müller of Dresden Technological University
(Müller et al. 2003) is placed here. The majority of
the species have restricted distributions—only F.
asplenioides and F. oblongifolius are widespread.
All species are easily recognized by the oblongi-
folius-type costa.
2004] PURSELL & BRUGGEMAN-NANNENGA: FISSIDENS
Subterranean, multicellular, irregularly globose,
rhizoidal gemmae have been reported in F. asplen-
ioides (Arts 1989) and F. oblongifolius (Imura &
Iwatsuki 1988). Six species have small and incon-
spicuous axillary hyaline nodules (Iwatsuki & Pur-
sell 1980): F. asplenioides (the nodules can at times
be large), F. fasciculatus, F. flabellatus Hornsch.,
F. maschalanthus, F. oblongifolius, and F. steno-
phyllus Ångstr. Fissidens pallidus, however, has
conspicuous axillary hyaline nodules. Caducous
leaves are present in F. radicans Mont. (Brugge-
man-Nannenga & Berendsen 1987; Pursell
1994a,b), and, although not demonstrated, these
structures most likely are involved in vegetative re-
production. The minor laminae in F. asplenioides
are mostly rounded and free, a feature also seen in
F. maschalanthus but to a lesser degree, and in per-
ichaetial leaves of species belonging to other sub-
genera. A weak limbidium of linear, oblong cells,
restricted to the proximal parts of the vaginant lam-
inae, can be found in F. asplenioides, F. fascicu-
latus, F. maschalanthus, F. plumosus, and F. sten-
ophyllus.
Laminal cell walls in this section are either con-
vexly thickened or flat and differ little from the cell
walls seen in the other two sections of the subge-
nus. Flat and evenly thickened walls throughout all
laminae are found in F. fasciculatus, F. pallidus, F.
petrophilus Sull., F. stenophyllus, and F. cagoui
(Müller et al. 2003). In most species, the laminal
cells are unistratose. However, in F. asplenioides
and F. maschalanthus these cells can be bistratose
along the costa. In F. fasciculatus, F. stenophyllus,
the freakish F. asplenioides var. aegrotus (Bizot)
Brugg.-Nann., Pursell & Z. Iwats., and other aquat-
ic expressions of F. asplenioides, all laminal cells
can be 2- to 3-stratose. In the leaves of F. sufflatus
I. G. Stone and F. pseudopallidus I. G. Stone there
are one or more thicker and bulging marginal rows
of cells, forming bands darker than the inner lam-
inal cells.
Thecae in the F. radicans-complex (F. dendro-
philus Brugg.-Nann. & Pursell, F. radicans, F. mi-
crocarpus Müll. Hal., and F. santa-clarensis
Thér.—Bruggeman-Nannenga & Pursell 1990) are
erect and radially symmetric while in the remaining
species of the section the thecae are usually some-
what curved and bilaterally symmetric. The simi-
liretis-type peristome is unique to this section and
even when reduced or broken is easily recognized
by the ‘beady’ appearance of the dorsal (outer) sur-
face of the undivided parts of the teeth. However,
four species, F. asplenioides, F. fasciculatus, F.
maschalanthus, and F. plumosus, are characterized
by the taxifolius-type peristome. In F. radicans and
F. microcarpus, the peristome teeth are undivided
or imperfectly divided and the number of files of
9
exothecial cells in these two species is usually less
than 40.
FISSIDENS subgenus PACHYFISSIDENS section CRIS-
PIDIUM (Müll. Hal.) comb. nov.
Fissidens section Crispidium Müll. Hal., Gen. Musc.
Frond. 64. 1901 [1900]. LECTOTYPE SPECIES (here des-
ignated): Fissidens zippelianus Dozy & Molk.
Plants small to medium, often robust, light to
dark green, unbranched or branched; branches with
basal rhizoids. Stem with basal and axillary reddish
rhizoids; subterranean, irregularly globose, multi-
cellular, rhizoidal gemmae not known; axillary hy-
aline nodules large, protruding; epidermis and outer
1–2 tiers of cortical cells small, thick-walled, often
pigmented; inner cortical cells larger, thin-walled,
hyaline; central strand differentiated or not. Leaves
usually imbricate, pinnately arranged or often fron-
diform, linear-lanceolate to elliptic, elimbate on
dorsal and ventral laminae, rarely weakly limbate
on proximal parts of vaginant laminae of perichae-
tial leaves, infrequently with 1–3 rows of swollen
marginal cells; costa ending 2–18 cells below apex
or percurrent to long-excurrent, taxifolius-type; dor-
sal lamina usually reaching insertion, sometimes
slightly undulate proximally; vaginant laminae
acute, slightly unequal; laminal cells unistratose or
regularly to irregularly bistratose, firm-walled,
walls lenticularly thickened, mammillose, indis-
tinctly or distinctly pluripapillose. All species di-
oicous (?); perigonia unknown in most species, ter-
minal on long branches when present; perichaetia
terminal on main stems or long axillary branches.
Sporophytes 1(–2) per perichaetium, reddish; seta
elongate; theca exserted, inclined, arcuate, bilater-
ally symmetric, stomatose, exothecial cells oblong,
vertical walls thicker than horizontal walls; peri-
stome zippelianus-type; operculum conic, long-ros-
trate (where known). Spores finely papillose to
smooth. Calyptra smooth, cucullate. n 5 6, 12
[based on the dioicous F. crispulus Brid. (as F. zip-
pelianus Dozy & Molk.), Fritsch 1991].
This is a small section consisting of 5–6 species
found in Africa, Malesia, Australasia, and Oceania
and characterized by a zippelianus-type peristome.
Another conspicuous feature is the highly devel-
oped axillary hyaline nodules. However, as a note
of caution, large axillary hyaline nodules are also
present in several species of section Pachyfissidens.
These axillary nodules also occur in other subgen-
era and sections where they usually are not as well
developed. Infertile specimens of section Crispi-
dium are most likely to be confused with those spe-
cies of section Pachyfissidens with hyaline axillary
nodules since both sections have elimbate leaves
and the same type of costa. In such cases, a peri-
10 THE BRYOLOGIST
stome is needed to establish the proper taxonomic
placement. Indeed, these two sections have strik-
ingly different peristomes, but these are not sup-
ported by consistent gametophytic differences.
Leaves of F. javanicus Dozy & Molk. have dark
marginal bands formed by 1–more rows of thicker-
walled cells that vary from 1–3 cells in thickness.
Leaves with bistratose laminal cells are found in F.
brevifrons Mitt. and F. planifrons. Pluripapillose
laminal cells are found in F. planifrons and F. su-
bangustus M. Fleisch. Laminal cells of F. brevif-
rons are smooth or indistinctly pluripapillose, while
F. crispulus and F. javanicus have mammillose
laminal cells.
Brotherus (1924) differentiated sections Serri-
dium (5 section Pachyfissidens) and Crispidium by
sporophytes on axillary branches in the former and
terminal sporophytes in the latter. However, at least
two species of section Serridium, F. osmundioides
Hedw. and F. gymnogynus Besch., have terminal
sporophytes whereas two species of section Cris-
pidium, F. planifrons Besch. and F. crispulus, have
both terminal and axillary (on long branches) per-
ichaetia.
FISSIDENS subgenus PACHYFISSIDENS section PA-
CHYFISSIDENS
Fissidens section Pachyfissidens Müll. Hal., Syn. Musc.
Frond. 1: 45. 1849 [1848]. TYPE SPECIES: Fissidens
grandifrons Brid.
Conomitrium Mont. section Sciarodium Müll. Hal., Syn.
Musc. Frond. 2: 526. 1851. LECTOTYPE SPECIES (here
designated): Conomitrium osmundioides (Hedw.) Müll.
Hal.
Pachyfissidens (Müll. Hal.) Limpr., Laubm. Deutschl. 1:
454. 1887.
Fissidens subgenus Aloma Kindb. section Adiantoidei
Kindb., Eur. N. Amer. Bryin. 2: 167. 1897. LECTOTYPE
SPECIES (here designated): Fissidens adianthoides
Hedw.
Fissidens subgenus Aloma section Osmundoidei Kindb.,
Eur. N. Amer. Bryin 2: 168. 1897. LECTOTYPE SPECIES
(here designated): Fissidens osmundioides Hedw.
Fissidens section Serridium Müll. Hal., Gen. Musc. Frond.
67. 1901 [1900]. LECTOTYPE SPECIES (designated by
Iwatsuki & Suzuki 1996): Fissidens taxifolius Hedw.
Conomitrium section Crispidiella Müll. Hal., Gen. Musc.
Frond. 75. 1901 [1900]. LECTOTYPE SPECIES (here des-
ignated): Conomitrium osmundioides (Hedw.) Müll.
Hal.
Fissidens subgenus Eufissidens section Laterales Grout,
Moss Fl. N. Amer. 1: 8. 1936. nom. illeg. incl. sect.
prior. LECTOTYPE SPECIES (here designated): Fissidens
taxifolius Hedw.
Fissidens section Marginatus Grout, THE BRYOLOGIST 44:
135. 1941. LECTOTYPE SPECIES (here designated): Fis-
sidens bourgaeanus Besch.
Fissidens subgenus Serridium (Müll. Hal.) Z. Iwats. in Z.
Iwats. & S. Inoue, J. Hattori Bot. Lab. 57. 354. 1984
and Iwatsuki & Suzuki, J. Hattori Bot. Lab. 79. 160.
1996.
Plants medium to large, light to dark green, usu-
[VOL. 107
ally branched; branches with basal rhizoids. Stem
with basal and axillary, reddish rhizoids, mostly
smooth, infrequently papillose; subterranean, irreg-
ularly globose, multicellular, rhizoidal gemmae
sometimes present; chlorophyllose, branched fila-
ments at bases of leaves rare; axillary hyaline nod-
ules well developed or not; epidermis and outer 1–
3 tiers of cortical cells small, thick-walled, often
pigmented; inner cells larger, thin-walled, hyaline;
central strand differentiated or infrequently not.
Leaves usually imbricate, pinnately arranged, ob-
long to lanceolate, often irregularly, coarsely and
distantly serrate distally, elimbate; costa ending
several cells below apex to short-excurrent, rarely
obscured by overlying chlorophyllose cells, taxifol-
ius-type; laminal cells unistratose, bistratose, or
rarely pluristratose, firm-walled, walls evenly thick-
ened, mostly somewhat bulging, infrequently mam-
millose or pluripapillose. Monoicous; perigonia
mostly gemmiform, basal, axillary, rarely terminal
on epiphyllous dwarf males, infrequently terminal
on longer stems; perichaetia terminal on short ax-
illary branches, rarely terminal on main stems. Spo-
rophytes usually one per perichaetium, reddish; seta
elongate; theca erect, radially symmetric or some-
what inclined, slightly arcuate, bilaterally symmet-
ric, mostly stomatose, exothecial cells mostly ob-
long, vertical walls thicker than horizontal walls;
peristome taxifolius-type; operculum conic, long-
rostrate. Spores finely papillose to smooth. Calyp-
tra cucullate, smooth. n 5 6, 8, 9, 12, 13, 16, 24
(Fritsch 1991).
This section consists of about 23 species found
in the Americas, Europe, Africa, Asia, Australasia,
and Oceania, often of limited distribution. It is well
represented in temperate regions. Fissidens adian-
thoides and F. taxifolius are widely distributed in
the Northern Hemisphere, but of limited distribu-
tion in the Southern Hemisphere.
Fissidens ovatus Brid. has large, protruding ax-
illary hyaline nodules and was therefore considered
to belong in section Crispidium (Iwatsuki & Pursell
1980, as F. glaucescens Hornsch.). However, its
taxifolius-type peristome places it in section Pachy-
fissidens.
Most reports of the subterranean, irregularly glo-
bose, multicellular, rhizoidal gemmae in Fissidens
have been found in species of section Pachyfssi-
dens: F. dubius (Arts 1987, as F. cristatus Wilson);
F. osmundioides (Arts 1989); F. polyphyllus Wilson
(Arts 1989); F. serrulatus Brid. (Arts 1989); F. tax-
ifolius (Bruggeman-Nannenga & Touw 1989; Cor-
rens 1899; Imura & Iwatsuki 1988; Landwehr
1984; Lorentz 1864; Whitehouse 1966), and F.
teysmannianus Dozy & Molk. (as F. adelphinus
Besch., Imura & Iwatsuki 1988). The clusters of
chlorophyllose, branched filaments produced near
2004] PURSELL & BRUGGEMAN-NANNENGA: FISSIDENS
leaf bases in F. aphelotaxifolius Pursell might func-
tion in vegetative reproduction (Pursell 1976). Fis-
sidens grandifrons and F. osmundioides have pa-
pillose rhizoids, rare in the genus.
The costa of F. grandifrons has a basic taxifol-
ius-type structure, although an occasional single file
of adaxial stereids occurs in some leaves, a feature
that led to placing the species in section Ambly-
othallia (Pursell & Allen 1994). A costa obscured
by overlying chlorophyllose cells is found in F.
gymnogynus Besch. and F. subbasilaris. According
to Iwatsuki and Suzuki (1982) stomata are lacking
in the sporophytes of F. grandifrons.
Fissidens bourgaeanus, F. luisieri, F. nobilis
Griff., and F. serrulatus have darker marginal lam-
inal bands. The cells of these marginal bands are
unistratose in F. bourgaeanus, F. luisieri P. de la
Varde, and F. serrulatus, but bistratose in F. no-
bilis. Lighter marginal bands are found in F. adian-
thoides, F. pacificus Ångstr., and sometimes F.
bourgaeanus and F. dubius. Although lamina cells
are usually bulging, there are several exceptions. In
F. nobilis, F. serrulatus, and F. taxifolius these
cells are mammillose (mainly in the vaginant lam-
inae in the last species), while in F. nothotaxifolius
Pursell & Hoe, F. teysmannianus (Japan, as F.
adelphinus) , and F. bushii (Cardot & Thér.) Cardot
& Thér. the cells in the vaginant laminae are indis-
tinctly pluripapillose in the corners. Moreover,
Iwatsuki and Suzuki (1982) reported that the cells
of the ventral laminae of F. dubius (as F. cristatus)
have thickened corners. Flat laminal cells are found
in F. obscurus Mitt. and in the vaginant laminae of
F. nobilis and F. luisieri. Laminal cells of F. no-
bilis, F. dubius, F. serrulatus, and F. geminiflorus
are irregularly bistratose. Fissidens grandifrons, F.
geijkesii Florsch., and F. perdecurrens are the only
species with pluristratose laminal cells. Epiphyllous
dwarf male plants are known in F. dubius (as F.
cristatus) and F. gymnogynus (Iwatsuki & Suzuki
1982).
The peristome of F. gymnogynus is not the typ-
ical taxifolius-type. Such teeth are reflexed when
dry and strongly inflexed when wet. The teeth of
F. gymnogynus, on the other hand, stand erect both
wet and dry. Iwatsuki and Suzuki described these
teeth with spiral thickenings, although the teeth are
rather strongly papillose and the thickenings rather
difficult to discern. Bruggeman-Nannenga and Ber-
endsen (1990) considered the peristome of this spe-
cies and that of F. anomalus to be reduced taxifol-
ius-type.
FISSIDENS subgenus OCTODICERAS (Brid.) Broth.,
Nat. Pflanzenfam. 1 (3): 361. 1909 [1901].
Octodiceras Brid., Muscol. Recent. Suppl. 1: 162. 1806.
TYPE SPECIES: Octodiceras fissidentoides Brid.
11
Fissidens section Hydrofissidens Müll. Hal., Syn. Musc.
Frond. 1: 44. 1849 [1848]. TYPE SPECIES (here desig-
nated): Fissidens julianus (DC) Schimp.
Conomitrium section Octodiceras (Brid.) Müll. Hal., Syn.
Musc. Frond. 2: 524. 1851, nom. illeg. prior. ut gen.
Osmundula Rabenh., Krypt.-Fl. Sachsen 1: 609. 1863.
TYPE SPECIES: Osmundula fissidentoides Rabenh.
Fissidens section Octodiceras (Brid.) Mitt., J. Linn. Soc.,
Bot. 12: 581. 1869.
Schistophyllum Lindb. subgenus Octodiceras (Brid.)
Lindb., Musci Scand. 13. 1879.
Plants aquatic, submerged but often periodically
emergent, delicate and feathery in appearance, to
12 cm or more long, yellow- to dark green at grow-
ing tips, mostly brown-black proximally, often en-
crusted with diatoms, usually much branched;
branches with basal rhizoids. Stem with smooth,
reddish axillary and basal rhizoids; axillary hyaline
nodules not differentiated; epidermis and 1–2 tiers
of outer cortical cells small, thick-walled, pigment-
ed; inner cortical cells larger, thin-walled, hyaline;
central strand not differentiated. Leaves usually dis-
tant, numerous, pinnately arranged, variable in size,
to ten times or more as long as wide, lanceolate to
linear-lanceolate, acute, frequently falcate; margin
6 entire, elimbate or weakly limbate on lower 1/3
of vaginant laminae, limbidial cells unistratose,
firm-walled, smooth; costa ending near apex or far
below apex, bryoides-type; dorsal lamina reaching
insertion or ending above; vaginant laminae acute,
6 equal, or unequal and minor lamina rounded or
nearly so, ending on or near costa; laminal cells
unistratose, or infrequently and irregularly bistra-
tose, firm-walled, smooth, flat or slightly bulging,
quadrate, short-oblong to hexagonal, eguttulate.
Monoicous; perigonia and perichaetia gemmiform
and axillary, or terminal on main stems and elon-
gate axillary branches. Sporophytes 1–more per
perichaetium, small, inconspicuous, sometimes ca-
ducous; seta short; theca usually exserted, infre-
quently immersed, erect, radially symmetric, esto-
matose; peristome undivided or divided, filaments
infrequently absent, reduced bryoides-type; oper-
culum conic, usually short-rostrate, infrequently
long-rostrate. Spores smooth, large for the genus.
Calyptra mitrate or cucullate, smooth. No chro-
mosome counts reported.
Subgenus Octodiceras is a group of species char-
acterized essentially by adaptations to its aquatic
habitat: stems long, weak, floating, often much
branched, central strand absent; leaves flaccid, lin-
ear-lanceolate; seta short, theca estomatose; and
peristome reduced. The subgenus shares the bryoi-
des-type costa with subgenera Fissidens and Aloma,
but is most closely related to subgenus Fissidens.
Both Octodiceras and section Sarawakia of sub-
genus Fissidens might be considered ‘aquatic
branches’ of section Fissidens subgenus Fissidens.
12 THE BRYOLOGIST
All three taxa share similar laminal cells and costae,
and, although most peristomes are reduced they can
be identified as the bryoides-type (illustrations in
Beever 1995; Pursell 1987; Pursell et al. 1988).
Sporophytes of F. berteroi (Beever 1995), F.
bessouensis Corb. (Pursell 1987), and F. fontanus
(Britton 1902; Melick 1985; Pursell 1987) are ca-
ducous. This small subgenus was revised by Pursell
(1987) and consists of six species distributed in
North America, South America, Europe, Africa,
and Australasia. There is no report of the occur-
rence of the subgenus in Asia.
FISSIDENS Hedw. subgenus FISSIDENS
Subgenus Fissidens is an assemblage of species
characterized by the bryoides-type costa and bryoi-
des-type peristome, although anomalous peristomes
are found in a few species. Typically, the leaves are
limbate on all laminae, less often only on the va-
ginant lamine of some or all leaves. Rarely are all
leaves elimbate. Laminal cells are smooth and flat
or bulging. The subgenus is divided into two close-
ly related sections.
FISSIDENS Hedw. subgenus FISSIDENS section FIS-
SIDENS LECTOTYPE SPECIES (designated by E. Brit-
ton in Britton, Fl. Bermuda 435. 1918): Fissidens
bryoides Hedw.
Heterodon Raf., Med. Repos., ser. 2, 5: 350. 1801. TYPE
SPECIES: Heterodon bryoides (Hedw.) Raf.
Fissidens section Eufissidens Müll. Hal., Syn. Musc.
Frond. 1: 50. 1849 [1848], nom. illeg. TYPE SPECIES
(here designated): Fissidens bryoides Hedw.
Fissidens section Bryoideae Mitt., J. Proc. Linn. Soc.,
Bot., Suppl. 1: 141. 1859, nom. nud.
Moenkemeyera Müll. Hal., Flora 69: 506. 1886. TYPE SPE-
CIES: Moenkemeyera mirabilis Müll. Hal.
Fissidens subgenus Aloma Kindb. section Obtusifolii
Kindb., Eur. N. Amer. Bryin. 2: 165. 1897. TYPE SPE-
CIES: Fissidens obtusifolius Wilson
Fissidens subgenus Eufissidens (Müll. Hal.) Kindb., Eur.
N. Amer. Bryin. 2: 166. 1897, nom. illeg.
Fissidens section Bryoidium Müll. Hal., Gen. Musc.
Frond. 54. 1901 [1900]. TYPE SPECIES: Fissidens bryoi-
des Hedw.
Fissidens section Heterocaulon Müll. Hal., Gen. Musc.
Frond. 54. 1901 [1900]. LECTOTYPE SPECIES (here des-
ignated): Fissidens geheebii Müll. Hal.
Fissidens section Pachylomidium Müll. Hal., Gen. Musc.
Frond. 60. 1901 [1900]. LECTOTYPE SPECIES (designated
by Bruggeman-Nannenga 1978): Fissidens rufulus
Bruch & Schimp. in Bruch, Schimp. & W. Gümbel
Conomitrium Mont. section Limbidium Müll. Hal., Gen.
Musc. Frond. 71. 1901 [1900]. LECTOTYPE SPECIES (des-
ignated by Bruggeman-Nannenga 1978): Conomitrium
smaragdinum Lorentz & Müll. Hal.
Conomitrium section Pycnothallidium Müll. Hal., Gen.
Musc. Frond. 75. 1901 [1900]. LECTOTYPE SPECIES (here
designated): Conomitrium lindigii Hampe
Simplicidens Herzog, Beih. Bot. Centrabl. 26: 58. 1909
[1910]. TYPE SPECIES: Simplicidens andicola Herzog
[VOL. 107
Nanobryum Dixon, J. Bot. 60: 101. 1922. TYPE SPECIES:
Nanobryum dummeri Dixon
Fissidens section Pycnothallia subsection Pycnophylli P.
de la Varde, Ann. Cryptog. Exot. 2: 277. 1929. LEC-
TOTYPE SPECIES (designated by Bizot & Pierrot 1964):
F. pycnophyllus Müll. Hal.
Fissidens subgenus Eufissidens section Terminales Grout
subsection Limbatus Grout, Moss Fl. N. Amer. 1: 8.
1936, nom. illeg. incl. sect. prior. LECTOTYPE SPECIES
(here designated): Fissidens limbatus Sull.
Fissidens subgenus Eufissidens section Terminales sub-
section Semilimbatus Grout, Moss Fl. N. Amer. 1: 8.
1936, nom. illeg. incl. sect. prior. LECTOTYPE SPECIES
(here designated): Fissidens obtusifolius Wilson
Sainsburia Dixon, THE BRYOLOGIST 44: 40. 1941. TYPE
SPECIES: Sainsburia novae-zealandiae Dixon
Fissidens section Limbatus (Grout) Grout, THE BRYOLO-
GIST 44: 131. 1941, nom. illeg. incl. sect. prior.
Fissidens section Semilimbatus (Grout) Grout, THE BRY-
OLOGIST 44: 132. 1941, nom. illeg. incl. sect. prior.
Fissidens section Semilimbidium subsection Sublucidi P.
de la Varde ex Bizot & R. B. Pierrot, Rev. Bryol. Lich-
énol. 33: 233. 1964. TYPE SPECIES: Fissidens valiae P.
de la Varde
Fissidens section Fissidens subsection Pachylomidium
(Müll. Hal.) Nork. in Gangulee, Mosses E. India 2: 454.
1971.
Fissidens section Semilimbidium Müll. Hal subsection
Bryolimbidium Nork. in Gangulee, Mosses E. India 2:
455. 1971. TYPE SPECIES: Fissidens bilaspurense Gan-
gulee
Plants small to large, light to sordid green, ter-
restrial, saxicolous, lignicolous, usually in humid
places, or aquatic, erect to decumbent, unbranched
or branched; branches with basal rhizoids; persis-
tent protonemata rare. Stem smooth, monomorphic,
more or less of equal size, or dimorphic, non-fertile
stems longer with a greater number of leaves, per-
ichaetial stems short with few leaves; axillary hy-
aline nodules differentiated or not; rhizoids basal
and axillary, tan to reddish; epidermis and outer 1–
4 tiers of cortical cells section small, thick-walled,
pigmented or not; inner cortical cells large, thin-
walled, hyaline; central strand differentiated or not;
subterranean and axillary, multicellular, irregularly
globose, rhizoidal gemmae rare; axillary, stalked
multicellular, clavate gemmae infrequent. Leaves
imbricate to distant, few to numerous pairs pin-
nately arranaged, variable in shape; margin entire
to serrulate, expecially at leaf apex; margin limbate,
limbidium on all laminae, wholly or in part, or, less
often, present only on vaginant laminae, infrequent-
ly confined to proximal parts of vaginant laminae
of perichaetial leaves, sometimes not differentiated,
limbidial cells uni- to pluristratose; costa variable
in length, bryoides-type; dorsal lamina ending well
above insertion to long-decurrent, mostly ending at
insertion; vaginant laminae acute, equal to slightly
unequal, minor lamina infrequently ending on cos-
ta; laminal cells usually quadrate to regularly hex-
agonal, rarely fusiform, unistratose to irregularly or
regularly bi- to tristratose, firm-walled, smooth, flat
2004] PURSELL & BRUGGEMAN-NANNENGA: FISSIDENS
or bulging, rarely unipapillose (one species), infre-
quently pluripapillose, eguttulate. Monoicous; po-
sition of antheridia variable, perichaetia terminal on
main stems or branches. Sporophytes 1–2 per per-
ichaetium; seta long; theca exserted, erect, radially
symmetric to 6 inclined, 6 arcuate, bilaterally
symmetric, stomatose or infrequently estomatose;
exothecial cells quadrate to oblong, vertical walls
often thicker that horizontal walls, often collenchy-
matous; peristome bryoides-type, sometimes re-
duced. Spores finely papillose to smooth. Calyptra
cucullate or mitrate, smooth.
The limited number of stable characters makes it
difficult to adequately circumscribe many species
in this section. For example, the widespread F.
bryoides is highly variable and taxonomically di-
verse. An examination of Index Muscorum (Wijk et
al. 1962) reveals there is no dearth of names that
have been applied to this complex. Considerable
variation is found in leaf size and shape, extent and
thickness of the limbidium, the number of strata of
laminal cells, size and shape of laminal cells, and
position of gametangia. As discussed by Brugge-
man-Nannenga (O’Shea et al. 2001), European
workers either treat expressions involved as species
(Ahrens 2000; Anderson et al. 1976; Bruggeman-
Nannenga & Nyholm 1986; Bruggeman-Nannenga
& Touw 1989; Casas et al. 2001; Cortini Pedrotti
2001; Nyholm 1954; Smith 1978) or as subspecies
(Frahm & Frey 1992; Pilous & Duda 1960; Sándor
& László 1983). Asian workers treat the different
expressions as varieties and/or forms (Chopra &
Kumar 1981; Iwatsuki & Suzuki 1982; Li & Iwat-
suki 2001) while in North America the expressions
have been given no taxonomic recognition (Crum
& Anderson 1981; Ireland 1971; Pursell 1994b) or
are treated as varieties (Pursell 1994a; Pursell &
Allen 1996), or species (Ireland 1982).
Persistent protonemata are found in F. gladiolus
Mitt. (Dixon 1922, as Nanobryum dummeri Dixon;
Potier de la Varde 1928, 1936, as N. dummeri), and
F. thorsbornei (I. G. Stone) Brugg.-Nann. Only one
species in this section, F. beckettii Mitt. has been
reported to have subterranean, multicellular, irreg-
ularly globose, rhizoidal gemmae (Arts 1998). Mul-
ticellular, axillary gemmae were reported by Gan-
gulee (1971) in F. bryoides.
Nanobryum, found in Africa, Australia, New Ca-
ledonia, and New Guinea, was reduced to the syn-
onymy of Fissidens (Pursell & Reese 1980b), al-
though it is recognized by some workers (Iwatsuki
& Suzuki 1989; Stone 1982). Leaves of this taxon
lack or nearly lack dorsal and ventral laminae, mak-
ing them similar to the lowermost leaves of many
other species. Laminal cells in this segregate are
linear-oblong or fusiform, similar to those seen in
some species of subgenus Aloma. However, the
13
bryoides-type costa and bryoides-type peristome
leave no doubt as to the taxonomic position of the
species.
Perichaetial leaves of F. jansenii Sérgio and Pur-
sell (2001, Fig. 2a) show an oblongifolius-type cos-
ta common to subgenus Pachyfissidens section Am-
blyothallia. The costa in F. schusteri Z. Iwats. & P.
C. Wu (Figs. 39–40; also Iwatsuki & Wu 1988, Fig.
j, and Li & Iwatsuki 2001, Plate 87, Fig. 10) ap-
proaches the bryoides-type, but because the en-
larged cells are bordered by stereid cells it is con-
sidered not to fit this type.
Most species of section Fissidens are limbate on
all laminae. Many species, however, e.g., F. bogo-
sicus Müll. Hal., F. bryoides s.l., F. crassipes
Bruch & Schimp. subsp. warnstorfii (M. Fleisch.)
Brugg.-Nann., F. rigidulus Hook. f. & Wilson var.
pseudostrictus Beever, and F. leucocinctus Hampe
have expressions in which the limbidium is limited
to the vaginant laminae. Indeed, F. taylorii Müll.
Hal., F. pygmaeus Hornsch., and F. diversifolius
are weakly limbate. In F. bryoides and F. bogosicus
there are elimbate expressions, and in F. amoenus
Müll. Hal. the leaves are elimbate except for a weak
limbidium on the lower margins of the vaginant
laminae.
Species with the limbidia restricted to vaginant
laminae, and with smooth, firm-walled laminal cells
are not restricted to subgenus Fissidens, but are also
found in some aquatic species of subgenus Aloma,
e.g., F. waiensis Beever (sporophyte unknown,
placed here provisionally), F. integerrimus Mitt., F.
sedgwickii Broth. & Dixon (in this species there are
more than 40 files of exothecial cells), and an Af-
rican species yet to be described by one of us (M.
A. B.-N.). Lacking peristomes (scariosus-type),
these species would doubtless be placed in subge-
nus Fissidens. It seems that partially limbate leaves
with smooth, firm-walled laminal cells are aquatic
adaptations found in several species that are not
closely related. In subgenus Fissidens this adapta-
tion has led to either a reduction of the limbidium
as in F. acacioides and expressions of F. leuco-
cinctus, or, to a thickening of the limbidium, as in
F. oediloma Broth, F. rigidulus, F. rufulus, F. ri-
vularis (Spruce) Schimp., and F. ventricosus Lesq.,
whereas in subgenus Aloma this aquatic adaptation
has led to a loss of papillae.
Although the laminal cells in this section are typ-
ically smooth, clear, and flat, there are species with
smooth, bulging laminal cells, and also species with
widely spaced, inconspicuous papillae on the lam-
inal cells (Potier de la Varde 1929b, 1931). The
laminal cells of F. megalotis Müll. Hal. (5 F. vit-
tatus Hook. f. & Wilson) vary from smooth to uni-
and pluripapillose, making this species difficult to
place. All these species, however, have a bryoides-
14 THE BRYOLOGIST
type peristome, bryoides-type costa, and more than
40 files of exothecial cells, which far outweigh the
presence of papillae found in some expressions.
Another example with a similar costa, peristome,
and exothecium is F. bogosicus in which the leaves
vary from completely limbate to elimbate, and the
large, clear laminal cells appear smooth but in
many specimens are minutely and distantly papil-
lose.
There are a few species in which the peristomes
are not the typical bryoides-type. Fissidens incisus
Herzog has a fragile peristome in which the teeth
are unequally divided, and are smooth proximally
but strongly spiculose distally. Fissidens excurren-
tinervis R. S. Williams also has a fragile peristome
in which the teeth are irregularly divided, smooth
proximally but spirally striate to spirally thickened
distally. The teeth in this latter species can also be
reduced so that only the proximal parts are present.
Fissidens scalaris Mitt. has undivided to imper-
fectly divided peristome teeth that are smooth prox-
imally but papillose distally. All the species men-
tioned in this paragraph also have dimorphic stems.
In F. andicola (Herzog) Brugg.-Nann., originally
assigned to the segregate Simplicidens, the teeth are
long, undivided, and papillose throughout. In the
segregate Sainsburia the teeth, undivided above
and below, are split along the center of the median
line. Moreover, these teeth are papillose proximally
but striolate distally.
FISSIDENS subgenus FISSIDENS section SARAWAKIA
(Müll. Hal.) comb. nov.
Conomitrium section Sarawakia Müll. Hal., Gen. Musc.
Frond. 72. 1901. TYPE SPECIES: Fissidens beccarii
(Hampe) Broth.
Fissidens subgenus Sarawakia (Müll. Hal.) Z. Iwats. in
Hara, Origin Evol. Div. Pl. & Pl. Comm. 140. 1985.
Plants sordid to black-green, aquatic, submerged,
to 10 cm or more long, branched profusely, with a
pleurocarpous-like habit, branches tightly attached,
without basal rhizoids. Stem with basal and axil-
lary, smooth, reddish rhizoids; axillary hyaline nod-
ules differentiated; epidermis and outer 1–4 tiers of
cortical cells small, thick-walled, pigmented; inner
cortical cells larger, thin-walled, hyaline; central
strand differentiated. Leaves imbricate, numerous
pairs pinnately arranged, lanceolate to ovate, acute
to obtuse, sometimes apiculate; margin 6 entire,
limbate on proximal 4/5 or less of vaginant lami-
nae, limbidial cells uni- to pluristratose; costa
bryoides-type; dorsal lamina ending at insertion or
above, or short-decurrent; vaginant laminae 4/5
length of leaf, acute, 6 equal; laminal cells unis-
tratose or irregularly bistratose, firm-walled,
smooth, irregularly quadrate to hexagonal, eguttu-
late. Monoicous (gonioautoicous). Sporophytes one
[VOL. 107
per perichaetium; seta short; theca immersed to
emergent, ovoid, erect, radially symmetric, stoma-
tose or estomatose, exothecial cells oblong, vertical
walls thicker than horizontal walls; peristome un-
divided, poorly developed, bryoides-type; opercu-
lum conic, long-rostrate. Spores smooth, large for
the genus. Calyptra smooth, mitrate. No chromo-
some counts reported.
This is a small section consisting of only two
species, each known from collections made at a sin-
gle site, F. beccarii (Hampe) Broth. (Sarawak) and
F. hydropogon Mitt. (Ecuador). Fissidens acacioi-
des Schrad. [as F. stissotheca (Müll. Hal.) Mitt.,
Pursell 1999], previously included in this section,
is now considered to be better placed in section
Fissidens. As in subgenus Octodiceras, this section
is characterized by adaptations to a submerged,
aquatic habitat. However, the plants are mostly
darker, the stems and leaves firmer, and the peri-
stome is not as reduced as it is in Octodiceras. The
growth form is pleurocarpous-like (likened to that
of Cryphaea, Iwatsuki 1985, and to Phyllogonium,
Pursell et al. 1988). Pleurocarpous growth-forms
are a common adaptation to an aquatic habitat (Vitt
& Glime 1984). Otherwise, the two species of sec-
tion Sarawakia differ from rheophilous species in
section Fissidens in having a reduced peristome
and an absence of rhizoids at the base of tightly
attached branches.
FISSIDENS subgenus ALOMA Kindb., Eur. N. Amer.
Bryin. 2: 165. 1897. LECTOTYPE SPECIES (here
designated): Fissidens pauperculus M. Howe.
Fissidens section Areofissidens Müll. Hal., Syn. Musc.
Frond. 1: 46 1849 [1848]. LECTOTYPE SPECIES (desig-
nated by Pursell 1988): Fissidens palmatus Hedw.
Conomitrium section Reticularia Müll. Hal., Syn. Musc.
Frond. 2: 525. 1851, nom. illeg. incl. sect. prior.
Conomitrium section Polypodiopsis Müll. Hal., Linnaea
39: 360. 1875. TYPE SPECIES: Conomitrium metzgeria
Müll. Hal.
Conomitrium section Schistostegiopsis Müll. Hal., Lin-
naea 39: 362. 1875. TYPE SPECIES: Conomitrium hyalin-
um (Hook. f. & Wilson) Müll. Hal.
Conomitrium section Antennidens Müll. Hal., Linnaea 39:
364. 1875. LECTOTYPE SPECIES (here designated): Con-
omitrium undatum Müll. Hal.
Polypodiopsis (Müll. Hal.) A. Jaeger, Ber. Thätigk. St.
Gallischen Naturwiss. Ges. 1874–1875: 132. 1876.
Fissidens section Antennidens (Müll. Hal.) Paris, Index
Bryol. 46. 1894.
Fissidens subgenus Aloma section Camptodontii Kindb.,
Eur. N. Amer. Bryin. 2: 165. 1897. LECTOTYPE SPECIES
(here designated): Fissidens pauperculus M. Howe
Fissidens subgenus Aneuron Kindb., Eur. N. Amer. Bryin.
2: 165. 1897. TYPE SPECIES: Fissidens hyalinus Hook. f
& Wilson
Fissidens section Polypodiopsis (Müll. Hal.) Paris, Index
Bryol. 992. 1898.
Fissidens section Pycnothallia Müll. Hal., Gen. Musc.
2004] PURSELL & BRUGGEMAN-NANNENGA: FISSIDENS
Frond. 59: 1901 [1900]. LECTOTYPE SPECIES (here des-
ignated): Fissidens subglaucissimus Broth.
Fissidens section Semilimbidium Müll. Hal., Gen. Musc.
Frond. 60. 1901 [1900]. LECTOTYPE SPECIES (here des-
ignated): Fissidens flavifrons Besch.
Fissidens section Aloma Müll. Hal., Gen. Musc. Frond.
61. 1901 [1900]. LECTOTYPE SPECIES (here designated):
Fissidens pellucidus Hornsch.
Fissidens section Crenularia Müll. Hal., Gen. Musc.
Frond. 62. 1901 [1900]. LECTOTYPE SPECIES (here des-
ignated): Fissidens prionodes Mont.
Conomitrium section Bryoidiopsis Müll. Hal., Gen. Musc.
Frond. 74. 1901 [1900]. LECTOTYPE SPECIES (here des-
ignated): Conomitrium monandrum (Mitt.) Müll. Hal.
Conomitrium Mont. section Alomidium Müll. Hal., Gen.
Musc. Frond. 74. 1901 [1900]. LECTOTYPE SPECIES (here
designated): Conomitrium elachistophyllum Müll. Hal.
Conomitrium section Weberiopsis Müll. Hal., Gen. Musc.
Frond. 74. 1901 [1900]. LECTOTYPE SPECIES (designated
by Pursell 1988): Conomitrium commutatum Müll. Hal.
Conomitrium section Crenulidium Müll. Hal., Gen. Musc.
Frond. 75. 1901 [1900]. LECTOTYPE SPECIES (here des-
ignated): Conomitrium wilsonii Müll. Hal.
Conomitrium section Semilimbidiella Müll. Hal., Gen.
Mus. Frond. 76. 1901 [1900]. LECTOTYPE SPECIES (here
designated): Conomitrium intramarginatum Hampe.
Fissidens subgenus Polypodiopsis (Müll. Hal.) Broth.,
Nat. Pflanzenfam. 1(3): 352. 1909 [1901].
Fissidens subgenus Eufissidens section Weberiopsis (Müll.
Hal.) Broth., Nat. Pflanzenfam. 1 (3): 353. 1909 [1901].
Fissidens section Pycnothallia Müll. Hal. emend. Broth.,
Nat. Pflanzenfam. 1(3): 355. 1909 [1901].
Fissidens section Semilimbidium Müll. Hal. emend.
Broth., Nat. Pflanzenfam. 1(3): 356. 1909 [1901].
Fissidens subgenus Areofissidens (Müll. Hal.) M. Fleisch.,
Musci Fl. Buitenzorg 1: 20. 1900–1902 [1904].
Fissidens subgenus Eufissidens section Reticularia Broth.,
Nat. Pflanzenfam. 1(3): 353. 1909 [1901]. LECTOTYPE
SPECIES (designated by Pursell 1988): Fissidens mollis
Mitt.
Fissidentella Cardot, Rev. Bryol. 36: 17. 1909. TYPE SPE-
CIES: Fissidentella perpusilla Cardot
Fissidens section Pycnothallia subsection Glauculi P. de
la Varde, Ann. Cryptog. Exot. 2: 281. 1929. LECTOTYPE
SPECIES (designated by Bizot & Pierrot 1964): Fissidens
glauculus Müll. Hal.
Fissidens subgenus Eufissidens section Terminales Grout
subsection Aloma (Kindb.) Grout, Moss Fl. N. Amer.
1: 8. 1936.
Fissidens subgenus Eufissidens section Terminales Grout
subsection Crenularia (Müll. Hal.) Grout, Moss Fl. N.
Amer. 1: 8. 1936.
Fissidens section Semilimbidium subsection Monosticti P.
de la Varde ex Bizot & R. B. Pierrot, Rev., Bryol. Lich-
énol. 33: 232. 1964. TYPE SPECIES: Fissidens mathieui
Cardot
Fissidens section Semilimbidium subsection Obscuri P. de
la Varde ex Bizot & R. B. Pierrot, Rev. Bryol. Lichénol.
33: 232. 1964. TYPE SPECIES: Fissidens tisserantii Broth.
& P. de la Varde
Fissidens section Pycnothallia subsection Pseudobryoidi
Bizot & R. B. Pierrot, Rev. Bryol. Lichénol. 33: 233.
1964. TYPE SPECIES: Fissidens monostictus Broth. & P.
de la Varde
Plants small to medium, light to sordid green,
often reddish, particularly in stems and costa, ter-
restrial, saxicolous, lignicolous, corticolous, rarely
epiphyllous, rarely epizoic, typically in humid plac-
15
es, or infrequently aquatic; unbranched or
branched; branches with basal rhizoids; persistent
protonemata infrequent. Stems with axillary and
basal, smooth, mostly reddish, but sometimes bright
red, tan or colorless rhizoids; axillary hyaline nod-
ules differentiated or not; epidermis and outer 1–2
tiers of cortical cells, thick-walled, or large, thin-
walled and collapsed when dry; inner cortical cells
large, thin-walled, hyaline; central strand differen-
tiated or not, hyaline or, infrequently red; axillary
and subterranean, irregularly globose, multicellular
gemmae infrequent; axillary, fiilamentous, multi-
cellular, rhizoidal gemmae infrequent; axillary
stalked, clavate, multicellular gemmae frequent; fil-
amentous, multicellular gemmae at stem apices in-
frequent; epiphyllous, branched, multicellular gem-
mae rare. Leaves imbricate to distant, few to several
pairs pinnately arranged or frondiform, elimbate or,
more often, limbate, limbidium, when present, on
all laminae, or more frequently more or less con-
fined to vaginant laminae, marginal or intralaminal,
sometimes inconspicuous, limbidial cells mostly
unistratose, rarely pluristratose; costa variable in
length, sometimes lacking or nearly so, bryoides-
type; dorsal lamina mostly ending at insertion; va-
ginant laminae variable, minor lamina acute, ending
near margin or between costa and margin, or infre-
quently rounded and free distally; laminal cells un-
istratose to irregularly or regularly bistratose, small
to large with firm walls or large, thin walls, col-
lapsed when dry, smooth, mammillose, uni- or plu-
ripapillose, guttulate or eguttulate. Monoicous; per-
igonia variable in position; perichaetia terminal or
main stems and branches, naked antheridia and ar-
chegonia infrequent in leaf axils. Sporophytes 1–
several per perichaetium, yellow, darkening with
age; seta mostly elongate; theca usually erect, ra-
dially symmetric, stomatose, exothecial cells fre-
quently 6 quadrate, usually collenchymatous, ver-
tical walls often thicker than horizontal walls; peri-
stome scariosus-type, infrequently reduced, rarely
none; operculum conic, usually short-rostrate.
Spores finely papillose to smooth. Calyptra cucul-
late, rarely mitrate, smooth or prorate. n 5 5, 6, 8,
10, 12, 15 (Fritsch 1991).
This subgenus includes the largest number of
species in the genus. Most species are tropical and
subtropical in distribution. On the whole, the sub-
genus is characterized by a scariosus-type peri-
stome, a bryoides-type costa, and fewer that 40 files
of exothecial cells. Gametophytically, the subgenus
is heterogeneous and at first glance seems to be
incongruous. However, when considered with a
worldwide perspective no sharp breaks among the
sections recognized by Brotherus are apparent. In a
given character there is intergradation from one end
of the spectrum to the other. Thus it is impractical
16 THE BRYOLOGIST
to make infrageneric separations on cell size, lam-
inal cell wall ornamentation, or costa structure (e.g.,
from a typical bryoides-type costa to the absence
of a costa). Groups of species with smooth laminal
cells, elimbate leaves (e.g., F. pellucidus Hornsch.)
grade into those groups with elimbate and limbate
leaves, and mammillose and unipapillose laminal
cells [e.g., F. pellucidus var. papilliferus (Broth.)
Pursell and F. saülensis W. R. Buck & Pursell]
which, in turn grade into those groups that are lim-
bate and have pluripapillose laminal cells [e.g., F.
elegans Brid.). There are many more species with
mammillose, unipapillose, and pluripapillose lami-
nal cells than there are species with smooth-walled
laminal cells.
The distinctions between species of Brotherus’
sections Weberiopsis, Reticularia, and Aloma are
blurred. For example, F. ornatus Herzog has lam-
inal cells of two distinct sizes, 3–4 rows of small
(6–14 mm long) marginal cells and larger (18–40
mm long) inner cells. Fissidens inaequalis Mitt. and
F. inaequiretis I. G. Stone fall into this category,
too. Brotherus’ ecostate subgenus Polypodiopsis is
not sharply distinct from Brotherus’ subgenus
(Eu)Fissidens section Reticularia, both of which
are characterized by large, thin-walled, smooth
laminal cells. Although leaves of some species in
subgenus Aloma, as we understand it, appear to be
ecostate, there is usually a proximal vestige of a
costa present as Salmon (1899) demonstrated in F.
dealbatus Hook. f. & Wilson, F. hyalinus Hook. f.
& Wilson, and F. usambaricus Broth. This series
of species, which illustrates changes in the length
of the costa, begins with F. hyalinus in which there
is a weak, short proximal vestige of a costa. This
species is followed by F. dealbatus in which there
is a somewhat longer, but still weak costa in the
proximal parts of the vaginant laminae of the per-
ichaetial leaves. The costa in F. usambaricus is
similar to that in F. dealbatus, but in some peri-
chaetial leaves the weak costa ends a short distance
beyond the distal ends of the vaginant laminae (ob-
served by M. A. B.-N. in several collections of F.
usambaricus including an isotype in S). Fissidens
wageri Dixon and F. amazonicus Pursell, species
with large fusiform laminal cells, also show a costa
that ends a short distance beyond the distal ends of
the vaginant laminae in all leaves, as does the costa
in F. gardneri Mitt, a species with pluripapillose
laminal cells. Fissidens subulatus Mitt. also has fu-
siform laminal cells but the costa is long-excurrent.
The costa can be reduced to a single file of large
cells in the distal part of the leaf, as in F. cylindro-
thecus (Pursell & Aguirre 1991, Fig. 8), or, rarely,
to two files of large cells, as in F. ornaticostatus
H. Rob. (Robinson 1974, Fig. 2).
To our knowledge, the only epizoic species of
[VOL. 107
Fissidens is F. brachypus Mitt., restricted to a
freshwater sponge in the Amazon Basin of South
America (Buck & Pursell 1980). Persistent proto-
nemata are known in F. exilis Hedw. (Steere 1950),
F. protonemaecola Y. Sakurai (Iwatsuki & Suzuki
1982), F. saülensis Pursell and Buck (1996), and F.
subulatus Mitt. (Pursell & Reese 1980a). Fissidens
pocsii Bizot and F. subplanifrons Bizot & Onr.
have been observed by M. A. B-N. to be rarely
epiphyllous.
Iwatsuki and Suzuki (1982) reported the pres-
ence of caducous rhizoidal branches in the terres-
trial F. obscurirete Broth. & Paris, which detach
easily and probably initiate new independent plants.
Beever (Beever & Stone 1999) found the same con-
dition to be true in the aquatic F. waiensis and F.
rigidulus var. pseudostrictus. This condition is
probably much more frequent throughout the genus
but has gone unreported.
Axillary, filiform, multicellular, rhizoidal gem-
mae have been reported in F. stenopteryx Besch.
(Pursell & Bruggeman-Nannenga 1991); axillary,
stalked, multicellular, clavate gemmae are known
in F. pallidinervis Mitt. (as F. microcladus Thwai-
tes & Mitt., Imura & Iwatsuki 1988), F. bogoriensis
M. Fleisch. (Imura & Iwatsuki 1988), F. splach-
nobryoides Broth. (Iwatsuki & Suzuki 1982), F. yu-
catanensis Steere (Pursell 1994a,b), and F. zollin-
geri Mont. (Dixon 1923; Sainsbury 1935, 1955).
The latter have also been observed by R. A. P. in
F. dissitifolius Sull. and F. lindbergii Mitt. and
American specimens of F. angustifolius Sull., and
by M. A. B.-N in African, Asian, and American
specimens of F. flaccidus Mitt. Filamentous, mul-
ticellular gemmae have been observed by M. A. B.-
N. at the stem apices of African specimens of F.
serratus Müll. Hall.; epiphyllous, branched, multi-
cellular gemmae have been reported in F. bryum
Müll. Hal. (Potier de la Varde 1929a); and, subter-
ranean, irregularly globose, multicellular gemmae
have been observed by M. A. B.-N. to be common
in F. letestui P. de la Varde.
Perichaetial leaves of F. pseudoplumosus Bizot
& Onr. are characterized by an oblongifolius-type
costa (Bruggeman-Nannenga 1990). Marginal lam-
inal cells in F. letestui, F. mariei (Besch.) Broth.,
expressions of F. porrectus Mitt., and F. splendens
Brugg.-Nann. are thick-walled, and sometimes bis-
tratose, forming margins that are bulging and dark-
er than the interior cells.
The characteristic large, thin-walled laminal cells
of some species can be restricted to juxtacostal
patches in the proximal parts of the vaginant lam-
inae, as in a series of species consisting of F. an-
gustifolius Sull., F. zollingeri Mont., F. yucatanen-
sis Steere, and F. dissitifolius Sull., that superfi-
cially resemble species in section Fissidens of sub-
2004] PURSELL & BRUGGEMAN-NANNENGA: FISSIDENS
genus Fissidens. Cells in the dorsal and ventral
laminae of these species are progressively larger,
with the smallest cells found in F. angustifolius and
the largest in F. dissitifolius. All of these species,
however, have the characteristic scariosus-type
peristome.
Some corticolous species are characterized by
long cylindrical capsules and anomalous peristomes
(Pursell & Aguirre 1991), e.g., F. cylindraceus
Mitt., F. cylindrothecus Pursell & Aguirre, F. sub-
planifrons Bizot & Onr., F. pseudoplumosus Bizot
& Onr., F. pocsii Bizot & Dury, and F. lagenarius
Mitt. In all these species, the peristome teeth stand
erect even when moist. In the first two species the
teeth are divided nearly their entire length into two
slender filaments. On the other hand, F. lagenarius
is characterized by undivided or imperfectly divid-
ed, papillose teeth. Other corticolous and terrestrial
species within the section that also have anomalous
peristomes are F. braunii (Müll. Hal.) Dozy &
Molk., F. ellipticus Besch., F. gardneri, F. henryae
I. G. Stone, F. macroglossus (Broth.) Brugg.-
Nann., F. macrosporus Dixon, F. michoacanus
Thér., F. parkii Mitt., F. sigmocarpoides P. de la
Varde, F. subradicans Broth., and F. termitarum
(Herzog) Pursell.
More than 40 files of exothecial cells are found
in F. dealbatus Hook. f. & Wilson, F. exilis Hedw.,
F. gymnostomus Brugg.-Nann., and F. sedgwickii
Broth. & Dixon (Bruggeman-Nannenga 1989,
1997, and Bruggeman-Nannenga & Berendsen
1990). Fissidens gymnostomus Brugg.-Nann. and
F. scleromitrius (Besch.) Broth. are the only two
species in the genus that lack peristomes. Both of
these species, however, have a limbidium on por-
tions of the vaginant laminae of all or most leaves,
bryoides-type costae, and mammillose laminal
cells. Although the number of files of exothecial
cells is twice the usual number characteristic for the
subgenus, there is no question that these species
belong in subgenus Aloma. Both species have cu-
pulate theca, which could signify either a close re-
lationship or convergence.
The calyptra is mitrate and vesiculose in F. hy-
alinus, differing from the cucullate, smooth calyp-
tra present in other species.
KEY TO THE SUBGENERA AND SECTIONS OF
FISSIDENS
1. Leaves essentially ecostate -------------- subgenus Aloma
1. Leaves prominently costate -------------------------------------- 2.
2. Costa in median region of vaginant laminae
showing three stereid bands (two lateral and
one ventral 5 oblongifolius-type) ---------------
--- subgenus Pachyfissidens section Amblyothallia
2. Costa in median region of vaginant laminae
showing two stereid bands (both lateral 5
taxifolius- and bryoides-types) -------------------------- 3.
17
3. Costa with four or more peripheral guide cells and
1–5 large central cells ----------------------------------------------- 4.
3. Costa with two peripheral guide cells and one
large central cell, often reduced ------------------------------ 5.
4. Large, protruding axillary hyaline nodules
present on stem; peristome zippelianus-type
------- subgenus Pachyfissidens section Crispidium
4. Smaller axillary hyaline nodules present on
stem or lacking; peristome taxifolius-type -----
-- subgenus Pachyfissidens section Pachyfissidens
5. Dorsal and ventral laminae poorly developed or
lacking ------------------------------------------------------------------------
--- subgenus Fissidens section Fissidens [Nanobryum]
5. Dorsal and ventral laminae well-developed on
most leaves ------------------------------------------------------------------ 6.
6. Plants aquatic ------------------------------------------------------ 7.
6. Plants terrestrial, corticolous, epixylic, or epi-
lithic ---------------------------------------------------------------------- 9.
7. Plants delicate; leaves often 10 or more times lon-
ger than wide, fragile when dry -------------------------
------------------------------------------------------ subgenus Octodiceras
7. Plants sturdy, sometimes somewhat coarse; leaves
wider, usually less than 10 times as long as wide;
firm when dry ------------------------------------------------------------- 8.
8. Plants profusely branched, with a pleurocar-
pous-like habit; leaves limbate only on vagi-
nant laminae; branches lacking basal rhizoids
----------------- subgenus Fissidens section Sarawakia
8. Plants unbranched or only moderately
branched, not pleurocarpous-like in habit;
leaves limbate to variable degrees on some or
all leaves; branches with basal rhizoids ---------
-------------------- subgenus Fissidens section Fissidens
9. Median dorsal lamina cells large, longer than 30
mm, often as long 120 mm -------------- subgenus Aloma
9. Medial dorsal lalmina cells smaller, less than 30
mm long --------------------------------------------------------------------- 10.
10. Laminal cells mammillose, unipapillose or
pluripapillose; peristome scariosus-type -----
------------------------------------------------------- subgenus Aloma
10. Laminal cells smooth, often bulging; peri-
stome scariosus- or bryoides-type --------------- 11.
11. Leaves limbate to variable degrees, on all lami-
nae of all leaves to only on vaginant laminae of
perichaetial leaves -------------------------------------------------- 12.
11. Leaves elimbate ------------------------------- subgenus Aloma
12. Peristome scarious-type or anomalous, ca
32 files of exothecial cells, leaves limbate
on vaginant laminae of some or all leaves,
infrequently limbate on all laminae, guttu-
late ------------------------------------------- subgenus Aloma
12. Peristome bryoides-type or anomalous, 40
or more files of exothecial cells, leaves typ-
ically completely limbate on all leaves, leaf
cells typically smooth, not guttulate ------
---------------- subgenus Fissidens section Fissidens
ACKNOWLEDGMENTS
We thank Bruce Allen, William R. Buck, and Dale H.
Vitt for reading the manuscript and offering suggestions
for its improvement.
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