Plants acrocarpous. Stems, except for initial stages, growing from a 2-sided apical cell. Leaves distichous, equitant, each consisting of 2 vaginant laminae clasping the stem, a ventral and dorsal lamina (except Nanobryum); costa single, usually well-developed, sometimes reduced or absent or nearly so. Peristome single, haplolepidous, endostomate, rarely absent; teeth 16, usually divided ½–⅔ their length, rarely undivided or irregularly divided or reduced.

The family has been previously divided into a number of segregate genera, but it is now usual to treat all in a single genus, Fissidens, which has been variously divided into subgenera and sections. Around 900 species names are listed in Index Muscorum, with more than 450 species currently accepted (Crosby et al., 2000) and together having an almost world-wide distribution, except for high Arctic and Antarctic regions. The greatest species diversity is found in the tropics.

Since the original manuscript was prepared for Australian Fissidens, there have been significant taxonomic changes at the subgeneric level and in species synonymy. Some 57 species and 16 infra-specific taxa of Fissidens are reported here from the mainland States and Territories of Australia, nine species and four varieties are apparently endemic, and many are restricted to coastal Queensland which has, historically, been the primary focus of collectors in Australia.

Plants can be erect or ± prostrate, scattered or gregarious, occasionally forming dense turves or cushions, terrestrial, rupestral, epiphytic, on soil, rock or bark, occasionally aquatic. Some species are important colonisers of bare soil, particularly roadside banks and even termite mounds.

Fissidentaceae Schimp., Coroll. Bryol. Eur. 20 (1856). Type: Fissidens Hedw.

Taxonomic Synonyms

Skitophyllaceae Mitt., J. Linn. Soc. Bot. 12: 11, 23, 580 (1869). Type: Fissidens semicompletus Hedw. (Lectotype designated by Pursell & Bruggeman-Nannenga, 2004).

Schistophyllaceae Lindb., Utkast Eur. Bladmoss 16 (1868), 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.

Nanobryaceae Schultze-Motel, Willdenowia 5: 386 (1969). Type: Nanobryum dummeri Dixon.

Fissidentaceae is characterised by the peculiar leaf structure which is essentially isobilateral (not dorsiventral as in most other mosses) and for which there have been several explanations that were fully discussed by Salmon (1899) and, more recently, by Robinson (1970). The most widely accepted interpretation (supported by Salmon) dates back to Robert Brown (1819) who proposed that the vaginant laminae represent the true leaf, with the addition of apical and dorsal appendicular outgrowths. Another theory, initiated by Spruce (1881), suggests that the whole Fissidens leaf constitutes the true leaf, the apical and dorsal laminae being the middle lobe of a trilobed leaf turned on its axis. Robinson (1970) elaborated on this by suggesting that a mutation caused a change in the mitotic spindle of the leaf primordium resulting in reorientation of the upper part of the leaf.

Relationships of the Fissidentaceae appear to be with the Dicranaceae (Bruggeman-Nannenga & Berendsen, 1990; La Farge et al., 2000; Buck & Goffinet, 2000). While both families share similar peristomes, the primary difference is in leaf structure. However, the peristome of F. taylorii var. sainsburiana is also similar to that found in some Grimmiaceae. The phylogenetic position of the Nanobryaceae, originally treated in this Flora as distinct, is perhaps equivocal. Pursell & Reese (1980) synonymised Nanobryaceae in Fissidentaceae and Nanobryum in Fissidens, a position supported by Bruggeman-Nannenga & Berendsen (1990). Stone (1982) argued strongly for the retention of the genus Nanobryum for those species lacking a “Fissidens-type” leaf (N. dummeri and N. thorsbornei). At least N. thorsbornei shares the limbate dorsal and apical laminae composed of large, smooth cells and features of the peristome with Fissidens sect. Fissidens. In the present treatment, Nanobryaceae and Nanobryum are treated as belonging to Fissidentaceae and Fissidens, respectively.

The family Archifissidentaceae Dixon & P. de laVarde, nom. illeg. was replaced by the Nanobryaceae Schultze-Motel (Schultze-Motel, 1969) with a single species, Nanobryum gladiolum (Mitt.) Bizot [= N. duemmeri Dixon (dummeri) fide Bizot (1963)] from Uganda and Cape Province, South Africa. The Archifissidentaceae was established to accommodate Nanobryum as an ancestral form of Fissidens. However, the family is based on two character states, viz. linear to linear-rhomboid lamina cells and a persistent protonema.

Based on morphological variation within Fissidens and Nanobryum, Pursell & Reese (1980) reduced Nanobryaceae to synonymy of Fissidentaceae and Nanobryum to synonymy of Fissidens. Iwatsuki & Suzuki (1989) apparently did not accept the transfer and reported Nanobryum thorsbornei from New Caledonia. Bruggeman-Nannenga & Berendsen (1988) agreed with Pursell & Reese (1980) and discounted the importance of the lack of dorsal and apical laminae in Nanobryum, placing greater emphasis on the bryoides-type peristome ornamentation which they suggested was found only in Fissidens. However, they acknowledged that the fasciculatus-type peristome was also found in genera of the Dicranales, as well as in Fissidens.

Stone (1982, 1990c) provided a cogent argument for the retention of Nanobryum and the family Nanobryaceae. Clearly, confusion persists around the placement of this minute and morphologically somewhat anomalous moss. Stems and leaves of very young plants or very short-stemmed species of Fissidens are often different to typical longer-stemmed forms. Molecular genetic studies may provide corroborative evidence for the retention of Nanobryaceae and Nanobryum or their retention in Fissidentaceae and Fissidens, respectively.

However, placing greater emphasis on peristome structure, N. thorsbornei is included here in Fissidens.

There have been two recent critical evaluations of the subgenera and sections of Fissidens. Pursell & Bruggeman-Nannenga (2004) recognise four subgenera, based on gametophyte and sporophyte characters: Aloma, Fissidens (with Sections Fissidens and Sarawackia), Octodiceras and Pachyfissidens (with Sections Amblyothallia, Crispidium and Pachyfissidens). A second infrageneric classification (Suzuki & Iwatsuki 2007) utilises morphological characters of the gametophytes and peristome teeth as well as cytotaxonomic evidence relating to sexuality. Those authors recognised six subgenera: Fissidens (with Sections Fissidens, Semilimbidium and Aloma); Aneuron; a new subgenus Neoamblyothallia (with Sections Neoamblyothallia and Crispidium); Pachyfissidens (with Sections Pachyfissidens and Serridium); Octodiceras; and Sarawakia.

In this treatment we have accepted the subgeneric division proposed by Pursell & Bruggeman-Nannenga (2004) with slight modification after Suzuki & Iwatsuki (2007). Several rearrangements of species in different sections have led to some confusion in the placement of species, and it is probable that future morphological and molecular phylogenetic comparison will result in further rearrangement.

A new classification of the Fissidentaceae has recently been published by Suzuki & Tsubota (2018). Based on the phylogeny and morphological reassessment, they recognise three subgenera: Pachyfissidens, Neoamblyothallia, and Fissidens. Subgenus Neoamblyothallia consists of two sections:  Neoamblyothallia, Crispidium; subgenus Fissidens consists of five sections: Fissidens, Polypodiopsis, Aloma, Areofissidens, Semilimbidium. "High diversity of the most derived section Semilimbidium in the tropics suggests that the evolutionary history of the genus is through adaptation and diversification in tropical regions.”

In this On-Line treatment of Fissidens in Australia the author has not used infrageneric categories.

Peristome structures:

Peristome characters were studied by Fleischer (1904) who recognised two types in which the filaments (forks) were, respectively, spirally thickened or papillose. Following scanning electron microscopic (SEM) observations, Allen (1980) recognised seven distinct types that were only partly correlated with the sections of Fissidens subg. Fissidens. Bruggeman-Nannenga & Berendsen (1990) investigated many species, including 28 identified as Australian (some by SEM, but most by light microscopy) and distinguished five basic types. They also recognised several other types of peristome found in very few species and not characteristic of any section, and they alluded to the potential taxonomic use of the number of columns of exothecial cells around mid-capsule. Thus, the scariosus-type peristome is correlated with (28–) 32 (–40) columns of exothecial cells around the capsule periphery while other peristome types usually have more than 40.

Key to main peristome types:

[Adapted from Bruggeman-Nannenga & Berendsen (1990) and Suzuki & Iwatsuki (2007)]

1. Peristome teeth usually divided into 2 equal filaments above; trabeculae distinct throughout the filament;
teeth broad and flat; lamellae of the abaxial (outer) side of filaments with rather high oblique riblets..........taxifolius-type

1: Peristome teeth divided above or not; trabeculae not distinct in the distal part of the filament; teeth various................2

    2. Trabeculae of abaxial side in undivided basal part of tooth higher than lamellae..........................................................3

    2: Trabeculae and lamellae of abaxial side in undivided part of tooth approximately equal in height..............................4

3. Trabeculae of abaxial side at bifurcation coarsely papillose, double at the margins, not forked;
distal part of filament appearing spirally thickened (actually oblique riblets on abaxial and adaxial surfaces).................................................................................................................................................................bryoides-type

3: Trabeculae of abaxial side at bifurcation smooth and double with forked ends;
distal part of filament with irregular squamae or spikes................................................................................zippelianus-type

    4. Distal part of peristome teeth entire, rimose or occasionally weakly split;
distal part coarsely and irregularly papillose...................................................................................................sainsburia-type

    4: Distal part of peristome teeth split into 2 filaments;
filaments appearing spirally ornamented or with deflexd squamae.....................................................................................5

5. Distal part of filament spirally ornamented; abaxial surface at the bifurcation with trabeculae,
and marginal vertical walls forming a continuous smooth ridge with rounded corners...............................scariosus-type

5: Distal part of filament with deflexed squamae; abaxial side at the bifurcation lacking marginal vertical walls;
lamellae with vertical striae, often papillose....................................................................................................similliretis-type

Costal structure

Stone (1990b) and Bruggeman-Nannenga (1990) independently studied the costal structure of Fissidens species in relation to classification and the latter defined the following main types using transverse sections of the costa in the mid-region of vaginant laminae of vegetative leaves:

oblongifolius-type, with 4 or more large guide cells in a U- or V-shape with 1–5 large central cells in 1–3 rows connecting to the dorsal lamina; an adaxial and 2 lateral bands of stereids or substereids; epidermal cells differentiated or not; junction of vaginant laminae and costa formed of laminal cells.

taxifolius-type, with 2–4 or more superficial, adaxial guide cells and 1–8 central connecting cells in 1 or 2 rows or a random mass; 2 lateral stereid or substereid bands; epidermal cells with broader lumina; junction of vaginant laminae and costa formed of guide cells and stereids.

bryoides-type, typically with 2 superficial adaxial guide cells and 1 large central connecting cell (always between the 2 guide cells) and occasionally more cells between this and the dorsal lamina; 2 lateral bands of stereids or small cells; epidermis differentiated or not; junction of vaginant laminae and costa formed of laminal cells.

Illustrations in Stone (1990b) demonstrate that the arrangement of the large central cells (which are morphologically similar to guide cells) in either 1 or 2 rows (occasionally a random mass) in the simple upper part of the leaf above the vaginant laminae matches that of the large connecting cells in the sheathing region.

Throughout this treatment of the family, the dimensions provided are of well-soaked plants: ‘minute’ plants are up to 3 mm tall; ‘small’ plants 3–10 mm; ‘medium-sized’ plants 10–20 mm; and ‘large’ or ‘robust’ plants more than 20 mm tall or long. Unless stated otherwise, cell details are for those from the mid-dorsal lamina opposite the junction of the vaginant laminae in a mid-stem (vegetative, not perichaetial) leaf; exothecial cell details are for those in mid-theca; costal structure refers to mid-vaginant lamina region: in ‘dorsal lamina tapered to the base’, the base refers to the leaf base; where the vaginant laminae are unequal, ‘minor’ lamina refers to the smaller of the two. Some species of Fissidens are dimorphic, the fruiting 4 plants being quite different in appearance from the vegetative plants, e.g. F. curvatus. Fruiting plants usually have short stems bearing a few pairs of juvenile leaves at their base. These leaves grade into larger subperichaetial and perichaetial leaves which surround a sporophyte at the stem apex, where further leaf production has ceased. Strictly vegetative leaves can be confined to a single pair on such plants, or they can even be completely lacking. Vegetative plants, on the other hand, after producing juvenile leaves at the stem base, can produce many pairs of vegetative leaves. These are uniform in size and smaller than perichaetial leaves.

Microscopic examination of specimens

Cell shape and size, limbidia (particularly where confined to the proximal part of the vaginant laminae), cell structure and surface ornamentation, costal structure and stem sections are best observed using cleared preparations. Whole moist leaves and thin hand-cut sections are mounted under a coverglass on a microscope slide and undiluted lactic acid carefully infiltrated under the coverglass before warming the preparation carefully and gently over a spirit lamp flame. This removes cellular contents and makes for much easier observation of critical structures, particularly surface papillae and mammillae or other wall thickening. Subsequent infiltration of an 0.5% Toluidine Blue O solution can enhance visibility after clearing tissues and sections. Another potentially useful aid to the identification of some taxa, although not employed here, is to observe the colour reaction of moist leaves to potassium hydroxide (KOH 10% solution), a technique that has proven to be particularly useful in Pottiaceae (Zander 1993).

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Stone, I.G. (1983b). Fissidens gymnocarpus, a new species from Queensland, Australia. J. Bryol. 12: 553–557.

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Adapted from R.D. Seppelt & I.G. Stone (2016), Australian Mosses Online 70. Fissidentaceae. (Australian Biological Resources Study: Canberra). Version 16 June 2016. http://www.anbg.gov.au/abrs/Mosses_online/70_V2_Fissidentaceae.html

This treatment of the family Fissidentaceae, originally intended for publication in Flora of Australia, is based on a manuscript prepared by Ilma Stone in 1992, and substantially updated in 1998. The authors of that early treatment, Ilma Stone and David Catcheside, are both deceased.

Since the preparation of the original manuscript, the infrageneric classification of Fissidens has altered considerably, along with the recognition of many new taxa and new synonyms, while the monogeneric Nanobryaceae has been subsumed into the Fissidentaceae. Therefore, an up-to-date, illustrated treatment of the most diverse moss genus in the Australian flora was clearly long overdue.

To accommodate recent revisions and records, a new treatment of the Australian Fissidentaceae has been prepared by Rodney D. Seppelt for publication in Australian Mosses Online, drawing heavily on the pioneering work of Ilma Stone. Descriptions of taxa have been expanded, all taxa are or will be fully illustrated, and a new key to species has been constructed. Over many years, Ilma Stone collected hundreds of Fissidens specimens, with particular emphasis on the tropical and subtropical regions of Australia. The subtropics support the richest Fissidens flora, and the majority of Ilma's specimens were of species that have proved to be notoriously variable and difficult to identify with certainty. Many challenges lie ahead, and updates to this treatment can be expected.