Multi-access key to the leafy liverworts of Victoria

Introduction

The multi-access key to the leafy liverworts of Victoria is a key to all 142 leafy liverwort taxa that are known to occur in Victoria. The leafy liverworts comprise plants with a stem bearing two or three ranks of leaves. In the case of species with a third rank of leaves, these leaves are positioned between the other two ranks on the side of the stem facing towards the substrate and are commonly referred to as underleaves. The sporophytes are produced at the end of a stem. Sometimes the stem is short making it appear as if the sporophytes are emerging directly out of the main stem below the stem apex. The sporophyte comprises a translucent seta and black capsule that dehisces by valves. Liverworts that do not have this combination of features are placed in the multiaccess key to thallose liverworts and hornworts. Most of those species have pad-like photosynthetic (green) parts that are appressed against the substrate, rather than a stem bearing two or three ranks of leaves. However, some genera excluded from this key have leaf-like structures that are not developmentally equivalent to the leaves of the leafy liverworts treated in this key (from subclass Jungermanniidae). These genera are Fossombronia, Petalophyllum and Treubia. Fossombronia and Petalophyllum have highly crisped leaf-like segments that are difficult to distinguish from one another and gives the overall plant the appearance of a small lettuce. Treubia comprises two ranks of leaf-like segments that are fleshy and multistratose throughout most of the leaf (most Victorian leafy liverworts with two ranks of leaves are unistratose) that grades into and is decurrent with the stem-like part at the base. Mosses are also often small plants that can superficially resemble leafy liverworts. Mosses in Victoria are never incised at the apex into lobed or lobules, they often have more than three ranks of leaves (which liverworts never do), they often have a multistratose costa (midrib) in the leaf (in liverworts the cells near the leaf centre may be abruptly differentiated from surrounding cells to form a vitta but are unistratose) and their sporophytes are easily distinguished. In many mosses the sporophyte has a capsule with an operculum, and often a peristome underneath, and the seta is coloured and not translucent.

This key was produced by Yennifer Longo during a 2022/2023 Willis Studentship placement offered by the Royal Botanic Gardens and funded by the Cybec Foundation. It contains 50 characters to help distinguish taxa from each other. It differs from other (dichotomous) keys provided in VicFlora by giving the user the flexibility to choose from one of the 51 characters provided rather than the user having to commit to investigate the particular features being questioned in the dichotomous keys. This has the benefit of:

avoiding having to answer questions relating to a feature that may not be

present in the sample when being identified, 2. potentially being able to identify a taxon without needing to comprehend the most technical terminology and 3. allowing the rapid identification of taxa with distinct features rarely encountered among the leafy liverworts

To use the key the user selects a feature from the features in the top left Features Available panel and clicks on the state present in their specimen to be identified. Once selected this feature and state will show as being selected in the Features Chosen panel directly below the Features Available panel. To undo a selection, click on the box next to the state chosen in the Features Chosen panel to erase the tick in the box. Once a feature state is chosen all the taxa that do not possess the chosen feature state are discarded in the bottom right Entities Discarded panel and those that do possess the chosen feature state are retained in the top right Entities Remaining panel. The user continues to choose further character states present in their specimen until identification is achieved. To restart the key when finished select the restart key icon represented by the two green arrows in the top left corner of the screen.

Go to Key Player

General helpful hints

For the most effective use of this key the user is encouraged to read the help guides for each character and state before submitting an answer for that character and state. Help guides and photographs can be found by clicking on the page icons beside characters and states. This help will reduce the chance of misinterpretation of the character and state. Some information and definitions required for interpreting a state may be given under character so users are encouraged to read the help guide for the characters before the states. When taking measurements or assessing which state a character exists in for the taxon being identified a typical plant and plant part should be used. Typical is here interpreted as a close representation of the average plant in terms of size and stature in a population and a plant part that represents the average condition for that character (e.g. average length, the most frequently observed shape etc.). Specimens should be hydrated to wet, not dry. For some of the characters it is possible for some taxa to possess more than one character state for the character. In such cases either correct character state can be chosen.

Both relatively straightforward and technical characters that require more expertise and access to compound microscopes are provided. Characters are grouped together in subjects, mostly based on the plant parts that they relate to. For each plant part, characters that can be assessed without a compound microscope are generally provided first followed by those that require a compound microscope. These latter characters are indicated with an asterisk. The least technical of the characters that don’t require a compound microscope are provided first for each plant part. These are often able to be determined without any visual aids. Characters that don’t require a compound microscope become increasingly more technical working down to the characters requiring a compound microscope (for plant parts where characters that require a compound microscope are provided). Similarly, among the characters that require a compound microscope, those that are least technical or prone to misinterpretation are provided first and become increasingly more technical working down toward the next plant part’s set of characters. Novices in liverwort identification are advised, if possible, to begin with the first characters provided for each plant part or subject first before moving into the more technically demanding characters for each plant part.

To help identify the specimen to be identified as quickly as possible the ‘best option’ tool can be used. This option highlights which characters should be used first to discard the maximum number of taxa among the remaining taxa. To use the best option, select the best icon given as a blue wand at the top of the screen. For some very similar species and varieties narrowing down to a single species may not be possible using this multiaccess key alone. In such instances the user can also use the dichotomous keys, descriptions and photographs provided on VicFlora to help distinguish between the remaining similar species.

Plant

A single character about the appearance of the overall plant.

Colour

This character refers to the overall colour of the plant, paying particular attention to the leaves (if the plants are large). It includes four states.

Stem

Four characters which refer to the main axis of the plant bearing the leaves.

Branching appearance

This character refers to how the branches are arranged in a plant. It includes four states.

Regular and pinnate

In this state branches are frequent from a main stem or stems, and are produced on both side of a main stem in a regular fashion.

Furcate or pseudo-dichotomous

In this state branching follows an evenly forked pattern, or nearly evenly forked.

Irregular or sparingly

In this state branching does not follow a particularly regular pattern of emergence, or is infrequent on main stems.

Leafy branches arising from stolons

In this state erect or ascending branches, bearing leaves, emerge from a system of creeping stems without normal leaves.

Branching type

This character refers to how a branch develops and the morphological consequences visible as a result and includes eight states.

Lateral-intercalary

In this state a branch emerges behind a stem apex on the sides in the zone where lateral leaves are attached. Emergence ruptures the stem epidermis and results in a minute unlobed collar of tissue around its perimeter at the branch base. This branching type is not associated with modified leaves.

Ventral-intercalary

In this state a branch emerges behind a stem apex on the underside in the zone where underleaves are attached or would be produced if the species produced underleaves. Emergence ruptures the stem epidermis and results in a minute unlobed collar of tissue around its perimeter at the branch base. This branching type is not associated with modified leaves.

Frullania-type

In this state a branch is formed at the apex of the stem and replaces the ventral half of the lateral leaf. This results in a branch that is associated with a lateral leaf on the stem of reduced width that is dorsal to the branch. The lateral leaf of reduced width (‘half-leaf’) can be simply narrower in the case of unlobed leaves, have half or around half the number of lobes in lobed leaves or be without a lobule in the case of leaves usually with lobules. In this type of branching there is no collar of tissue at its base.

Microlepidozia-type

In this state a branch is formed at the apex of the stem and replaces the dorsal half of the lateral leaf. This results in a branch that is associated with a lateral leaf on the stem of reduced width that is ventral to the branch. The lateral leaf of reduced width (‘half-leaf’) can be simply narrower in the case of unlobed leaves, have half or around half the number of lobes in lobed leaves or be without a lobule in the case of leaves usually with lobules. In this type of branching there is no collar of tissue at its base.

Acromastigum-type

In this state a branch is formed at the apex of the stem and replaces half of the underleaf. This results in a branch that emerges beside a narrower underleaf and in many cases an underleaf with half or around half the number of lobes. In this type of branching there is no collar of tissue at its base.

Radula-type

In this state a branch is formed at the apex of the stem but does not replace part of a leaf. This results in a branch with only fully formed leaves around it and that lacks a collar of tissue at its base.

Lejeunea-type

In this state a branch emerges behind a stem apex on the sides in the zone where lateral leaves are attached. Emergence ruptures the stem epidermis and results in a lobed collar of tissue around its perimeter at the branch base. This branching type is not associated with modified leaves.

Zoopsis-type

In this state a branch is formed at the apex of the stem and completely replaces a leaf. In this type of branching there is no collar of tissue at its base.

Surface

This character refers to the roughness of the stem surface and includes two states.

Smooth or papillose

In this state the stem is completely smooth or has minute bumps only visible under compound microscope.

Hispid

In this state the stem has obvious spicule-like protuberances.

Paraphylllia presence

This character refers to whether or not the stem has leaf-like appendages, much smaller than the lateral leaves, scattered along the stem.

Leaf

A set of 16 characters that refer to the main broadened photosynthetic segment of the plant, the lateral-leaves that are attached along the sides of the stem and branches.

Length

This character refers to the distance from the point where the lateral leaf is attached to the stem to the leaf tip, or tip of the longest lobe (both stem or branch leaves). Lateral leaves from mature branches or stems away from reproductive structures should be used. Often the lateral leaves at the shoot apex and base of a sporophyte (bracts) differ in size from the typical leaves and should be avoided. Instead focus on using lateral leaves midway along an elongate stem or branch. Length is divided into five ranges.

Shape

This character refers to the shape of the lateral leaves (both stem or branch leaves) when flattened and includes ten states.

Undivided, rounded to truncate

This state refers to intact, undivided lateral leaves which are circular, oblong or squarrose/rectangular in shape. The apex is rounded or flat, not shaped into a point.

Undivided, pointed

This state refers to intact, undivided lateral leaves which taper off into a point at their apex.

Bifid

This state refers to lateral leaves which are split at the apex into two main lobes.

3-fid

This state refers to lateral leaves which are split at the apex into three main lobes.

4-fid

This state refers to lateral leaves which are split at the apex into four main lobes. Segments that are toothed but smaller than other lobes have been referred to as laciniae. In this key they are also regarded as lobes.

More than 4-fid

This state refers to lateral leaves which are split at the apex into more than four main lobes. Segments that are toothed but smaller than other lobes have been referred to as laciniae. In this key they are also regarded as lobes.

Insertion

This character refers to how the angle that the leaves are attached to the stem. It has four states.

Succubous

In this state the leaves are inserted at an angle on the stem so that the margin facing the stem apex (acroscopic margin) is attached toward the ventral side of the stem and the margin facing the stem base (basiscopic margin) is attached toward the dorsal side of the stem. When the leaves overlap, this results in the bottom (basiscopic) margins of one leaf overlapping the top (acroscopic) margin of the leaf below it. This creates a shingle-like effect where water would run down and off the leaves, like the tiles of a roof.

Incubous

In this state the leaves are inserted at an angle on the stem so that the margin facing the stem apex (acroscopic margin) is attached toward the dorsal side of the stem and the margin facing the stem base (basiscopic margin) is attached toward the ventral side of the stem. When the leaves overlap, this results in the bottom (basiscopic) margins of one leaf being overlapped by the top (acroscopic) margin of the leaf below it. This creates a shingle-like effect where water would run down into the leaves if trickling from apex to base.

Transverse

In this state leaves are inserted at right angles to the axis of the main stem.

Acroscopic margin

This character refers to the dentition along the margin attached closest to the stem apex or in the case of transverse leaves the dorsal margin. It includes three states.

Entire

In this state the edge of the leaf is a smooth line without teeth or the margin is rough due to papillae on the cell surfaces.

Ciliate

In this state the edge of the leaf has thin hair-like projections, often one cell thick either at their base or along the majority of their length. These can be long or short.

Dentate

In this state the edge of the leaf is jagged with cells forming triangular tooth-like projections.

Basiscopic margin

This character refers to the dentition along the margin attached closest to the stem base or in the case of transverse leaves the ventral margin. It includes the three states given for the previous character (see there).

Total number of marginal teeth or cilia*

This character refers to how many teeth or cilia or scattered along the entire lateral leaf margin. Ot includes seven states.

Adaxial Cuticle*

This character refers to whether the surface of the lateral leaves facing upwards or toward the stem apex is textured or not and includes three states.

Smooth

In this state the adaxial surface of the lateral leaves are devoid of any ornamentation.

Papillose, striolate or verruculose

In this state the adaxial surface of the lateral leaves have bumps, grooves or warts.

Cellular protuberances

In this state the adaxial surface of the lateral leaves have cellular protuberances.

Abaxial cuticle*

This character refers to whether the surface of the lateral leaves facing downwards or toward the stem base is textured or not and includes the same three states as the previous character (see there).

Lamellae presence*

This character refers to whether or not there is a wing of tissue that is attached to the surface of the lateral leaf, appearing like a fin.

Lobe length

This character refers to the size of the largest lobe relative to the overall length of the leaf and includes two states.

Largest lobe apex

This character refers to the shape of the end of the largest lateral leaf lobe and includes two states.

Acute

In this state the leaf lobe comes to a point.

Rounded

In this state the end of the lobe is rounded, and in some cases the largest lobe is circular.

Largest lobe width

This character refers to how many cells wide most of the largest lateral leaf lobes are and comprises two states.

Mostly uniseriate

In this state the largest lateral leaf lobe comprises a single row of cells for most of its length.

Most of length at least two cells wide

In this state the largest lateral leaf lobe is at least two cells for most of its length. In many cases it will be far more than two cells wide.

Lobe orientation

This character refers to how the lobes are oriented relative to each other and to the unlobed (disc) section of the lateral leaf. It includes three states.

Not folded

In this state all the lobes are in the same plane as each other and the disc or the lobes form a continuously curved surface with leaf disc.

Folded with abaxial lobule

In this state the base of the smaller lobe folds and lies on the ventral or abaxial side of the main lobe forming a lobule.

Folded with adaxial lobule

In this state the base of the smaller lobe folds and lies on the dorsal or adaxial side of the main lobe forming a lobule.

Lobule form

This character refers to the 3-d structure and shape of the lobule and includes five states. It is only applicable to species with lobules (lobes that are folded to rest dorsally/adaxially or ventrally/abaxially over the main lobe).

Flat

In this state the lobule does not form a sac or is not inflated in any part.

Inflated or saccate, fused along basal margin with main lobe

In this state the lobule is broadly fused with the main lobe along its basal margin and is inflated at least near this line of fusion but can by completed saccate in some cases.

Saccate, separate from main lobe and clavate, taller than wide

In this state the lobule is saccate, forming a water-sac, and is only scarcely attached to the main lobe, if at all, and so appear quite separate from the main lobe. Lobules in this state are club-shaped, and are taller than wide (i.e. the axis of the lobule more or less parallel with the stem or branch is greater in size than the axis of the lobule that is approximately perpendicular to the stem or branch).

Saccate, separate from main lobe and falcate or beaked

In this state the lobule is saccate, forming a water-sac, and is only scarcely attached to the main lobe, if at all, and so appear quite separate from the main lobe. Lobules in this state are sickle shaped or look like a parrot’s beak, and are wider than tall (i.e. the axis of the lobule more or less perpendicular with the stem or branch is greater in size than the axis of the lobule that is approximately parallel to the stem or branch).

Saccate, separate from main lobe and galeate, similar height as width

Lobule margins*

This character refers to the number or teeth or cilia along the margins of the lobules. It contains six states. It is only applicable to species with lobules (lobes that are folded to rest dorsally/adaxially or ventrally/abaxially over the main lobe).

Lobule tooth length*

This character refers to how many cells long the longest tooth on the margins of the lobule are. It includes three states. It is only applicable to species with lobules (lobes that are folded to rest dorsally/adaxially or ventrally/abaxially over the main lobe).

All unicellular

In this state lobule will only ever produce teeth that are one cell long.

Two cells long on some lobules

In this state on an average stem there will at least one lobule that will have teeth two cells long. There will also likely be lobules on the same stem that have teeth formed by only a single cell.

Three or more cells long on some lobules

In this state on an average stem there will at least one lobule that will have teeth three cells long.

Underleaf

A set of seven characters that refer to the row of leaves situated on the ventral surface of the stem, between both sets of lateral leaves.

Presence

This character refers to whether or not the plant has underleaves.

Size

This character refers to the overall size of the underleaf in relation to the lateral leaves. It includes four states.

Form

This character refers to the shape of the underleaf when flattened and includes six states.

Undivided and rounded

This state refers to undivided underleaves which are circular, oblong or rectangular in shape. The apex is rounded and does not come to a point.

Undivided and pointed

This state refers to undivided underleaves which taper into a point at their apex. Leaves in this state are often lanceolate or triangular.

Bifid

This state refers to underleaves which are split at the apex into two main lobes.

3-fid

This state refers to underleaves which are split at the apex into three main lobes.

4-fid

This state refers to underleaves which are split at the apex into four main lobes. Segments that are toothed but smaller than other lobes have been referred to as laciniae. In this key they are also regarded as lobes.

More than 4-fid

This state refers to underleaves which are split at the apex into more than four main lobes. Segments that are toothed but smaller than other lobes have been referred to as laciniae. In this key they are also regarded as lobes.

Margins

This character refers to whether the edges of the underleaf are toothed or ciliate or not and includes three states.

Lobe size

This character refers to the length of the lobe relative to the overall length of the underleaf and includes two states.

Connection to lateral leaves

This character refers to whether the underleaves are connected at their base to the lateral leaves or not. It contains three states.

Free of lateral leaves

This state describes underleaves which are only connected to the stem of the plant, and do not connect to the lateral leaves.

Bilaterally united to lateral leaves

This state describes underleaves that are connected to the main stem and both of the lateral leaves closest to it at their base. This connection may only be by a single row of cells.

Unilaterally united to a lateral leaf

This state describes underleaves which are connected to the main stem and only one nearby lateral leaf at their base. This connection may only be by a single row of cells.

Ratio of underleaves to leaves

This character refers to how many underleaves there are relative to the number of lateral leaves and comprises two states.

One underleaf for every two or more lateral leaves

In this state a pair of lateral leaves are separated by a single underleaf. In some species that comply with tis state the underleaves are sporadically produced and may be absent between some lateral leaves.

One underleaf for every leaf

In this state near the base of each lateral leaf there is an underleaf. In Victoria this state exists only in Colura pulcherrima and Diplasiolejeunea plicatiloba.

Stylus

A single character referring to a separate lobe that occurs in few species that appears as an appendage between the main lobe and lobule of a lateral leaf.

Presence of conspicuous stylus composed of 8 or more cells*

This character refers to the presence and size of a stylus.

Conspicuous stylus present

In this state the stylus is usually triangular in shape and is composed of more than eight cells make it reasonably noticeable with a compound microscope.

Stylus absent or inconspicuous

In this state the stylus is completely absent or if it is present, it is filiform and composed of less than eight cells.

Leaf Cells

A set of ten characters relating to the cells of lateral leaves. In species with lobules, these characters relate to cells of the main lobe.

Trigone size*

This character contains two states which refer to the presence or absence of prominent triangle-shaped wall thickenings at the junction where three or more cells meet.

Massive, often coalescing

In this state the trigones are large and obvious and often adjacent trigones fuse together (coalesce), but if not coalescing are at least wider that the non-thickened cell wall between them. The result of these large trigones is that the area of trigones can approach or be a similar area of the cell lumens (cavities) in a leaf.

Small or absent

In this state the cell walls do not become thickened in the corners and trigones seem to be absent or if the trigones are present, then they are small and do not contribute a significant proportion of the length of the cell wall between the corners (i.e. the intervening non-thickened section of the cell wall is longer than the width of the trigone) and they do not contribute a significant proportion of the leaf area.

Vitta*

This character refers to the presence or absence of a distinct strip of elongated cells down the center of the leaf. These cells are much more elongate than rows of cells immediately beside them.

Cell wall thickness*

This character refers to the relative thickness of the cell walls and contains two states.

Thin

In this state the cell wall apart from any trigones is thin such that if you were to illustrate the cells it would be more accurate at around x100 magnification to represent them as a single line.

Thick

In this state the cell wall is thick enough that if you were to illustrate the cells it would be more accurate at around x100 magnification to represent them as two lines to define the edges of the lumen and cell walls.

Uniformity*

This character refers to how the dimensions of the laminal cells varies throughout the lateral leaf. When assessing this character care needs to be taken to cut or brake leaves off at the attachment to the stem rather than tearing the leaves off. If leaves are carelessly torn off some of the basal rows of cells may remain on the stem which is an issue for the states in which the basal cells rows differ from those elsewhere in the leaf. This character comprises five states.

Mid-leaf cell diameter*

This character refers to the distance across the lumen of cells in the middle of the leaf for unlobed leaves or in the middle of the disc for lobed leaves and includes three range states. Measure the axis roughly parallel with the stem.

Presence of intermediate thickenings in cell wall of mid-leaf*

This character refers to whether or not there is a swelling or circular to elliptic node in the cell wall around half way between the corners of the cells.

Oil body presence*

This character refers to whether or not there are oil bodies in each cell. Oil bodies are organelles which contain oil and can have a brown, grey or hyaline colour. They can be easily confused with chloroplasts which are usually numerous and green when fresh but can turn to brown with age. Oil bodies usually degrade rapidly after a plant is collected and so oil body characters should only be assessed on fresh material.

Oil body number*

This character refers to the number of oil bodies that occur within an individual leaf and comprises seven states.

Oil body colour*

This character refers to whether the oil bodies are distinctly dark and opaque or whether they are mostly hyaline or very pale.

Oil body texture*

This character refers to the appearance of the oil body and includes two states.

Smooth, homogenous

In this state the oil body does not have any surface ornamentation and it is not composed of granules or globules.

Granular or segmented

In this state the oil body can have a granular surface and can be composed of granules or globules and resemble a compact cluster of grapes (i.e. botryoidal).

Rhizoid

A single character about the hair-like structures that provide anchorage.

Location

This character refers to the presence and location of rhizoids. When assessing this character, rhizoids will most likely be produced where the stem is in contact with substrate so it is in these sections of stem that the rhizoids should be searched for.

Base of underleaf or where underleaf would be if absent

In this state the rhizoids are produced in a tight clump immediately below or on the base of the underleaf or in the position on the stem where an underleaf would be expected to be if the species had underleaves.

Base of lateral leaf

In this state the rhizoids are produced in a tight clump at the base of a lateral leaf.

Lobule

In this state the rhizoids are produced in a clump from the outer surface of the lobule.

Scattered along stem

In this state rhizoids are not clearly aggregated into clusters but are instead scattered, either sparsely or densely, along the stem.

Excluded from leafy stems and restricted to stolons

Species that have stems differentiated into creeping leafless stolons and erect or ascending leafy axes will belong to this state. In this state the leafy stems are not in contact with the substrate, whereas the leafless stolons are and so it is here that the rhizoids are produced.

Reproduction

A set of eleven characters relating to the reproductive structures and sexuality of the plant.

Position of female structure

This character refers to where on the plant the female reproductive structures are produced and includes four characters. The female part of the plant is easily identified from where the sporophyte (capsule and the seta) is when sporophytes are visible. Often these wont be present but structures associated with the sporophyte such as perianths, bracts, marsupia or coelocaules (see female structures present character) may be and can be used to define where the female structures are produced.

Terminal on long leafy axis

In this state the female structures are at the end the stem or a branch with several normal leaves. When the branch or stem produces a sporophyte afterwards it can branch below it eventually resulting in the female structures appearing as if it was produced on the side of the stem.

On short lateral branch

In this state the female structures are produced on very short branches without normal leaves but with the structures associated with the sporophyte such as those given as states for the female structures present character. These short branches emerge from the main stem along the sides in the zone where the leaves are attached. The short size of the branch can give an initial impression that the sporophyte is produced directly on the side of the main stem.

On short ventral branch

In this state the female structures are produced on very short branches without normal leaves but with the structures associated with the sporophyte such as those given as states for the female structures present character. These short branches emerge from the main stem on the ventral surface in the zone where the underleaves are produced or would be expected to be produced if absent. The short size of the branch can give an initial impression that the sporophyte is produced directly on the main stem.

On short basal branch

Bracteole presence near female reproductive structure

This character refers to whether or not the modified rank of leaves equivalent to the underleaves (bracteoles) are present immediately below the sporophyte.

Female structures present

This character refers to the type of structure that forms the main role in surrounding and protecting the developing sporophyte and includes five states.

Bracts (and sometimes bracteoles) only

In this state the only well-developed structure that surrounds the developing sporophyte are bracts and sometimes bracteoles. Bracts are modified leaves that are usually a different size and have a different degree of dissection than the normal vegetative leaves, but will otherwise look similar to the leaves.

Perianth

In this state the developing sporophyte is surrounded by a protective leaf-like envelope, the perianth. The perianth are similar to bracts but the perianth differs in being a more fused structure rather than being composed of separate parts like individual bracts.

Coelocaule

In this state the stem below the sporophyte enlarges and swells around the sporophyte so that the sporophyte is embedded into a hollow enlarged stem. Coelocaules follow the same axis of growth as the stem below and bear bracts on their surface because they are derived from stem tissue. The occurrence of bracts and the general fleshy and non-herbaceous appearance of coelocaules helps them to be distinguished from perianths.

Marsupium

In this state the stem becomes swollen near the developing sporophyte, envelops it and grows downwards, usually perpendicular to the axis of the rest of the stem. Its surface is often covered in rhizoids. The fleshy texture and direction of growth allows marsupia to be distinguished from other reproductive and when they are covered with rhizoids are unmistakable.

Shoot calyptra or perigynium

In this state the developing sporophyte is surrounded by tissue derived from the female organ that produces the eggs (archegonium) and cells immediately surrounding the archegonium. They are fleshy, often tubular, continue to axis of growth of the stem, but do not bear bracts over their surface like in coelocaules. They often bear vestigial perianths at their apex.

Perianth surface

This character applies to species that produce a perianth (see Female structures present character). It refers to whether or not there are bristly or tooth-like outgrowths over its external surface.

Perianth cross section near apex

This character refers to how the perianth (see Female structures present character) would look if it was cut perpendicular to its axis of growth near the apex of the perianth and the cut is viewed looking down. This character applies only to species that produce perianths and contains five states.

Terete

In this state the cross section does not have clear angles but is instead more or less circular.

Bilabiate

In this state the perianth has two ridges or wings and two intervening surfaces. This results in a cross section that has two angles and curves connecting them.

Trigonous

In this state the perianth has three ridges or wings and intervening surfaces. This results in a cross section that has three angles and edges or curves connecting them, like a triangle.

4 or 5-plicate

In this state the perianth has four or five folds, ridges or wings at least near the apex of the perianth. The perianth may be terete toward its base.

6- or more plicate

In this state the perianth has six or more folds, ridges or wings at least near the apex of the perianth. The perianth may be terete toward its base.

Marsupium surface

This character refers to whether or not the surface of the marsupium (see Female structures present character) is covered in rhizoids or not. It only applies to species that produce marsupia.

Bracteolar antheridia presence

This character refers to whether or not modified underleaves (bracteoles) among the male reproductive part of the stem (androecium) bear antheridia. Antheridia are the male organs that produce sperm and are produced on the adaxial surface and appear like spheres on a short and narrow stalk.

Antheridia number per bract

This character refers to the number of male organs (antheridia) produced on each bract in the male reproductive part of the stem (androecium). Antheridia appear like spheres on a short and narrow stalk.

Plant sexuality

This character refers to which sexes can be present in an individual plant and consists of two states.

Dioicous

In this state male organs are produced on completely separate plants to the plants that produce female organs.

Monoicous

In this state male and female organs are produced on the same plant.

Asexual gemmae presence

This character refers to whether or not asexual propagules like gemmae are produced. Asexual gemmae are often small few-celled granule objects produced on the surfaces or margins of leaves, often near the stem apex, or can be in the form of leaves, teeth or lobes that readily detach and become new plants. It is advised that this character be used if structures that are definitely gemmae are observed because it is possible that a plant could produce asexual gemmae (and so would match the state of present for this character) but was currently not producing gemmae at the time it was observed. In this case if absent was selected, misidentification would likely result.

Capsule wall thickness*

This character requires the user to make a thin cross section through the capsule and look down on the cross section with a compound microscope and count the number of cell layers that form the wall.