Jungermanniales H.Klinggr.
Artificial key to genera of Jungermanniales of New Zealand 1
1. This key is adapted, with considerable modification, from Schuster (2000a). Extraterritorial genera that are known only from southern South America or the subantarctic islands are included, but placed in brackets, since a) all are included in various discussions within this flora, and b) a possibility exists that these genera eventually may be discovered in New Zealand.
General Key
1
Vegetative gametophyte lacking recognizable leaves (but gametangial branches leafy)
Group A
2
Leaves varied in form but never clearly complicate-bilobed (abaxial face never ridged or winged) and never with a water-sac of ventral margin, or forming a ventral lobe
Leaves complicate-bilobed, the two lobes clearly folded over each other; or leaves, on ventral side, with a distinct water-sac or lobule (which is hidden from view, in dorsal aspect of plant); the lobule reduced or vestigial in a few taxa (then leaves appearing simply incubously shingled)
3
Leaves with lamina absent or under 1 cell-tier high, divided to base into 2–4(6) uniseriate lobes; plants usually confervoid
Group B
Leaves, lobed or unlobed, with distinct laminae, usually 2–many cells high (even when leaves are deeply lobed)
5
Rhizoids (usually frequent) scattered over ventral face of stem, or at ventral bases of leaves; underleaves 1–2-lobed, or lacking, lobes often ciliate. Leaves 2–3- or 3–4(5)-lobed only, always alternate; sexual branches never reduced and ventral-intercalary (exc. Jackiella). (Lophozioideae, Plagiochilaceae and Acrobolbaceae)
see Group H
Rhizoids (often few or very local) only at bases of the distinct underleaves, or at bases of reduced appendages of stolons and/or flagella; underleaves often 3–4-lobed or more, lobes rarely ciliate, rarely reduced, then of 2 minute segments
6
Vegetative (leafy) shoots, and/or flagelliform, microphyllous axes and/or sexual branches all, or in part, ventral-intercalary (at times such branches infrequent, but always present if searched for)
Group C
Vegetative (leafy) shoots with branches all lateral (usually 1–2–3-pinnate, regularly or irregularly so); gametangia always on elongated leafy axes (acrogynous; acroandrous); geotropic, microphyllous, flagelliform branches never produced
Group D
7
Leaf insertion (or at least orientation) clearly incubous. Underleaves nearly always distinct, large, as broad as stem or usually broader; rhizoids confined to fascicles from underleaf bases
Group E
Leaf insertion (and orientation) transverse or succubous. Underleaves large to none; rhizoids scattered, or in fascicles from underleaf or leaf bases
8
Isophyllous or subisophyllous: underleaves similar to lateral leaves in size and form. Usually erect in growth, with leaves transverse or ± incubous. Rhizoids usually in fascicles from underleaf bases
Group F
Anisophyllous to distichous: underleaves lacking or differing in form and size from lateral leaves, at most 0.1–0.5 (rarely to 0.55) lateral leaves in area
9
Rhizoids (usually readily located) clearly limited to ± sharply defined fields, at underleaf bases, or ventral lateral leaf bases (if scattered, leaves clearly opposed).Underleaves rarely wholly lacking; leaves alternate or opposed
Rhizoids (sometimes scarce) freely scattered (usually on normal, leafy stems), never fasciculate. Underleaves distinct varying to lacking; leaves clearly alternate
10
Underleaves distinct: usually at least 0.1–0.2 area of lateral leaves, most often bifid. Rhizoids always at underleaf (never leaf) bases; leaves usually alternate, opposite in some taxa
Group G
Underleaves lacking (or mere minute, few-celled vestiges, at most minute lanceolate or subulate segments, the underleaves in a few taxa small and lamelliform, these then with free slime papilla formation)
Group H
11
Leaves vertically oriented; their insertion nearly transverse (no more than 5–15°) succubous, as regards dorsal half of insertion). Leaves usually shallowly to conspicuously bifid, rarely entire and unlobed
Group I
12
Leaves bilobed, shallowly to deeply so (sometimes with accessory teeth; rarely sporadic leaves trilobed)
Group J
Leaves unlobed, never bilobed in vegetative regions (but sometimes conspicuously ciliate)
Group K
13
Underleaves of mature stems lacking; rhizoids scattered, or from lobule of leaf, or in small fascicles from stem (from loci where underleaf development was suppressed)
Group L
Underleaves present, conspicuous; rhizoids always at bases of the underleaves, from stem or from underleaf base
14
Underleaves unlobed, at most the apex toothed or lobulate
Group M
15
Plants without water-sacs or lobules: leaves simply complicate-bilobed (the dorsal lobe usually slightly to clearly, sometimes greatly, smaller than the ventral). Leaf lobes usually variously dentate or ciliate; marsupium or coelocaule developed at apices of leading shoots; coelocaule at most with a reduced perianth at apex
Group N
Plants developing distinct water-sacs (lobules), usually ± longer than broad, typically lying parallel to stem or weakly divergent
16
Lobules of stem leaves (and/or branch leaves) galeate, or helmet-shaped, longer than wide, only feebly attached to (and often remote from) stem; water sac formed only by the inflated sac-like lobule. Plants usually deep- or brownish green, or tinted reddish to purplish (only Lepidogyna pale yellowish green). Branching smooth, without basal collar (except sometimes in Frullania spp.)
Group O
Lobules (water sacs) of leaves formed by the infolded ventral lobe + opposed basal portion of the lobe, the lobules attached by an elongate keel to the lobe. Plants pale to yellowish to grass-green. Vegetative branches thecal, with a distinct basal sheath (except sometimes in Nephelolejeunea)
Group P
Group Keys
Group A. Mature vegetative plants essentially leafless and thalloid. Axis of 6 (or more) large cortical cell rows + many small inner cell rows; leaves reduced (to lateral 1- or 2-celled projections, each bearing a 1–2-celled papilla); rhizoids in fascicles from vestigial underleaves.
Only Zoopsis p. p. will key here.
Group B. Plants with leaves formed of 2–4 (wholly or largely) uniseriate, ciliiform lobes; disc lacking or less than 1 cell-row high.
Only Telaranea p. p. will key here.
Group C. Plants able to produce ventral-intercalary branches from axils of ± distinct underleaves (sometimes only ♂ and ♀ produced on ventral-intercalary branches). Plants usually regularly with leaves (3)4–12-lobed (rarely only lobulate), whose margins may be dentate or ciliate. Ventral (less often lateral) branches flagelliform or stoloniform in many taxa. Rhizoids always from underleaf bases, sometimes local or rare. (Leaf insertion variable:incubous, succubous, or transverse.)
Key
1
Gametangia (♀, usually also ♂) all or mostly on ± abbreviated ventral-intercalary branches that mostly lack normal leaves; microphyllous flagelliform axes usually produced. (Lepidoziaceae + Neogrolleaceae + Trichotemnomaceae)
Gametangia produced on leafy, elongated or leading shoots (♂ usually becoming intercalary; ♀ sometimes or often innovating if unfertilized). Microphyllous geotropic axes absent or present.(Plants usually tristichous, often ± isophyllous.)
4
Cells large, leptodermous, smooth; with lateral-intercalary branches
Paracromastigum (p. 463) + Hyalolepidozia (p. 472)
5
Leaves deeply (2)3-fid, with long-acute lobes; underleaves deeply 2(3)-lobed
6
Leaves transverse, hemispherically concave, apex irregularly (2)3–4-lobulate + dentate. Plants subisophyllous, the underleaves leaf-like; cells with coarse nodose trigones, the surface coarsely striate-papillose
7
Leaves 2–3-lobed for at least 0.5 their length, formed of rectangulate cells lacking trigones; lateral branching monopodial; plants subhyaline, pale green
Leaves shallowly (0)2–3-dentate or -lobulate. Branching pseudodichotomous (except Bazzania exempta); vigorous terminal branches laterally displacing main stem. Copious ventral flagella but lateral branches never flagelliform
8
Cells firm-walled, small, lacking oil-bodies; lateral branches all intercalary; plants with a system of basal, creeping, small leaved or microphyllous rhizomatous or stoloniform axes
Cells thin-walled, large, with oil-bodies; lateral branches terminal, Frullania type; plants lacking basal rhizomatous or stoloniform axes
10
Plants not dendritic, without a creeping caudex; lateral (vegetative) branches normally all Frullania type
Plants dendritic: a copiously branched erect shoot arising from a caudex, which freely produces geotropic flagelliform axes; lateral branches of Frullania + Microlepidozia types
Leaves (7)8–12(13)-lobed. Otherwise as in Telaranea but lacking a hyaloderm
12
Leaves (more or less) strongly asymmetrical, with dorsal sector larger than ventral; disc usually as high, or higher than lobes; cells firm-walled, ± opaque; stem lacking a conspicuous hyaloderm
Leaves ± symmetrical; disc usually under 0.5 height of leaf; cells thin-walled, ± rectangulate; stem with hyaloderm distinct
13
Leaves with lobes unarmed (except sporadically at base); cells lacking trigones; plants with perianths, a marsupium lacking
Leaves with lobes and disc copiously armed with pluricellular cilia/laciniae; cells with coarse trigones; plants with marsupia, a perianth lacking. Leaves asymmetrically 4-lobed, the dorsal lobe smaller
14
Leaves (5)6–8-lobed, each lobe ending in a hyaline, erect, finger-like cell. Oil-bodies absent; disc cells small, thick-walled; lateral branches all of Frullania type
15
Microlepidozia- type branching present; plants opaque, firm, dull green to fuscous; cells small, firm-walled
16
Leaves strongly asymmetrical, shallowly (3)4–5-lobed, dorsal lobe smaller; surface conspicuously papillose
Leaves symmetrically (3)4–6-lobed; surface smooth or slightly roughened or striolate
Leaves rigid, horny, polystratose; terminal branches lacking. Plants often blackish. Southern South America
[Herzogiaria]
19
Plants pale green; bracteolar antheridia present; perianth absent, a coelocaule present. Southern South America
[Vetaforma]
Plants brown to fuscous; bracteolar antheridia lacking; perianth conspicuous, a coelocaule absent. Branching malleable: Acromastigum-Microlepidozia-Frullania- type terminal + intercalary branches. Capsule wall 4–5-layered. Southern South America
[Pseudolepicolea]
20
Leaves usually over 0.5 (3)4-lobed, transverse to strongly succubous (if transverse, then with lobes ciliate and/or ending in setaceous lobes); leaves, if asymmetric, with dorsal lobe smaller. Plants lacking geotropic stolons; perianth conspicuous, androecial bracteoles lacking antheridia
Leaves usually under 0.35 2–3–4-lobed (or leaves incubous), the lobes/margins entire to toothed with broad-based teeth, never ciliate; leaves often asymmetric, then with larger dorsal lobe
Leaf cells with walls thin, the angles forming conspicuous trigones. Leaf lobes setaceous, rigid; perianth contracted to mouth; elaters 1-spiraled, the spiral broad
22
Perianth mouth wide open; leaves (at least near gynoecia) with opposed teeth and/or cilia
Perianth mouth strongly contracted; leaf (+ underleaf and bract) margins edentate/eciliate (at most 1 tooth of disc). Southern South America
[Archaeochaete]
23
Lateral branches all axillary, intercalary (rare exceptions aside); perianth conspicuous, long-emergent; ventral-intercalary stoloniform axes common; leaf cells shorter, the length:width under 2:1
Lateral branches of Frullania type; perianth vestigial, an Isotachis- type rigid perigynium present; ventral stolons usually lacking; leaf cells elongated
24
Non-julaceous; shoot tips usually cernuous; leaf cells under 4–5:1
Isotachis (vol. 2)
Julaceous, fragile, leaves very flimsy; shoot tips erect; leaf cells to 8–10:1
Eoisotachis (vol. 2)
Group D. Plants unable to form ventral-intercalary branches; both vegetative and gametangial axes lateral in origin (very rarely from terminal-ventral branches); plants predominantly, often regularly, 1–2- to 3–4-pinnately branched.Leaves variable; 3- or 4-lobed, sometimes bisbifid; never 5-lobed or more, sometimes strongly asymmetric; never forming geotropic, microphyllous stolons or flagella. Plants isophyllous to anisophyllous.
Key
2
Leaves transverse to succubous, never bisbifid, symmetric or subsymmetric, copiously armed with or dissolved into cilia; cells rectangulate in lamina, often tiered, lacking trigones, the surface ± roughened
Leaves somewhat to clearly incubous in insertion and/or orientation; cells not clearly rectangulate, never tiered, with walls firm (with coalescent trigones) or thin (with coarse, nodose trigones), the surface smooth
Perianth a low cap or urn, at summit of a distinct coelocaule; plants yellow-green. Never with ventral-intercalary branches
5
Leaf lobes caudate, the margins copiously armed with pairs of opposing cilia, the cilia repeatedly dichotomously branched; ventral-intercalary type branching lacking
Leaf lobes acute to lanceolate, the margins entire or with opposed spinescent teeth or cilia, the armature normally unbranched; ventral-intercalary type branching typically present
6
Branching usually 2–3(4)-pinnate, the branches never tapered, not cernuous; plants vigorous, prostrate to pendulous; Acromastigum- type branching never present
Branching 1-pinnate (primary pinnae very rarely with a short secondary branch), the pinnae tapered, often flagelliform, often cernuous; plants small (to 1 cm tall), erect or suberect; Acromastigum- type branches rare
7
Leaves symmetrically or subsymmetrically bisbifid, the lobes lanceolate to long-acuminate, entire or sporadically ciliate; plants greyish; perianth absent, a clavate, hispid coelocaule present
Leaves 3–4(5)-lobed, usually asymmetric (dorsal lobe largest); lobes triangular, rarely lanceolate, never acuminate, usually toothed and/or ciliate; plants brown to red-brown; perianth present, a coelocaule not developed
8
Lateral branches attenuate, becoming flagelliform; gynoecia all terminating weak lateral branches. Stem underleaves quadrifid; shoots strongly complanate, lax, often pendulous
Dendromastigophora (vol. 2)
Lateral branches never attenuate and flagelliform; gynoecia terminating main axes
Ptilidium (vol. 2)
9
Branching regularly 1–2(3)-pinnate, the branches freely developed; ultimate branches with leaves + underleaves fused in distinct rings; plants never gemmiparous
Herzogianthus (vol. 2)
Branching irregular, the branches sparing; both main axes and branches with leaves and underleaves free; plants with reddish brown stellate gemmae developed at shoot apices
Chaetophyllopsis (vol. 2)
Group E. Plants with leaves incubously oriented (and insertion usually at least weakly incubous), usually bilobed or bidentate (rarely unlobed). Rhizoids limited to bases of underleaves and/or geotropic stolons.
Key
1
Leaves never vittate, 0.05–0.55 bilobed (less often unlobed or 2–3-lobulate or 2–4-denticulate at apex); lobes, if distinct, never lanceolate and/or acuminate. Perianth bluntly trigonous, or lacking
Leaves vittate, 0.55–0.85(09) bifid, the vitta of elongate cells extended into each lobe; leaves often falcate-lobed, the lobes lanceolate to acuminate. Perianth fusiform, bluntly triplicate, mouth deeply 6-lobed
3
Branching all or mostly ventral-intercalary; gynoecia and androecia on short, ventral-intercalary branches; leaves unlobed, apices entire or bidentate
Branches normally lateral-terminal, Frullania type; gynoecia and androecia on leading leafy shoots; leaves irregularly lobulate. Southern South America
[Xenocephalozia]
4
Vegetative branches all lateral-intercalary. Leaves lepidozioid, usually 2(3)-lobed, few-celled; underleaves similar, usually smaller
Leaves with a distinct field of elongated, thin-walled ± inflated basal cells (in contrast to the small, firm-walled isodiametric distal cells); leaves strongly canaliculate. Leaf apices minutely (2)4-dentate
7
Stolons all axillary from underleaf axils; stem without hyaloderm, with thick-walled cortical cells in numerous rows; leaves (often obscurely) (2)3-dentate
Stolons all or many terminal, not axillary; stem with a firm hyaloderm, the cortical cells in 7 rows, at least on branches; leaves 0–2-lobed
8
Leaves planodistichous, unlobed or bidentate at apex. Leafy shoots giving rise to ventral-intercalary, geotropic flagelliform axes; leaf apex entire
Leaves concave, 2–3-lobed; leafy shoots giving rise to geotropic flagelliform axes
see Isotachis (vol. 2)
Group F. Plants usually erect or stiffly ascending, isophyllous or weakly anisophyllous; underleaves over 0.7 area of lateral leaves, usually similar in form, both unlobed or 2(3)-lobed or -dentate. Rhizoids only at underleaf bases (except Anthelia, p. p.). Leaves and underleaves mostly unlobed and edentate, sometimes bilobed. Plants either pigment-free, brownish, or fuscous; never reddish. Asexual reproduction uniformly lacking. (All on soil or rocks.)
Key
2
Cells with coarse nodular or lobed trigones, large (35–40 µm wide in middle). Leaves shallowly (under 0.25) 2–3–4-dentate or -lobulate; underleaves identical, unlobed, retuse or lobulate. (If leaves and underleaves unlobed, stiffly spreading, see Acromastigum.)
Cells thin- to firm-walled, at most with large but concave-sided trigones, smaller (median under 34 µm wide)
3
Leaves 2(3)-lobulate; underleaves spoon-shaped, unlobed; ♂ and ♀ gametangia on leading leafy axes; branches only lateral-intercalary; surface smooth. Southern South America
[Pleurocladopsis]
Leaves variably 3–4-lobulate + dentate distally; underleaves similar; both almost hemispherical; ♂ and ♀ gametangia on short, axillary branches; branching malleable, ventral- + lateral-terminal + intercalary; surface strongly striate-papillose
5
Leaves (isolated exceptions aside) bilobed or bidentate; wall pigments absent, or brown to reddish brown; cells never strongly elongated
Leaves ± oblong, variably and shallowly 2–3–4-lobed; underleaves similar but 2-lobed; fuscous; cells strongly elongated, thin-walled, to 8–10:1 basally. Plants julaceous subaquatic, lax
Eoisotachis (vol. 2)
6
Leaf with surface covered by large, hyaline, dome-shaped thickenings over cell wall intersections (not over lumina). Plants tiny, pale or whitish green, with spreading leaves and underleaves at most subcontiguous; rhizoids at underleaf bases only; branches freely terminal, Frullania type; perianth trigonous. Southern South America
[Pigafettoa]
Leaves smooth or minutely papillose. Leaves contiguous or imbricate (stem usually hidden), erect or suberect
8
Leaves and underleaves identical, both over 0.55 bifid, with smooth margins; ♂ and ♀ gametangia on leading axes (ours paroecious); leaves with age often becoming decolorate and whitish
Leaves 2(3)-fid, underleaves bifid, to 0.35, the margins ± denticulate; ♂ and ♀ on short ventral-intercalary branches (dioecious); leaves not becoming decolorate and whitish
9
Leaves cuneiform, transverse to weakly succubous, bifid to 0.5–0.75, the lobes abaxially canaliculated below; perianth present, conspicuous; capsule wall 3–4-stratose; terminal branches sometimes present, at times frequent
Leaves oblong-ovate (at times narrowly so), transverse to weakly incubous, bifid to 0.35–0.5, the lobes plane; perianth lacking, the innermost gyre of large bracts + bracteole appressed to calyptra; capsule wall 2-stratose; terminal branches lacking. Southern South America
[Grollea]
10
Plants filiform, with remote ovate-triangular leaves and, optimally similar, ± triangular underleaves; rhizoids scattered; no stolons. All branches lateral-intercalary. Prince Edward Is.
[Invisocaulis]
Medium-sized, with contiguous to imbricate, ovate-orbicular leaves and underleaves; rhizoids at underleaf bases; with stolons. Some ventral-intercalary branches
11
Plants bright green; ♂ and ♀ gametangia on short, leafless intercalary branches; leaves never stiffly spreading, basally strongly concave
Plants deep green to olive-green to almost black; gametangia on ± elongated, leafy axes; leaves rigid, ± spreading, not or little concave
Pachyglossa (vol. 2)
12
Branching exclusively intercalary, lateral and ventral; cells, all or most, lacking oil-bodies; stem with hyaloderm lacking; leaves not cucullate
Branching frequently terminal; cells with oil-bodies; stem with a hyaloderm surrounding smaller, firm-walled intracortical cells; leaves cucullate
Group G. Plants anisophyllous to almost distichous, with leaves alternate to (at most) subopposite, succubous or transverse, 0–2-lobed (rarely 2[3]-lobed).Underleaves usually under 0.5 (rarely 0.5–0.6) leaves in area (if, rarely, larger, very different from leaves in shape). Rhizoids in fascicles: from bases of underleaves, never scattered (except in a few taxa with sharply opposed leaves). (An artificial group, including taxa of Lepidoziaceae, Geocalycaceae, Lophocoleaceae, Jungermanniaceae.)
Key
1
Stem, in cross section, with a conspicuous hyaloderm (medullary cells usually distinct by transmitted light as an opaque central strand); cortical cells 1.5–4× medullary in diam. or even larger
Stem without a perceptible hyalodermis; cortical cells thin- or firm-walled, never conspicuously larger than intracortical in cross section
2
Leaves subhorizontally or obliquely oriented; cells large, hyaline, leptodermous; hyaloderm very conspicuous; underleaves reduced, under 0.2 area of leaves, few-celled, bifid. Leaves with a symmetrically bilobed, rectangulate lamina, 4 or more cells broad, conspicuously bifid (Lepidoziaceae)
Leaves transverse or almost so, vertically oriented; cells firm, small, with ± distinct trigones; hyaloderm poorly demarcated, the enlarged cortical cells thick-walled; underleaves to 0.5(0.65) area of leaves
4
Leaves with lobe apices sharp, persistent; sinuses descending under 0.25 leaf length
Leaves with long-tapered apices, of moniliform cells, fragmenting; sinus descending over 0.25 leaf length
5
Leaves entire or shallowly 2(3)-lobed/lobulate, their surfaces smooth; ventral-terminal branches present
Leaves to 0.5 2(3)-fid, covered by coarse, convex, lenticular thickenings over cell wall intersections; ventral branches only intercalary. Southern South America
[Pigafettoa]
6
Ventral-intercalary microphyllous stoloniform axes absent. Acromastigum- type branching absent; plants usually large, with collenchymatous leaf cells; leaves 0–2-lobed and underleaves 0–2-lobed, or with lateral teeth or laciniae large enough to simulate lobes
Ventral- and/or lateral-intercalary, microphyllous-rhizoidous, usually strongly geotropic branches freely developed (often axillary from lower leaves or leafy shoots) and/or with a creeping, rhizoidous caudex
8
Perianth conspicuous; no marsupium; surface smooth or with solitary papillae (plants never conspicuously dull when dry)
Perianth lacking; a marsupium developed; surface densely covered by papillae (plants diagnostically dull when dry)
Saccogynidium (vol. 2)
9
Plants whitish to pure green; perianth trigonous
Chiloscyphus (p. p.) (vol. 2)
Plants brown to fuscous; perianth bilabiate, laterally compressed at least above
Leptoscyphus (p. p.) (vol. 2)
10
Asexual reproduction lacking, or via gemmae, or via caducous leaves, cells with few (under 12) oil-bodies (the oil-bodies at times botryoidal)
Asexual reproduction via caducous or fragmenting teeth of distal portions of the bilobed leaves; cells leptodermous, each with numerous (20–40 or more) minute, finely papillose oil-bodies. Leaves flat, neither with convex dorsal margin, nor concave, even basally; plants maximally flattened, delicate, fragile; Frullania- type branches freely formed; usually sterile
Leptophyllopsis (vol. 2)
11
Plants with leaves ± constricted at base, freely caducous, the mature plants extensively denuded. Plants brownish; perianth laterally compressed at the closed mouth; branching mostly intercalary; leaves ± obdeltoid or obovate. Southern South America
[Anomylia]
12
Plants strongly laterally compressed, with broadly rounded leaves. Underleaves free, minute, divided to near base into 2–3 linear segments, the flanks unarmed; perianth 2-lipped, compressed; rhizoids in part scattered. Southern South America
[Pedinophyllopsis]
13
Perianth 2-lipped, laterally compressed at least above (with broad, bilobed and/or toothed lateral faces, the ventral, third lobe, narrowed, usually abbreviated, often infolded)
Perianth symmetrically 3-sided (ventral face as broad as lateral faces) or inflated and unkeeled, the mouth equally 3-lobed (each lobe often again bilobed)
14
Leaves 1-stratose throughout; dorsal-intercalary branching never present; underleaves broader than stem, the lamina margins usually conspicuously armed
Leaves, except marginally, 2–3-stratose; dorsal-intercalary branching (Andrewsianthus -type) present; underleaves narrower than stem, small, the lamina margins entire or with 1(3) small teeth. Leaves unlobed, entire. Southern South America
[Evansianthus]
15
Plants green, chiloscyphoid; perianth with longitudinal, ± undulate, toothed wings, in addition to the 3 major keels
Lamellocolea (vol. 2)
Plants brown to fuscous; perianth surfaces unarmed (at most keels ± winged)
Leptoscyphus (vol. 2)
17
Gametangial branches all highly reduced; both ♂ and ♀ branches short, the ♂ spicate, compact. Cells typically with coarse, often nodose trigones
Heteroscyphus (?incl. Amphilophocolea) (vol. 2)
Gametangial branches unspecialized; androecia and gynoecia on (usually) elongated, or if short, leafy branches. Cells with usually thin walls, with small or very small trigones
Chiloscyphus (vol. 2)
18
Leaf insertion, including at dorsal end, clearly succubous, leaves obliquely oriented (concavity turned up, or toward shoot apex); leaves usually decurrent dorsally; ♀ bracteole usually smaller than bracts; perianth usually inflated, convex-sided
Clasmatocolea (vol. 2)
Leaf insertion dorsally transverse or feebly incubous, the leaves incubously oriented (concavity turned ± toward ventral face of plant); leaves non-decurrent; ♀ bracteole subequal to bracts in size, obovate, large; perianth faces flat or concave. Southern South America
[Xenocephalozia]
19
Leaves deeply (usually 0.3–0.65 or more) 2- or 2–3-fid, formed of ± elongated, thin- or equally firm-walled, trigone-free cells; underleaves 2-, 2–3-, or 3-fid; Acromastigum- type branches usually present
Leaves unlobed (rarely retuse or emarginate), entire-margined or 2- to 3–4-denticulate at apices; Acromastigum- branches absent
21
Leaves succubous, never fleshy; dorsal-intercalary branches present, from dorsal end of leaf axils; plants closely prostrate. Southern South America
[Evansianthus]
Leaves vertical, transverse or weakly succubous, fleshy; dorsal-intercalary branches lacking, the lateral branches clearly axillary in leaves; plants erect in growth
22
Rhizoids purplish. Plants chestnut-brown, branching typically all ventral-intercalary, leaves polished and nitid, strongly dorsally assurgent
Acroscyphella (vol. 2)
Leaves chiloscyphoid, with deflexed dorsal leaf margins, the leaves not adaxially concave
Stolonivector (vol. 2)
24
Leaves clearly to closely imbricate, hiding stem from view, elongated, shallowly 2(3)-lobed or 2–4-denticulate at apex, vertical, transversely inserted at dorsal ends; leaves with median-basal or at least basal cells conspicuously larger; oil-bodies lacking
Leaves remote or contiguous, saucer-shaped, strongly concave, rounded, unlobed, subvertical, dorsal insertion ends oblique; leaves lacking an area of enlarged cells in basal sector; oil-bodies (1)2–3 in some/many cells. Underleaves very inconspicuous, appressed to stem
Hepatostolonophora (vol. 2)
25
Leaves spreading, naviculariform, the shoot complanate; leaf tips narrowly 2–4-lobulate/dentate
Leaves suberect, closely imbricate, the shoot (more or less) julaceous; leaf apices shallowly 2(3)-lobed, with broad sinuses
Group H. Plants like those of Group G, but essentially distichous; ventral merophytes narrow, underleaf-free, or with minute, subulate vestiges, or—at most—small lamellae with many small cilia or papillae. Leaves alternate or exactly opposed, unlobed, or 2– 4-lobed, entire-margined or variously toothed and/or ciliate. Rhizoids either at the bases of the underleaf vestiges or at ventral leaf bases, almost never scattered (or leaves 3–4-lobed). (Artificial group including taxa of Lophozioideae, Plagiochilaceae, Jackiellaceae, Acrobolbaceae, Lophocoleaceae.)
Key
2
Stem without a hyalodermis. Leaf cells usually ± clearly collenchymatous, with coarse trigones, with 1–few, large granular oil-bodies per cell; rhizoids scattered or often in fascicles at ventral leaf bases
Stem with a ± distinct hyalodermis
see Group G
4
Leaf insertions not extending to stem midline dorsally, a leaf-free strip present; plants usually deep brown to fuscous; ♂ and ♀ on short ventral-intercalary branches lacking normal leaves
Leaf insertions extending to stem midline dorsally, a leaf-free strip lacking; plants lacking secondary pigments or, if present, then reddish or purplish or (Austrolophozia) with distal parts of leaves in sun reddish brown or golden brown; ♂ and ♀ on leading shoots
5
Secondary pigments lacking; leaf surface conspicuously papillose (smooth in A. concinnus); rhizoids often confined to ventral base of leaves, not forming a ventral mat; ♂ bracts moderately concave to ventricose at base, the antheridia covered by the bracts
Acrobolbus (vol. 2)
Secondary pigments present, reddish or purple brown or (Austrolophozia) tinged with reddish brown or golden brown in distal parts of leaves; leaf surface (lamina) smooth; rhizoids in a dense ventral mat forming a decurrent bundle; ♂ bracts leaf-like, the antheridia exposed
6
Leaves unlobed; plants with reddish or purple-brown pigments; gemmae lacking; marsupium very long and slender, buried in substrate
Goebelobryum (vol. 2)
Leaves (2)3–4(5)-lobed; plants in sun with reddish brown or golden brown pigments in distal parts of leaves; gemmae present; marsupium conoidal, elevated above substrate
Austrolophozia (vol. 2)
8
Vegetative branches lateral-intercalary and/or of Frullania type, never ventral-intercalary
Plagiochila (vol. 2)
Vegetative branches all ventral-intercalary, never lateral-intercalary. Leaves ± erect, laterally appressed
Acrochila (vol. 2)
9
Rhizoids (sometimes infrequent) always scattered; perianth inflated, plicate, ± narrowed to mouth; asexual reproduction common
Rhizoids frequent, in fascicles at bases of reduced or minute underleaves; perianth wide at mouth, 3-lobed (ventral lobe sometimes narrower); asexual reproduction absent
10
Leaves (in ours) 2–3-lobed, the 2–3-lobed leaves in about equal frequency, the lobing at times asymmetric, the dorsal-most lobe smaller and shorter; ventral leaf base entire, without cilia; underleaves lacking
Leaves ± symmetrically (3)4-lobed; ventral leaf base with 1–several cilia; underleaves distinct. Southern South America
11
Leaves mostly convex, 2(3)-dentate at apex, their insertion lines not attaining dorsal stem midline (with a “leaf-free” strip); perianth with third, ventral face narrow. Plants strongly dorsiventrally flattened
Pseudolophocolea (vol. 2)
Leaves concave, often strongly so, usually unlobed or edentate, sometimes pluridentate, the insertion dorsally attaining stem midline; perianth trigonous or inflated, the ventral face not or little narrowed
Clasmatocolea (p. p.) (vol. 2)
12
Perianth sharply laterally compressed; ♀ bracts free from bracteole, leaf-like; plants usually brownish
Plagiochilion (vol. 2)
Perianth not or weakly laterally compressed, contracted to mouth; ♀ bracts + bracteole united, often forming a sheath; plants often purplish or reddish, or reddish brown
13
Leaves erect-appressed, reniform-orbicular, the plant strongly laterally compressed; branching mostly lateral (intercalary + Frullania type), rarely ventral-intercalary. Cortical cells, like medullary, thin-walled. Southern South America
[Protosyzygiella]
Leaves loosely to strongly laterally patent, plants not laterally compressed; branches all or predominantly ventral-intercalary. Cortex 1–3-stratose, of thick-walled cells. [Kerguelen]
[Syzygiella]
Group I. Plants with leaves transversely oriented (vertical), always alternate, usually bilobed (less often unlobed), the dorsal half of insertion lines within 5° of being transverse (sometimes decurrent). Underleaves usually very small (under 0.3 leaf area) or none. Perianths distinct (except in Gymnomitrion); cells with oil-bodies (except in Metahygrobiella, which lacks stolons).
Key
2
Leaves remote to contiguous to imbricate, never tightly appressed; leaf perimeters, in situ, easily evident; shoots not filiform-julaceous. Cells never armed with high lenticular papillae
Leaves remote, scale-like, bifid, closely appressed, often almost unresolvable; shoots filiform-julaceous, tiny
3
Leaves clearly bifid, the transverse dorsal ends interlocking; perianths included, short, or lacking. Plants freely stoloniferous; cells with 2–3 granular oil-bodies. (Gymnomitriaceae, p. p.; all alpine)
Leaves, usually bifid or bilobed, with dorsal insertion lines extending to, but not beyond, stem midline; perianth free, conspicuous, externally evident
4
Perianth well developed, sheathed closely by the erect bracts, without a perigynium at its base; plants procumbent, with stiffly spreading leaves with abaxially sulcate lobes; surface coarsely papillose. Autoecious
Perianth absent or a short tube hidden within the erect bracts; plants erect (ours usually fuscous to piceous), with leaf lobes not abaxially sulcate; surface smooth or weakly papillose.Leaves with margins flat or nearly so; branches all from middle or lower ends of leaf axils; gynoecium with a low perianth + distinct perigynium
5
Stem lacking a hyaloderm; cortical cells usually firm-walled, no larger than medullary; cells with oil-bodies present
Stems subhyaline, with (at least dorsally) a hyaloderm formed of few rows of large, hyaline cells surrounding a medulla of (usually) firmer-walled, mostly smaller cells; with stolons; leaf cells thin- or equally firm-walled, usually 18–38 µm wide; oil-bodies lacking. Underleaves absent, but gynoecia tristichous
6
Cells minute or very small (usually 9–18 µm broad), usually thick-walled and lacking trigones; plants small or minute, with small, usually remote leaves. Terminal branches absent or rare
Cells larger: normally 17–35 µm wide or more in leaf middle, usually with conspicuous or nodose trigones; plants usually relatively large; over 600 µm wide with leaves. Leaf lobes sharp (leaves all bilobed); many or all branches intercalary, lateral or dorsal; plants mostly ± brownish or infuscated to purplish, relatively rigid
7
Underleaves distinct, to 0.5 area of lateral leaves; perianth bluntly trigonous; gemmae lacking. Seta with 8+4 cell rows; otherwise Cephaloziella -like. Southern South America
[Amphicephalozia]
Underleaves lacking or very small (rarely larger in Cephaloziella; this with a 4+4-seriate seta); perianth 4–5-plicate or at apex of a perigynium; gemmae present, 2-celled, elliptical (but infrequent or lacking in most species of Allisoniella). Seta of the 4+4 or 8+4 type
8
Leafy stems ± procumbent, not arising from a caudex, leaf lobes 2–9(12) cells broad; minute plants, with non-sulcate leaf lobes; seta with 4+4 cell rows
Leafy stems erect, arising from an extensive caudex; leaf lobes 15–25 cells wide, usually abaxially sulcate; small but not minute, fuscous or purplish plants, with rigid stems; seta of the 8+4–12 type
9
Shoot apices not running out into microphyllous, pigmented stolons; shoot tips never abruptly geotropic; branches Frullania type and/or lateral-intercalary, not displaced to dorsal end of leaf insertion. Gemmae sometimes present; leaves always bifid
Shoot tips of normal leafy stems running out into microphyllous, often geotropic (or strongly arched) flagelliform axes that are often pigmented; Frullania- type branches usually lacking, the branching in most taxa with intercalary branches from dorsal side of stem. Gemmae never present; leaves 2-lobed, rarely 0- or 3-lobed
11
Shoot apices not abruptly geotropic and microphyllous; branches never from dorsal end of leaf axils. Leaves dorsally interlocking; plants slender, julaceous and ± suberect, with leaves unlobed or to 0.35 bilobed; perianth absent or short, usually included; gemmae lacking; underleaves lacking or vestigial; cells firm-walled
Shoot apices abruptly geotropic and becoming microphyllous/leafless; branches from dorsal end of leaf insertions. Surface coarsely papillose
Perianth none; innermost bracts variable, reduced, ± scale-like, hidden within larger outer bracts
13
Leaves crenulate with thin-walled, delicate cells, or smooth-margined; perianth pluriplicate, not compressed; dioecious
Leaves irregularly denticulate with knob-like, thick-walled projecting marginal cells; perianth hardly plicate, narrowed and laterally compressed toward mouth; autoecious
Group J. Plants with leaves succubous, non-complicate, alternate and bilobed (margins edentate or dentate); rhizoids scattered, usually frequent (sometimes none on upper shoot sectors). Underleaves usually small or lacking. (Artificial group, including Lophozioideae, Acrobolbaceae, Cephaloziellaceae, Cephaloziaceae, etc.)
Key
2
Underleaves rectangulate, deeply bifid, lacking lateral cilia or teeth; perianth lacking, a subterranean, rhizoidous marsupium developed. Branches ventral-intercalary only; plants opaque, yellowish green
Geocalyx (vol. 2)
Underleaves undivided and lanceolate, or 2- or 3-fid, if divided, with ± cilium formation; perianth distinct, without a marsupium
3
Dorsal leaf margins not deflexed; leaf margins (except basally) edentate. Plants lacking stolons or flagella; perianth terete, plicate and contracted above
Dorsal leaf margins ± deflexed (a cnemis developed); leaf apices and/or margins usually toothed or shallowly bilobed. Androecia spicate, tapered
Leaf surface with papillae low or none, the stem smooth. Trigones not knot-like; ♂ bracts neither laciniate nor reduced in size; rhizoids all dispersed; ventral-intercalary branches lacking; leaves 0.25 or more bilobed; often with gemmae
5
Trigones coarse, knot-like; stems paraphyllose; perianth plicate to base, not twisted; ♀ bracts not reduced
Trigones not nodose; stems smooth; perianth plicate above only, the plicae twisted; ♀ bracts reduced in size, laciniate. Southern South America
[Roivainenia]
6
Perianth laterally compressed, wide at mouth; leaf apices ± toothed; underleaves free, usually ciliiform or ciliate
Plagiochila (vol. 2)
Perianth plicate, narrowed above; leaf apices to 0.2 bilobed, edentate; underleaves connate with leaves on one side, lanceolate. Southern South America
[Nothostrepta (p. p.)]
8
Perianth lacking, a pendulous marsupium developed; branching wholly intercalary; plants green to yellow-green, the stems whitish with age. Surface usually rough; oil-bodies large, often opaque
Perianth present, a marsupium lacking; branching variable; plants not opaque green above and whitish on older sectors
9
Plants creeping, with rhizoids on leading leafy axes, stem under 14 cells in diam., without a distinct cortex, a system with a leafless/microphyllous caudex and stolons lacking; antheridia 1(2) per bract
Acrobolbus (vol. 2)
Plants with ascending or erect leafy shoots from a leafless/microphyllous pale caudex, often with stolons or flagella, with rhizoids restricted to leafless axes below, the stem 16 cells or more high, with ± distinct cortex; antheridia 2–15 per bract
10
Gametangial branches usually abbreviated, then often whitish, from lower ends of leafy shoots; ♀ bracts often mere inconspicuous scales; antheridia 2–4(5)
Marsupidium (p. p.) (vol. 2)
Gametangia on leading, leafy, erect stems; ♀ bracts large, leaf-like, the innermost ones distinct; antheridia 5–15
Tylimanthus (p. p.) (vol. 2)
11
Perianth wide at the laterally compressed mouth; leaves often with a cnemis, the dorsal margin abruptly decurved; leaf margins and/or lobes usually sparingly to copiously dentate, the teeth usually sharp, spinose. Leafy shoots from a rhizoidous basal caudex
Perianth not laterally compressed; leaves never with a cnemis; leaf margins usually edentate. Perianth plicate, moderately contracted to mouth; seta of the general type: at least 6–8 cells in diam.
12
Leaf surface mostly smooth, at most finely papillose, the papillae low
Plagiochila (vol. 2)
Leaf surface coarsely papillose, the papillae dense, hemispherical, distinctly protruding
Dinckleria (vol. 2)
13
Leaves with ventral margins not conspicuously decurved; seta reduced, of 4 or 8 outer + 4 inner cell rows
Leaves with ventral margin strongly deflexed, leaf apices shortly bifid; seta ca. 10 cells in diam. Southern South America
[Nothostrepta (p. p.)]
14
Dorsal end of leaf insertion essentially transverse, extended to stem midline dorsally; gemmae usually lacking; perianths on leading axes
Dorsal end of leaf insertion oblique, not extended to stem midline dorsally; gemmae often present; perianths usually on abbreviated branches
Group K. Leaves with insertion (and usually orientation) perceptibly succubous, the adaxial lamina usually ± turned toward the observer; leaves alternate, unlobed, edentate or variously dentate/ciliate; rhizoids always scattered. Branches normally all intercalary. Most taxa with vestigial or no underleaves.
Key
1
Cells on abaxial surface of leaf with large dome-like to mammilliform thickenings; gametangia on abbreviated lateral-intercalary branches or on leading shoots. Perianth present
Cells smooth or, if papillose, without dome-like thickenings associated with each cell; gametangia on abbreviated ventral-intercalary branches. Plants suberect to erect
2
Surface of leaf with large dome-like to mammilliform thickenings each positioned over the cell lumen, the dome-like thickenings with dense hemispherical to clavate papillae; gametangia on abbreviated lateral-intercalary branches; plants prostrate and loosely creeping, typically corticolous (very rarely saxicolous)
Brevianthus (vol. 2)
Surface of leaf with large dome-like thickenings each positioned over the radial walls (not over the cell lumen), the dome-like thickenings smooth; gametangia on leading shoots; plants erect, in cliff crevices or over soil
Bragginsella (vol. 2)
3
♂ and ♀ Branches reduced in size, abbreviated, normally all ventral-intercalary (usually produced only from older, lower sectors of, often, small-leaved or microphyllous axes). Leaf cell trigones lacking to large, rarely nodose
♀ + ♂ Gametangia not on reduced branches, the ♀ terminal on long leafy shoots, the ♂ becoming intercalary on leafy shoots. Branches often of lateral-intercalary and Frullania types, usually not exclusively ventral-intercalary
4
Perianths lacking; a pendent marsupium developed, or an erect, fleshy perigynium on weak ventral-intercalary branches. Androecia small, spicate, ± determinate; vegetative branches all or mostly ventral-intercalary
Perianths developed, usually conspicuous, trigonous. Plants erect, the leafy shoots smaller-leaved below, arising from a creeping, microphyllous, rhizoidous caudex; underleaves lacking or vestigial; leaves unlobed, bidentate or pluridentate; seta massive, over 6 cells in diam.
5
Plants green to fuscous, but not pitch black, the leafy shoots erect; with leafy shoots arising from an extensive system of plagiotropic and geotropic leafless axes; trigones ± concave-sided
Plants brown or black, creeping and forming compact mats, the shoots all leafy, without a small-leaved caudex (but geotropic stolons often present); trigones nodose. Gynoecia producing marsupia
6
Gynoecium producing an erect, fleshy, ± clavate calyptral perigynium, invested beneath by an isomorphic ring of bracts + bracteole
Gynoecia producing a pendent marsupia, at summit with vestigial bractlets. Plants green, older sectors (more or less) ivory white
Marsupidium (vol. 2)
7
Stem to 25 cells in diam,, with a 2–3-layered firm-walled cortex; plants small, usually olive-green to brownish; gemmae often present. Capsule wall 4–6-stratose
Adelanthus sect. Calyptrocolea (p. 616)
Stem to 35–40 cells in diam., with a collapsing hyaloderm surrounding thick-walled cells within; plants vigorous, pure green; gemmae lacking. Capsule wall 6–7-stratose; leaves copiously toothed
8
Leaf apices entire or variously toothed, never bidentate; stems usually fuscous pigmented, darker than the leaves, the cortex 2–3-layered, the cortical cells very thick-walled and with greatly reduced lumina; plants often developing gemmae
Leaf apices (in ours) with 2 strong spiniferous teeth; stems unpigmented (except P. aureocinctum), usually not darker than leaves, the cortex in 1(2) layers, the cortical cells at most moderately thick-walled; plants lacking gemmae
9
Leaves narrow, elongated, acutely pointed, edentate, the summit long-protracted. Perianth terete, plicate; leaves spreading, in drying dorsally ± secund; ♂ bracts with lobule long-acuminate distally
12
Leaves inserted dorsally to the stem midline, or beyond, a leaf-free dorsal strip lacking; cortex well defined, medulla mycorrhiza-free; caudex and/or stolons freely formed; leaves mostly dentate/ciliate; dioecious
Plagiochila (vol. 2)
Leaf insertion not attaining stem midline dorsally; cortex scarcely distinct, medulla mycorrhizal; caudex and stolons lacking; leaves unlobed or retuse, edentate; autoecious
Pedinophyllum (vol. 2)
13
Branches all, or predominantly, terminal-lateral, or axillary (in middle or ventral halves of leaves; gynoecia with bracts unlobed, edentate, without accompanying bracteole; plagiotropic caudex lacking, but some taxa with stolons. Basal perigynium usually at least weakly developed (strongly developed in some taxa); perianth with beak present or absent, if present, then the rostellum never depressed and situated at the truncate summit of the perianth; perianth mouth crenulate or entire; plants nearly always developing reddish or vinaceous pigments; leaves various, mostly orbicular or ovate; underleaves wholly absent, even in gynoecial regions
Branches all, or predominantly, intercalary, from ventral end of leaf axils or ventral (in some taxa some terminal-lateral, Frullania -type branches developed); gynoecia with bracts + bracteole usually variously lobed and/or incised, often reduced in size; plagiotropic microphyllous axes and stolons usually frequent (except in Jamesoniella)
14
Leafy shoots bearing or arising from a caudex and/or stolons; terminal branches usually absent. Trigones not coarsely nodose, not bulging and prismatic; perianth mouth crenulate or entire; plants nitid, deep brown to red-brown
Leafy shoots not arising from a caudex; stolons lacking; terminal, Frullania- type branches present
Leaves opposite
see Couplet 12, Group H
Leafy shoots erect or stiffly ascending, lacking rhizoids except on small-leaved bases and caudex
Tylimanthus (vol. 2)
17
Stem 15 or more cells in diam.; ♂ bracts with 3–9 or more antheridia; marsupia slenderly cylindrical, subterranean
Stem slight: ca. 8 cells in diam.; ♂ bracts 1-androus; marsupia conoidal, not penetrating substrate. Leaves round, edentate, concave
Acrobolbus subg. Lethocoleopsis (vol. 2)
18
Leaves edentate; plants ± flat, green to warm brown; gemmae often present
Lethocolea (vol. 2)
Leaves (more or less) toothed, rarely entire; plants julaceous, the leaves strongly concave, ± purplish pigmented; gemmae absent
Goebelobryum (vol. 2)
Group L. Plants with leaves complicate-bilobed (often abaxially winged) and/or with elaboration of a ventral, specialized water sac. Underleaves lacking.
Key
1
Rhizoids either scattered on stem or in bundles at ventral bases of leaves; infra-axillary branches lacking
Rhizoids either in bundles from the ventral lobe (lobule) of the leaf or occurring in small groups (2–few) at intervals on the stem; branches all infra-axillary, arising posterior to a leaf. (Taxa with strongly complicate-bilobed leaves, with ventral lobe smaller, folded under the larger, incubous, dorsal lobe. Usually yellow-green or green; often with asexual reproduction.)
3
Keel elongated, 0.35–0.5 leaf length, the leaf spatulate, U-shaped in section; cells with trigones coarse (often confluent) in distal parts of leaf, at base narrowly rectangulate, forming a vitta; gemmae and underleaves lacking
Keel short, 0.15–0.35 leaf length, sharply folded and complicate, not spatulate; cells thin- to equally firm-walled, the leaf lacking a distinct vitta; gemmae on erect, microphyllous shoots with underleaves
4
Perianth lacking, a coelocaule present; dorsal lobe ± larger than ventral (if not, a wide wing separating the two); rhizoid apices often digitately branched; asexual reproduction absent
Perianth distinct, plicate and/or flattened dorsiventrally; dorsal lobe 0.25–0.7(0.85) area of ventral lobe; rhizoid apices never digitate; asexual reproduction via catenulate gemmae
5
Dorsal lobe flat, slightly larger than ventral, broadly margined with vitreous, hyaline cells; monandrous. New Caledonia
[Perssoniella]
Dorsal lobe usually convex, not larger than ventral, not margined with hyaline cells; polyandrous
Paraschistochila (vol. 2)
6
Leaves with keel U-shaped, the leaves basally spatulate, the keel at most sharp near its apex; gemmae polygonal or stellate, or lacking; perianth plicate, moderately compressed
Leaves sharply folded in keelar region, the bases not sheathing, the keel usually ridged or winged (at least distally); gemmae usually ellipsoidal, smooth; perianth usually strongly compressed, smooth
7
Leaf lobes unequal: the dorsal at most 0.75 the ventral in size; leaf lobes non-squarrose, or dorsal squarrose; plants green to brown, sometimes with red pigments, never piceous. Gemmae present, stellate or polygonal
Diplophyllum subg. Diplophyllum (p. 747)
Leaf lobes subequal, both strongly squarrose; plants often dark brown to fuscous, opaque
Diplophyllum subg. Austrodiplophyllum (p. 747)
8
Rhizoids lacking from stem, arising as small fascicles from the ventral leaf lobes (lobules); perianth strongly dorsiventrally flattened, wide and truncate at mouth
Radula (vol. 3)
Rhizoids occurring isolated, or in bundles of 2–few, at regular intervals on stem (at points where underleaf formation has been suppressed); perianth 5-angled or ± compressed, but always contracted abruptly into a small beak
9
Branches all with a low basal collar; gemmae usually intramarginal; leaves never sharply dimorphic
Cololejeunea (vol. 3)
Branches athecal, smoothly joined to stem; gemmae usually marginal; leaves dimorphic: some lobulate, others elobulate
Aphanolejeunea (vol. 3)
Group M. Plants strongly complanate, with leaves complicate-bilobed, with 1 (2 in Goebeliella) smaller, inflexed ventral lobe(s), hidden in dorsal aspect of plant, the lobules ± transformed into a water sac (except Porella). Underleaves large, conspicuous, unlobed; branching only lateral; rhizoids only at underleaf bases. Perianths (but no other perigynoecial structures) developed. (Lejeuneaceaesubfam.Ptychanthoideae + Porellaceae + Goebeliellaceae.)
Key
1
In ventral aspect, a single lobule visible; leaf cells in distal part of lamina lacking hyaline, perforated cells
In ventral aspect, each leaf is provided with 2, curved, horn-like, inflated lobules; leaf cells, distally, dimorphic: some hyaline and perforate; at base leaves with linear cells (Goebeliellaceae)
Goebeliella (vol. 3)
2
Lobule oriented nearly parallel to stem, not converted to a water sac, lying at nearly right angles to lobe, with margins reflexed and organs all on short lateral branches
Porella (vol. 3)
Lobule with longer axis oriented parallel to lobe, united to it by a long keel, inflated at least along keel, with free margins never reflexed and organs (in ours) always on elongated branches or at apices of main stems. (Lejeuneaceae subfam. Ptychanthoideae)
3
Perianths never sharply 5-carinate: with 3 plicae or keels, or with 4 winged or armed keels, or up to 8–10 plicate. Usually deep brown to blackish
Perianths sharply 5-carinate (dorsal keel sometimes low and short), the carinae smooth. Plants dull, deep green to brown; with 1–2 innovations; oil-bodies many (in ours to 20–40 per cell), small, few-segmented; stem lacking a distinct hyaloderm of enlarged cells. (Our spp. dioecious.)
Archilejeunea (vol. 3)
4
Stem with cortex of thick-walled cells in many rows (underleaves attached to stem by many cells). Cells with 2–8(10) large, botryoidal, opaque oil-bodies. Leaves and underleaves often dentate. Subfloral innovations present
Stem with cortex of thin- or slightly thick-walled cells (diam. superior to that of medullary cells), in few, usually 12–16 rows (underleaves attached by usually 2–4 ventral cortical cells). Oil-bodies numerous, very small, homogeneous and glistening, or few partite with age. Leaves and underleaves edentate. No subfloral innovations
5
Branching regularly, thecal (Lejeunea type); perianth 3-carinate
Mastigolejeunea (vol. 3)
Branching all or mostly athecal (Frullania type); perianth pluriplicate
Ptychanthus (vol. 3)
6
Plants golden to inky black, glossy, nearly flat when dry and when moist; gynoecia with toothed bracts; perianth flattened, with 4 winged (and sometimes armed and/or lamellate) keels; underleaves orbicular to reniform; trigones, if distinct, never cordate. (Dioecious or autoecious.)
Lopholejeunea (vol. 3)
Plants golden brown or fuscous to brown-black, never inky, not nitid, when dry with leaves convolute, when moist spreading; gynoecia with neither bracts nor perianth keels toothed or lamellate; perianth with ca. 8–10 rounded plicae, the plicae smooth, rounded; underleaves obdeltoid; trigones conspicuous, cordate. (Our taxa paroecious.)
Acrolejeunea (vol. 3)
Group N. Similar to Group M (leaves basically complicate-lobed, often with a smaller, inflexed ventral lobe = lobule); branching only lateral (in Balantiopsaceae sometimes also ventral-intercalary). Differing in the 2- or 4-lobed underleaves. (Balantiopsaceae, Schistochilaceae p. p., see Group L for Paraschistochila.)
Key
1
Leaf cells strongly elongated, rectangulate, often tiered; branches never infra-axillary, in part ventral-intercalary; marsupium developed
Balantiopsis (vol. 2)
Leaf cells neither strongly elongated nor tiered; branches often infra-axillary (and never ventral-intercalary); marsupium absent, a coelocaule developed
2
Rhizoids scattered, claret-red, their apices often dendritically, ramified but not digitate, their branches often coalescent, not or sparingly septate; leaves 1-stratose, often pigmented, often ± toothed or ciliate
Schistochila (vol. 2)
Rhizoids in fascicles from underleaf and ventral leaf bases, colorless, apices often divided into multiseptate, finger-like digits; leaves normally polystratose, at least locally, clear green, mostly edentate or with few, broad-based teeth
Pachyschistochila (vol. 2)
Group O. Complanate plants, with complicate-bilobed lateral leaves; lobules of stem leaves (and/or branch leaves) galeate or helmet-shaped, longer than wide, only feebly attached to (and often remote from) stem; water sac formed only by the inflated sac-like lobule; underleaves distinct, 2- or 4-lobed. (Lepidolaenaceae, Jubulopsaceae, Frullaniaceae.)
Key
1
Leaf lobes (and usually underleaves) variously ciliate or toothed; underleaves 2- or 4-lobed; perianth reduced or lacking, a clavate, paraphyllose or lamella-bearing coelocaule produced
Leaves with lobe margins entire (except Frullaniachevalieri, which has lobe margins crenulate denticulate); underleaves 2-lobed, lobes edentate. Perianth conspicuous, without a coelocaule at base
Frullania (vol. 3)
2
Underleaves of stems and branches all 2-lobed, with reflexed margins, no underleaf lobes converted into water sacs. Coelocaule hidden by imbricate, mutually involute bracts + bracteoles
Gackstroemia (vol. 2)
Underleaves of main stems, at least, all deeply 4-lobed. Coelocaule not wholly obscured by bracts + bracteoles
3
Branch underleaves never with lobes converted to water sacs, 2- or 4-lobed. Coelocaule club-shaped, covered with multiform lamellae
Jubulopsis (vol. 2)
4
Plants soft-textured, pale green, spongy, never reddish. Main stems paraphyllose. Coelocaule slenderly clavate. (On peaty ground or soil.)
Lepidogyna (vol. 2)
Plants firm, deep green, but usually reddish to red-brown pigmented, much like Frullania. Stems smooth. Coelocaule obpyriform to short-clavate. (Usually epiphytic.)
Lepidolaena (vol. 2)
Group P. Plants with stem leaves developing distinct, galeate lobules (water sacs) formed by the infolded ventral lobe and the opposed basal portion of the lobe, the lobules attached by an elongate keel to the lobe. Plants pale to yellowish to grass-green. Vegetative branches thecal, with a distinct basal sheath (except at times in Nephelolejeunea). (Lejeuneaceae.)
Key
1
Underleaves normal in number: one per pair of lateral leaves (segmentation helical); underleaves usually not with divergent slender lobes
Underleaves duplicated: one for each lateral leaf (segmentation pendular); underleaves with very slender, usually divergent, lanceolate to linear-lanceolate lobes. Leaf insertion on a short base, the leaves constricted at base. (Sometimes with discoid gemmae.)
2
Leaves constricted basally, with abbreviated, subtransverse insertion on stem: lobule attached by only 1–2(3) cells to stem; lobes narrow, usually obovate, elongated; underleaves remote, small, narrow, with very narrow lobes; sometimes with discoid pluricellular gemmae from surface cells. (Tuyamaelloideae)
Leaves with a J-shaped insertion line: the insertion not constricted, the lobe insertion running up stem (longer arm of J); lobule broadly attached to stem by 5–6 or more cells (short arm of J); never with discoid gemmae (in New Zealand)
Lobule with 3 distinct teeth distally, the median largest. Lobule obovoid-cuneate, with hyaline papilla entally displaced; lobes narrowly obovate, rounded distally
Austrolejeunea (vol. 3)
4
Lobule linear, the apical tooth formed of 2 rounded, projecting cells, with hyaline papilla inserted between the teeth, thus externally visible; lobe spatulate-obovate, rounded distally
Siphonolejeunea (vol. 3)
Lobule obovoid-cuneate, the apical tooth single, formed of 2 superposed cells, the hyaline papilla entally displaced to base of tooth; lobe rounded or sharply pointed
Nephelolejeunea (vol. 3)
5
Leaf margins neither lobulate nor toothed; lamina smooth, unarmed; underleaf bilobed but lobes and disc unarmed on margins; mostly not epiphyllous
Leaf margins variously lobulate-dentate or dentate (at least above); lamina smooth or hispid; underleaves with margins crenulate or dentate; always epiphyllous, on mosses, ferns, and seed plants. Lobules inflated, strikingly so, the entire free (anterior) margin conspicuously involute, the apical tooth never apparent without dissection
6
Underleaves subrotundate to ovate or obovate, with erect lobes (which are always over 2 cells wide at base on stem underleaves); leaves without ocelli (except in Rectolejeunea, which has caducous leaves); lobes usually rounded (except Stenolejeunea), not elongated: length:width ca. 1.1–1.2:1
Underleaves narrow-based, obdeltoid, with strongly divergent lobes; leaves with ocelli of lobe base (and often also laminar ocelli); lobes elongated, elliptical to falcate (then acute to acuminate), length:width ca. 2–4:1. Caducous leaves lacking: asexual reproduction, if present, via short caducous branches from leading axes (= cladia). (Small or tiny, rather pellucid, light green, delicate plants.)
7
Lobules of stem leaves with hyaline papilla inserted in a faint notch just proximad of the 1-celled apical tooth; plants usually clear to yellowish to whitish green, mostly rather translucent; cells with (1–3)4–many small, glistening or granular oil-bodies
Lobules (always well developed) with hyaline papilla in sinus, distad of the apical tooth; plants rather firm: walls thick, or trigones conspicuous, often rather opaque and whitish green or grey-green; cells with 1–2(4) very large, coarsely botryoidal oil-bodies
Cheilolejeunea (vol. 3)
8
Leaf lobes rounded, never with sharp tips ending in 2 superposed cells; no asexual reproduction (or, Rectolejeunea, with caducous leaves); surface (in ours) smooth
Leaf lobes, especially on branch leaves, apiculate, sharp, ending often in 2 superposed cells; with thalloid, elongating propagulae from lobe margins (which also develop marginal rhizoids); surface minutely granular
Stenolejeunea (vol. 3)
9
Leaves persistent, lacking ocelli; perianth 5-keeled or inflated, never compressed; marginal cells over 12 µm
Lejeunea (vol. 3)
Leaves caducous, the lobes ocellate; perianth compressed, the dorsal keel vestigial or weak; marginal cells only ca. 8 µm
Rectolejeunea (vol. 3)
10
Ocelli (in ours) basal, (0)1–2 per lobe; leaf lobes sharp to acuminate; gynoecia always with 1–2 innovations; median cells (in ours) with 1 or 2–4 granular oil-bodies; underleaves without a conspicuously small-celled basal field, the marginal cells not differentiated; plants never intensely fragrant when alive
Ocelli 6–8(9) per lobe, in a broken (interrupted) median line + a few ocelli distally scattered; leaf lobes elliptical-lingulate, blunt to rounded, in drying standing stiffly away from substrate; gynoecia lack innovations, on very short branches; median cells with 7–11 oil-bodies; underleaves transversely rectangular, margined by 6 very large cells contrasted to a small-celled field within; lobes uniseriate, widely separated, few-celled; plants intensely fragrant when alive
Leptolejeunea (vol. 3)
11
Median cells (in ours) with 2–4 oil-bodies; leaf lobes decurved, cuspidate; underleaf lobes blunt to rounded, ca. 4 cells wide at base; gynoecia (in ours) on long to short shoots, with 1–2 Lejeunea -type innovations (first appendage of innovation a lateral leaf)
Harpalejeunea (vol. 3)
Median cells (in ours) with 1 oil-body; leaf lobes attenuate, subacuminate, erect to suberect; underleaf lobes few-celled, long-acute; gynoecia (in ours) on short branches, with 1 innovation of Pycnolejeunea type (first appendage an underleaf)
Drepanolejeunea (vol. 3)
12
Lobes appearing lobulate-crispate, due to reflexed, lunate marginal sectors, and lobes distally denticulate; lobe and perianth surfaces smooth; underleaves with entire lobe margins, lobes divergent and rounded, margins crenulate
Kymatolejeunea (vol. 3)
Lobes not crispate and lobulate, merely gently convex, with both margins and surfaces spinose-armed; perianth surface also spinulose; underleaves with margins spinulose, the lobes sharp and erect
Echinolejeunea (vol. 3)
13
Leaves obovate or rotundate-obovate, with lobule smaller than and clearly distinct from lobe; oil-bodies granular
Diplasiolejeunea (vol. 3)
Leaves complex: the enlarged (hardly recognizable) lobule forming a water sac, opening by a flap, formed jointly with the reduced dorsal lobe: the lobule apex produced as a tumid or horn-like, inflated sac; oil-bodies homogeneous
Colura (vol. 3)
