Liverworts v1 (2008) - A Flora of the Liverworts and Hornworts of New Zealand Volume 1
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Lepidozia (Dumort.) Dumort.

Lepidozia (Dumort.) Dumort.

Pleuroschisma sect. Lepidozia Dumort., Syll. Jungerm. Europ. 69. 1831.

Lepidozia (Dumort.) Dumort., Recueil Observ. Jungerm. 19. 1835, nom. cons.

Herpetium sect. Lepidozia Nees, Naturgesch. Eur. Leberm. 3: 31. 1838.

Lepidoziopsis E.A.Hodgs., Rec. Domin. Mus. 4: 105. 1962.

Type: Lepidozia reptans (L.) Dumort. (≡Jungermannia reptans L.), a Holarctic, widespread species.

Plants anisophyllous, firm, creeping to loosely prostrate to, more often, somewhat ascending, rarely stiffly erect, yellowish or whitish to olive-green, rarely brownish in life, rarely glaucous, medium-sized to vigorous, usually 0.5–2.5(4) mm wide. Branching irregular to regular, copiously 1- to 2- or 2–3-pinnate, of Frullania type, the lower terminal branches partly or mostly attenuate at the apices, often becoming rhizoid-bearing; Acromastigum -type branching present in a few species; Microlepidozia -type branches lacking; ventral-intercalary microphyllous or leafy branches in some taxa; lateral-intercalary branches lacking. Stem firm to rigid, with (18)20–32 or more rows of firm-walled cortical cells, typically little or at most moderately larger in diameter than those of adjoining medulla, not forming a discrete hyaloderm; subepidermal layers occasionally present, composed of thick-walled cells; medullary cells equal to or slightly smaller than cortical, firm-walled (the outer sometimes strongly so). Rhizoids of normal-leaved axes usually rare or absent, except at shoot bases, arising from underleaf bases; rhizoids on flagelliform shoots from all three merophyte rows. Leaves almost without exception 4-lobed or 4-lobulate on main shoots, incubous to subtransverse, the insertion extending to stem midline dorsally, the leaves typically asymmetrical, the dorsal half (and dorsal 1–2 lobes) larger than the ventral, the lobes usually many-celled (at bases 3–4 or more cells broad in most taxa), the cells similar to those of disc, the lobes usually sharp, often deflexed or inflexed in most taxa; disc margins entire or toothed to spinose-dentate or ciliate. Cells of disc irregularly arranged, not tiered, firm-walled, the lumina often rounded, the trigones concave-sided and mostly obscurely defined; median cells of disc mostly 18–25(40) µm wide; surface smooth or clearly papillose. Oil-bodies present in all leaf cells, colorless, usually (2)5–16 per cell, relatively large, usually botryoidal, less often subhomogeneous, often vestigial in subg. Glaucolepidozia. Asexual reproduction lacking or (Lepidozia fugax, occasionally in L. digitata) via fragmenting or caducous leaf lobes. Underleaves usually 0.5 or less the lateral leaves in area, transversely inserted, symmetrically 4-lobed on main shoots.

Dioecious, rarely (generitype, Lepidozia reptans) autoecious. Gametangial branches all short, devoid of normal leaves and underleaves, ventral-intercalary in origin (the androecia less frequently terminal on primary and secondary branches). Androecia typically spicate, smaller than vegetative shoots, the bracts concave, ± imbricate, lobed like (but usually more shallowly than) vegetative leaves; antheridia 1–2 per bract, the stalk 2-seriate; bracteoles small, lacking antheridia. Gynoecia with bracts and bracteoles identical, 4(6)-lobed to -lobulate, occasionally 2–4-dentate to -lobulate. Perianth fusiform or ellipsoidal-fusiform, bluntly trigonous in at least the narrowed distal sectors, the mouth usually crenulate to crenate-denticulate.

Seta with (8)12–16 large epidermal + 16 or more rows of much smaller internal cells. Capsule ellipsoidal to obloid, the wall 3–5(6)-layered; outer layer of cells often imperfectly tiered, the primary longitudinal and all transverse walls normally lacking pigmented thickenings, the secondary (longitudinal) walls with conspicuous radial (nodular) thickenings, often ± coalescing; inner layer of cells narrower, with radial (nodular) thickenings, ± connected by weak, often vestigial, tangential bands.

Spores papillose-vermiculate, less than 1.5× the diam. of the bispiral elaters.

Artificial Key to Species 1

1 Modified from Engel and Schuster (2001).

Key

1
Branching irregularly to regularly 1-pinnate (except L. serrulata), secondary branches rarely or sporadically developed, if present only 1(2) per primary branch (branches sometimes developing into new leafy shoots); leaves imbricate to contiguous, the insertion weakly to distinctly incubous
2
Branching regularly and consistently 2-pinnate, the branches slender and drooping; leaves typically distant, vertically oriented, the insertion transverse. Plants erect, with rigid, woody stems
21
2
Plants opaque and ivory-white, becoming pale brownish with age; leaves and stems with a dull, waxy, water-repellent surface (at least on new growth)
3
Plants green, yellow-green, brownish to dark brown, but without a dull, waxy, water-repellent surface
6
3
Dorsal margin of leaf moderately to distinctly ampliate; cells of disc and lobes not papillose (but with a fine, granular coating); leaves broadly inserted and clasping the stem, distinctly concave, with involute lobes and margins; leaf insertion strongly incubous to horizontal
4
Dorsal margin of leaf ± straight; cells of disc and lobes striate-papillose, the papillae obscured by the continuous waxy coating; leaves narrowly inserted, the disc flat, with spreading lobes; leaf insertion weakly to distinctly incubous. Disc and lobe cells distinctly thick-walled; first branch underleaf undivided or 2-lobed (rarely 4-lobed). Chatham Is. + South Is. (150–300 m) + North Is. (Waipoua area, 340–540 m)
4
Disc cells with evenly thickened walls, trigones never present; dorsal margin of leaf strongly ampliate, distinctly auriculate at the insertion, the dorsal margin sinuate to distantly toothed; leaf lobes subcaudate, ending in a uniseriate row of 3–5 cells. Rare (South Is.: Fiordland, Paparoa Ra.)
Disc cells lax, nearly thin-walled, trigones lacking to medium; dorsal margin of leaf moderately ampliate, ± clearly cordate at the insertion, the dorsal margin normally entire; leaf lobes acute to apiculate, ending in a single cell or a uniseriate row of 2–3(4) cells
5
5
Plants medium, to 1.5 cm wide (including branches); leaves 0.8–1.3 mm long and broad, the disc 15–22 cells high at dorsal sinus, the median lobes 6–10 cells wide at base; underleaves inserted on 9–10 rows of stem cells, the disc 5–9 cells high; first branch underleaf 2- to 4-lobed, transversely inserted on the stem, often somewhat below the branch. Typically in upper montane forests to penalpine to alpine
Plants small, to 0.6 cm wide (including branches); leaves at most 0.55 mm long and broad, the disc 7–13 cells high at dorsal sinus, the median lobes 3–5 cells wide at base; underleaves inserted on 3–4 rows of stem cells, the disc 2–4 cells high; first branch underleaf 2-lobed (or rarely undivided), obliquely inserted at branch base. Forest plant; 950–1390 m
6
Dorsal margin of leaves serrate or dentate-laciniate or ciliate
7
Dorsal margin of leaves entire (rarely with a basal tooth)
10
7
Dorsal pair of lobes caudate, the distal portion ciliiform and with a uniseriate row of 4–15 cells
8
Dorsal pair of lobes acute to acuminate, the distal portion not ciliiform, the uniseriate row comprised of 2–3 cells
9
8
Branching 1-pinnate; dorsal margin of leaves with numerous (up to 19) spinose teeth or cilia; lobe margins often with paired cilia; underleaf lobes truncate, the summit with several cilia ± equal in size
Branching 2-pinnate; dorsal margin of leaves denticulate-dentate; lobe margins entire or sparingly denticulate; underleaf lobes acute or if ± truncate, then with accessory cilia smaller
9
Dorsal margin of disc and contiguous margin of lobe irregularly dentate to coarsely serrate; leaf lobes acute; underleaves divided to at most 0.5 (the disc 12–16 cells high), the lobe margins entire or sparingly toothed
Dorsal margin of disc and confluent margin of lobe ± regularly spinose-dentate, with discrete, often curved, acuminate spines; leaf lobes acuminate; underleaves divided to 0.55–0.8 (the disc 6–9 cells high), the lobe margins laciniate-multifid (rarely entire)
10
Leaf lobes setaceous, terminating in a uniseriate row of 6–10(12) elongated capillary cells. Leaf insertion distinctly incubous; trigones large and bulging to knot-like; underleaves distinctly connate on one side
Leaf lobes never setaceous, the lobes acute or terminating in a short uniseriate row of up to 2–5 cells
11
11
Leaves appressed to stem, distinctly ventrally secund, ± equally and shallowly (0.25–0.4) lobed, the two median lobes wider than the outer lobes. Slender, wiry plants with strongly ventrally secund branches
Leaves spreading, not appressed to stem, the two median lobes not clearly wider than outer lobes. Leaves often asymmetrically lobed, the dorsal lobes often ± united in a perceptible pair, the ventral lobes usually shorter and often somewhat divergent from the dorsal pair
12
12
Leaves and underleaves variously appendaged, with accessory lobes and often lamellae; underleaves plicate, the lobes ventrally sulcate, the sinus bases strongly reflexed. Plants brown; branches spreading. Penalpine–alpine species
Leaves and underleaves not appendaged (rarely with marginal teeth); underleaves plane
13
13
Plants with brown pigments, typically erect, with distinctly ventrally secund branches (occasionally greenish with spreading branches in var. parvula). Disc cells often distinctly thick-walled, brownish; leaf and underleaf lobes often bluntly rounded. Montane to alpine species
Plants green, typically prostrate to procumbent, with spreading or ± decurved branches
14
14
Median cells of leaf disc large (26–41 mm wide); plants medium-sized; underleaves inserted on 10 rows of stem cells; leaves 0.8–1 mm wide × 0.6–0.9 mm long. Leaves variable in shape, the ventral lobes typically not much shorter than the dorsal (the branch leaves ± equally 4-lobed). Surface of leaf disc smooth; median disc-cells typically with trigones straight to large and bulging
Median cells of leaf disc smaller: 25 µm wide or less (except sometimes in L. bidens); plants small to minute; underleaves inserted on 9 or fewer rows of stem cells; leaves usually less than 0.5(0.6) mm in longest dimension
15
15
First branch underleaf consistently unlobed. South Is. (Westland Prov., 990m) + North Is. (Rotorua area)
16
First branch underleaf of mature shoots 2–4-lobed or mixed 1–2-lobed. Leaves mostly bisbifid: median sinus deeper, dorsal sinus usually shallow, usually strongly asymmetric, the lobes always strongly unequal
17
16
Leaves deeply (0.5–0.6) quadrifid, the lobes lanceolate; dorsal sinus descending at least 0.5 leaf length; lobes ending in ± elongated cells, the lobe cells mostly rectangulate; first branch underleaf subulate; stem half-leaf usually 0.45–0.55 bifid; branch leaves mostly 3-lobed
Leaves under 0.5 quadrifid, the lobes triangular, under 3× as long as broad; dorsal sinus usually descending to 0.2–0.3; lobes ending in non-elongated cells, the lobe cells mostly subquadrate; first branch underleaf ovate-lanceolate; stem half-leaf under 0.25 bifid; branch leaves 4-lobed
17
Leaves ± conspicuously papillose (at least on the lobes) or if indistinctly papillose (suboptimal populations of L. laevifolia) then the dorsal margin of disc ampliate and the outer wall of the marginal cells of disc and lobes not thickened
18
Leaves smooth (rarely finely and indistinctly papillose); outer wall of marginal cells of disc and lobes distinctly thickened
20
18
Marginal cells of disc and lobes with free wall not differentially thickened; dorsal margin of disc ampliate, the margin cordate to auriculate at the insertion; leaves usually distinctly concave, wet or dry. Leaves coarsely papillose, the disc closely and distinctly striate-papillose; lobes with uniseriate row (when present) of non-elongated cells. Common; usually not below 300 m
Marginal cells of disc and lobes with a strongly thickened free wall (the wall often bulging into the cell lumen); dorsal margin of disc ± straight to moderately ampliate proximally, cordate at insertion, not clearly auriculate; leaves plane or only moderately concave, the lobes spreading or incurved. Rare or infrequent species
19
19
Lobe tips caducous, the cells turgid, barrel-shaped, the septa constricted; lobes with uniseriate row of 2–4(5) cells, the cells ± isodiametric, the terminal cell rounded at the tip; median disc cells 17–24 mm wide; underleaves inserted on 7 rows of stem cells. Margins of disc and lobes occasionally toothed. South Is., 560–900 m
Lobe tips not caducous, the cells with straight or concave walls, the septa dilated; lobes terminating in a single cell or at most 2 superposed cells, the cells elongated (to 2:1), the terminal cell ± tapering to a point; median disc cells 11–16 µm wide; underleaves inserted on 4–6 rows of stem cells. South Is. + North Is.; forest species
20
Plants medium-sized (the main shoots, 700–900 mm wide, leaf tip to leaf tip); underleaves inserted on 8 rows of stem cells; leaf disc 8–10 cells high at dorsal sinus, the cells ± longitudinally elongated; leaves with dorsal lobes ending in a uniseriate row of elongated (up to 2:1) cells. Plants of forests (0–330 m in South Is.; to 920 m in North Is.)
Plants minute to small (the shoots 365–490 µm wide, leaf tip to leaf tip); underleaves inserted on 5–6 rows of stem cells; leaf disc 9–12 cells high at dorsal sinus, the cells quadrate or slightly longer than wide; dorsal lobes with cells of uniseriate row ± isodiametric or at most slightly longer than wide. Plants of forests + penalpine zone + (var. alpina) alpine zone (570–1945 m); South Is. + North Is.
21
Leaves inconspicuous, scale-like, narrower than stem, strongly erect and commonly ± appressed to stem, divided to 0.4–0.5 (median sinus)
Leaves conspicuous, wider than stem, stiffly patent or erect-patent from a spreading base, divided to 0.65–0.9
22
22
Leaves ± symmetrical, the lobes 6–8 cells wide at base, ending in a uniseriate row of 7–12 cells; dorsal margin of disc straight to slightly concave; underleaf disc 3–6 cells high, the lobes with a uniseriate row of 7–16 cells
Leaves asymmetrical, the lobes 14–24 cells wide at base, with a uniseriate row of 2–4(6) cells; dorsal margin of disc broadly ampliate; underleaf disc 14–24 or more cells high, the lobes with a uniseriate row of 1–3(4) cells, the lobes ± parallel, ± ventrally sulcate

Lepidozia is a large and polytypic genus, with perhaps ca. 80 species worldwide. Over half of these occur in an area extending from New Zealand (with 23 species) into Southeast Asia (e.g., New Guinea has ca. 17 species). Much of the morphological diversity found within the genus occurs in New Zealand. Of the seven subgenera of Lepidozia, five are present in New Zealand (see Engel and Schuster, 2001; Schuster, 1973). Absent from New Zealand are subg. Chaetolepidozia R.M.Schust., which ranges from Costa Rica south to Colombia and Ecuador, and subg. Cladolepidozia R.M.Schust. of the Paleotropics.

The species show a diverse ecology: they occur principally on soil, peat, or decaying logs, but also on soil-covered rocks, rarely a few invade even the lower trunks of trees. The large bulk of taxa occur in sheltered loci, but several ascend into the alpine zone, where they occur in damp sites of alpine grasslands, or on soil in crevices of crags or cliffs. All of our species are unisexual, but ♂ and ♀ plants often occur intermingled, and in many species sporophytes are produced, if infrequently so.

References: Hodgson (1956); Engel and Schuster (2001); Engel (2004a).

The following treatment of taxa is adapted and modified from Engel and Schuster (2001).

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