Cephaloziella varians (Gottsche) Steph.
Jungermannia varians Gottsche, Ergebn. Deutsch. Polar-Exped. 2(16): 452. pl. 7, f. 1, 2. 1890.
Cephalozia varians (Gottsche) Steph., Résult. Voy. S. Y. Belgica 1897–1899, 6(5): 5. 1901.
Lepidoziella varians (Gottsche) Mahu, Bol. Antárt. Chileno 7: 3. 1987, nom. inval.
Type: South Georgia, Penguin Bay, Will 35.
Cephalozia divaricata var. verrucosa C.E.O.Jensen, Meddel. Gronland 15: 374. 1898 (1897).
Cephalozia verrucosa (C.E.O.Jensen) Bryhn & Kaal. in Bryhn, Rep. Second Norweg. Arctic Exped. (2) 11: 45. 1906, hom. illeg. non C. verrucosa Steph., Hedwigia 32: 1893.
Cephaloziella verrucosa (C. Jens.) Bryhn & Kaal. in Bryhn, Vidensk.-Selsk., Forh. Kristiania 1908 (5): 4. 1908.
Type: Greenland, Scoresby Sound, Hartz.
Type: Canada, Ellesmere Is., Simmons.
Cephaloziella alpina Douin, Rev. Gén. Bot. 28: 269. 1916.
Type: ?
Cephaloziella glacialis Douin, Soc. Bot. France, Mém. 29: 71. 1920.
Type: Scandinavian Alps.
Cephaloziella antarctica Douin, Soc. Bot. France, Mém. 29: 78. 1920.
Lectotype (fide Bednarek-Ochyra et al., 2000): “…Terres antarcticques Canal de la Belgique, sur les rochers isolées au milieu d’un glacier à som… Exp. ant. belg. 12 feb. 98.”
Cephaloziella varians (Gottsche) Steph. fo. minor Grolle, Bull. Brit. Antarct. Surv. 28: 83. 1972, nom. nud.
Type: South Sandwich Is., Candlemas Is., western end of eastern lagoon, Longton 597.
Plants delicate, creeping to suberect, brownish green to deep violet to purplish brown or purplish black, small, 180–300 µm wide, 2–5 mm long. Branching sparse, the branches normally exclusively lateral-intercalary. Stems rather rigid, the cortical cells ± thin- to strongly thick-walled and guttulate; medullary cells somewhat smaller. Rhizoids hyaline, rare or rather common. Leaves usually purplish or ± violet-brown, at least near tips, erect to suberect to obliquely spreading, distant to approximate, transverse to subtransversely inserted, weakly to moderately concave (often loosely sheathing stem), mostly 1.5–2× as wide as stem, subquadrate to short-oblong when flattened, 170–225 µm wide and long, bilobed to 0.5–0.7; lobes little divergent, narrowly ovate to ovate-triangular, erect and often ± incurved at tips, relatively broad, the base usually 6–9, occasionally 10–12(14), cells wide, terminating in 1(2) superposed cells, or blunt to ± rounded, the lobes entire; sinus sharp, never gibbous or reflexed at base. Cells with walls thin to ± strongly thick-walled, moderately small, usually 11–14 µm wide × 12–16 µm long; surface usually smooth (in pachydermous forms ± verruculose). Oil-bodies (Schuster, 1980b sub Cephaloziella arctica) 2–7 per cell, glistening and conspicuous, nearly homogeneous in appearance, 2–3.2 × 2.5–3.6 µm, ovoid to ellipsoid, averaging somewhat smaller than chloroplasts (which are 3.5–4 µm long). Underleaves usually present throughout on mature, robust shoots (on weak shoots often sparse and very small), small and lingulate to lanceolate or bilobed, 3–6 cells high × 2–4 cells wide, larger in androecia. Gemmae usually violet, 2-celled, smooth or with a weak projection at each end, ellipsoidal-fusiform, 10–12 × 18–22 µm.
Autoecious, usually fertile. Androecial branches usually numerous and conspicuous, often arising from beneath unfertilized ♀ inflorescences, at other times as ventral branches a short distance beneath apices of fertilized ♀ shoots; androecia terminal but becoming intercalary, compactly but narrowly spicate; bracts almost always deep violet or violet-black (even when rest of plant is only slightly pigmented), in 4–10 pairs, imbricate, erect, bilobed, with ± incurved lobe apices, usually entire-margined, ventricose at base; antheridia 1 per bract. Gynoecia almost always on ± elongate, leafy shoots, often with 1–several subinvolucral innovations (which are more often ♂ than ♀); bracts often vinaceous or purplish but with margins decolorate, ovate to subquadrate, bilobed to 0.45–0.6, the lobes broadly ovate-triangular to ovate, ca. 9–14 cells wide, nearly entire-margined or finely and remotely denticulate; bracteole usually considerably smaller than bracts, 2(or irregularly 3–several)-lobed, connate for 0.1–0.3 with bracts. Perianth oblong to oblong-clavate, the apical region, at least, formed of slightly to ± strongly thick-walled, rectangular cells, decolorate in apical 2–5 or more rows; mouth cells crenulate by cells that are laterally free only near the summit and thus ± weakly projecting, the cells ca. 12 µm wide × 18–24 µm long.
Capsule ovoid-ellipsoidal, the outer layer of wall with nodular thickenings; inner layer of cells with semiannular bands, the bands narrow.
Spores 9–12 µm in diam., delicately verruculose. Elaters 7–8 µm in diam., 2-spiral.
Distribution and Ecology : Bipolar, in the Northern Hemisphere probably pan-Holarctic, with a probable continuous distribution in the Arctic (often in basic-rich areas).Apparently very widespread and the most ubiquitous species of the genus in the Arctic; on the Arctic coast of Ellesmere Island it is one of the two commonest hepatics (Schuster, 1980b, p. 150) and it “is perhaps the commonest and most widespread Cephaloziella in Greenland and is one of the most abundant species of Hepaticae found there” (Schuster and Damsholt, 1974, p. 326). In the Antipodes in New Zealand, 1615 m on South Island, South Georgia (type of Cephaloziella varians) and in the Antarctic, where the species “is the commonest, most widespread and the most abundant species of liverwort in Antarctica. It is the only liverwort known to occur in the Continental Antarctic...” (Bednarek-Ochyra et al., 2000, p. 87).
In New Zealand the species is known only from near the summit of the “Old Man” (Old Man Ra., Otago), where it grew on peaty soil along a small alpine rill, with Lophozia excisa, L. autoica, Cephalozia austrigena and Hygrolembidium acrocladum (Schuster, 1972a).
Comments : Schuster (1980b) deals with this species in detail and provides numerous figures, and Schuster and Damsholt (1974) also provided a discussion. Bednarek-Ochyra et al. (2000) provided useful comments on this species. The species is known from New Zealand only from the following, which was described from a collection from the Old Man Ra.
