Flora of New Zealand Series

Cephaloziaceae Mig., Krypt.-Fl. Deutschland 1: 465. 1904.

Plants predominantly markedly anisophyllous, rarely subisophyllous (Hygrobiella), often soft-textured and/or lax, usually creeping in growth but sometimes with ascending gemma-bearing axes (lacking in ours), often green, hyaline, the exposed sectors often with reddish, purplish, brownish or fuscous pigments, small to medium in size. Branching sparing and irregular, the branches for the most part ventral-intercalary; lateral-intercalary branches present in some taxa; Frullania -type terminal branching in a few taxa; Andrewsianthus type present in Odontoschismatoideae; stolons and flagelliform axes often developed. Stems various, the cortical cells often larger than medullary cells, at times forming a distinct hyaloderm, at times thick-walled. Rhizoids hyaline to pale brown, scattered, at times rather dense. Leaves alternate, bifid (in ours; 3[4]-lobed in Cephalozia chilensis) to bidentate (Alobiella) to unlobed, the insertion mostly succubous, at times nearly transverse, not extending to stem midline dorsally and delimiting a leaf strip (extending to stem midline in Metahygrobiella). Cells thin-walled to, less often, firm-walled (excluding Odontoschismatoideae), mostly devoid of trigones; surface smooth. Oil-bodies generally (always in ours) lacking, small and granular in a few taxa, large in Odontoschismatoideae. Underleaves bilobed, small to vestigial and then reduced to slime papillae or to a small scale-like lamella bearing slime papillae (the underleaves subequal to leaves in Hygrobiella). Asexual reproduction frequent, often via 1(2)-celled gemmae produced from reduced leaves of tapered branches.

Dioecious, often autoecious, rarely paroecious. Androecia usually on short, ventral-intercalary branches, less frequently on leading shoots, typically spicate; bracts usually much smaller than leaves, usually bilobed, concave at base to strongly ventricose; antheridia normally 1 per bract, the jacket-cells elongated, tiered (except Odontoschismatoideae and Schiffneria), the stalk 2-seriate; bracteoles, when present, lacking antheridia. Gynoecia isophyllous, at apices of weak ventral-intercalary branches, less often terminating long leafy axes; bracts in 1–3 series, nearly uniformly bifid; bracteoles free from bracts, identical to bracts or nearly so. Perianth well developed, terete below, bluntly trigonous distally, with 2 keels lateral + 1 ventral, the dorsal face flat; perianth usually unistratose but at times with up to 8 strata below. Coelocaule absent. Calyptra thin, never fleshy.

Foot lacking a well-developed haustorial collar. Seta usually with 8(9) rows of outer cells and 4(5) rows of internal cells. Capsule wall 2–3-stratose; outer layer of cells normally oblong, with two-phase development, the alternating walls with nodular to nodular-confluent thickenings, or (Metahygrobiella) with two-phase ontogeny indistinct.

Spores papillose to papillose-vermiculate or (Schiffneria) areolate.

There are 16 genera in the family, which is worldwide in distribution. The family is well represented in the Neotropics, with nine genera occurring there (Gradstein et al., 2001). The Cephaloziaceae include six subfamilies, only one of which ( subfam. Cephalozioideae) occurs in New Zealand and is represented only by a few species in two genera (Metahygrobiella, Cephalozia). The other genera in subfam. Cephalozioideae are Cladopodiella Buch (two species), Nowellia Mitt. (10 species), Pleurocladula Grolle (one species) and Schofieldia J.D.Godfrey (one species). The other five subfamilies do not occur regionally and are as follows.

1) subfam. Alobielloideae R.M.Schust. has three genera, one, Alobiellopsis R.M.Schust., with five species, two represented in South Africa (A. heteromorphus (Lehm.) R.M.Schust., A. pillansii (Sim) R.M.Schust.); the other two genera are Alobiella (Spruce) Schiffn. (monotypic) and Iwatsukia N.Kitag. (four species).

2) subfam. Hygrobielloideae R.M.Schust. has one genus, Hygrobiella Spruce, which is monotypic and is subarctic to low arctic in distribution.

3) subfam. Odontoschismatoideae Buchex Grolle, with two genera, Odontoschisma (Dumort.) Dumort. (ca. 12 species) and Anomoclada Spruce (monotypic).

4) subfam. Schiffnerioideae R.M.Schust., with only Schiffneria Steph.

5) subfam. Trabacelluloideae (Fulford) R.M.Schust. with only the monotypic Trabacellula Fulford. Schuster (2002a) also included Fuscocephaloziopsis Fulford (two species) and Haesselia Grolle & Gradst. (two species) in this subfamily, but Engel and Braggins (2005) showed that further studies are required prior to a decision as to subfamily placement of the last two genera. Note that Gradstein et al. (2001) regard Fuscocephaloziopsis as a synonym of Cephalozia subg. Macrocephalozia.

Separation of genera and subgenera at times may be difficult within our area, and a separate key is included, as follows.