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Published In: Index Bryologicus 557. 1896. (Index Bryol.) Name publication detailView in Biodiversity Heritage Library

Project Name Data (Last Modified On 10/24/2011)
Acceptance : Accepted
Project data     (Last Modified On 10/24/2011)

67. HENNEDIELLA           Plates 95, 96, 97, 98.

Hennediella Par., Ind. Bryol. 557, 1896, nom. nov. for Hennedia R. Br. ter. Type: Hennediella macrophylla (R. Br. ter) Par.

Hennedia R. Br. ter, Trans. New Zealand Inst. 25: 285, 1893, hom. illeg. non Harv., 1860. Type: Hennedia macrophylla R. Br. ter, lectotyp. nov.

Beckettia C. Müll., Hedwigia 37: 77, 1898. Type: Beckettia bruchioides C. Müll.

Neobarbula Dus., Bot. Not. 1905: 299, 1905. Type: Neobarbula magellanica Dus.

Bauriella Warnst., Hedwigia 57: 88, 1915, nom. inval. prov. Type: Tortula polyseta (C. Müll.) Warnst.

Henediella Paris ex Fleisch., Musci Fl. Buitenzorg 4: 1697, 1923, orthogr. var.

Willia subg. Schistidiella (C. Müll.) Broth., Nat. Pfl. 1(3): 417, 1902. Type: Willia marginata (Hook. f. & Wils.) C. Müll.

Hennediella subg. Beckettia (C. Müll.) Roth, Aussereur. Laubm. 219, 1911.

Hennediella subg. Hennedia Roth, Aussereur. Laubm. 220, 1911, nom. illeg. incl. typ. gen.

Willia sect. Schistidiella C. Müll., Gen. Musc. Fr. 424, 1900. Type: Willia marginata (Hook. f. & Wils.) C. Müll.

Tortula sect. Neobarbula (Dus.) Card., Wiss. Ergebn. Schwed. Südpol. Exp. Nordenskjöld 4(8): 95, 1908. Type: Tortula magellanica Dus.


            Found on soil and rock, commonly in wet areas, nearly worldwide but most diverse in temperate areas of the austral region.


            This previously monotypic genus has been treated as a synonym of Pottia (Dixon 1923; Sainsbury 1955; van der Wijk et al. 1959–69); however, a clear sporophyte reduction series involving other species (here transferred to Hennediella) and the distinctive morphology of gametophyte warrant recognition at the generic level. Support for recognition of distinctive characters of at least the gametophyte includes Sainsbury's (1955) comment that Hennediella be treated as “a section of Pottia distinguished by the toothed and bordered leaves and by the very large, usually mitriform, calyptra¼”; A. Fife's comment (pers. comm.) that Pottia macrophylla (Pl. 95, f. 1–8) deserves recognition (as Hennediella); Corley et al. (1981) suggested Tortula stanfordensis (Pl. 98, f. 7–11) should be in a separate section or genus; and Matteri (1977a, 1977b, 1988a) pointed out that Pottia austrogeorgica (Pl. 96, f. 1–7), which she regarded as a good species, was vegetatively similar both to the P. heimii (Pl. 96, f. 18–21) complex and to Hennediella macrophylla. Recently, Mishler (1990), in a phylogenetic analysis of 23 Tortula s. lat. species from Mexico, found that T. stanfordensis, T. polyseta (Pl. 98, f. 1–6) and T. leiostoma, all here recognized in Hennediella, are “likely to be monophyletic.” Blockeel (1991) appreciated the relationships of H. macrophylla, T. stanfordensis and P. austrogeorgica, and provided a detailed evaluation recognizing Hennediella as a good genus for at least these three species, making the appropriate combinations (and mentioned my sketchy generic key that included Hennediella as conceived here, Zander 1989, as support).

     Characters here considered of most importance are the often large and commonly broadly lanceolate leaves (Pl 95, f. 3–4) bordered by a band of usually elongate thick-walled cells, with usually serrulate to dentate and almost always plane (narrowly recurved in H. limbata, Pl. 97, f. 1–6) upper laminal margins, upper medial laminal cells large, superficially rather flat (Pl. 95, f. 7, 11, 16), thin-walled and strongly hollow-papillose (rarely epapillose, as in H. denticulata, Pl. 96, f. 12–14), costa usually with a dorsal epidermis, stereid band elliptical, or semicircular to rounded (Pl. 95, f. 7, 11, 16—like Tortula and unlike Syntrichia, which has lunate dorsal stereid band sections), sporophytes in a reduction series (also like Tortula and unlike Syntrichia), operculum often long and narrowly rostrate (Pl. 98, f. 6), calyptra enlarged, sometimes inflated (Pl. 95, f. 8; 97, f. 7; 98, f. 17), and KOH color reaction red. The perigonia are borne terminally on lateral branchlets on the archegoniate stems of H. macrophylla and H. longirostris. Hennediella densifolia is also monoicous but has gemmate perigonia sessile on the main axis. Dixon (1923) noted that the calyptra of H. macrophylla is campanulate but tears basally as it matures, while Sainsbury (1955) observed that in this species the inflated calyptra may appear campanulate in some collections and cucullate in others with much variability and intergradation in shape.           The costal guide cells are in what may be interpreted as two layers, the ventral layer being of smaller and somewhat thicker walled cells than those of the dorsal guide cell layer. Because the stereid band is made up of substereid cells, the ventral layer of guide cells, although intermediate in appearance between the larger layer of guide cells and the dorsal substereids, may be interpreted as indication of a second stereid band. This, plus the common lanceolate leaf shape may imply a relationship to the Barbuleae. Its closest relationship, however, is probably with Tortula species having leaves with similar lax areolation and differentiated border (e.g. Tortula solmsii), but which differ by the generally broader leaves, more convex superficial surface of the laminal cells, lack of marginal teeth, and yellow KOH reaction (see, however, Cladograms 11 and 14 for alternative hypotheses). The upper laminal cells of Hennediella are rather large, but the genus may be distinguished from Chenia, Dolotortula and Sagenotortula, similar genera with red KOH reactions and very large upper laminal cells, by the papillose upper laminal cells, except in the case of the epapillose H. denticulata, which may be distinguished by its unistratose border of elongate cells that are denticulate in the upper portion of the leaf.

     Like Pseudocrossidium, Hennediella is an essentially austral genus with extensions of some species into the north temperate zone probably through migration routes along north-south cordilleras.


Number of accepted species: 20
Species Examined: H. acletoi (US), H. acutidentata (NY), H. angustifolia (NY), H. arenae (NY, PC), H. austrogeorgica (NY), H. bellei Bartr. (FH), H. densifolia (NY), H. denticulata (NY, BM), H. heimii, H. heteroloma (BUF, TENN), H. kunzeana (NY), H. limbata (NY), H. longipedunculata (NY), H. longirostris (BM), H. macrophylla (BM), H. marginata (BM), H. oedipodioides (NY), H. polyseta (BUF, FH, TENN), H. serrulata (NY), H. stanfordensis (BUF, NY), H. steereana (BUF).


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            Plants forming loose turf or cushions, green above, reddish brown below. Stems branching irregularly, ca. 0.5–1.0 cm in length, transverse section rounded-pentagonal or elliptical, central strand present, sclerodermis absent or represented as an outer cortex of substereid cells, hyalodermis absent or weakly developed, not collapsed; axillary hairs of ca. 5 cells, the basal 1–2 cells occasionally somewhat thick-walled; rhizoids usually common. Leaves appressed or incurved when dry, spreading and sometimes reflexed at midleaf when moist, ovate to long-lanceolate, occasionally ligulate or spathulate, comparatively large, (1.5–)3–4(–7.0) µm in length, upper lamina plane or broadly channeled across leaf, margins plane or only rarely narrowly recurved, often dentate above, less commonly entire, with a border (occasionally intramarginal) of short-rectangular to elongate cells, these usually less papillose or smooth and occasionally thick-walled but rarely bi‑ or tristratose; apex acute or rarely obtuse, rarely fragile; base not differentiated in shape or ovate and somewhat sheathing, commonly bordered by cells similar to those of the upper border; costa percurrent or short-excurrent as a flattened, denticulate mucro, rarely ending 3–4 cells below apex, costa with lamina inserted laterally, superficial cells quadrate to short-rectangular and papillose ventrally, dorsally elongate and smooth or papillose, 4–5 rows of cells across costa ventrally at midleaf, costal transverse section round to elliptical, stereid band one, of substereid cells and elliptical to round in shape, ventral epidermis present, dorsal epidermis usually present (occasionally present only laterally on the costa or sometimes absent, guide cells 2–4 per layer in (1–)2 layers, hydroid strand present, usually large; upper laminal cells relatively large, quadrate to hexagonal or short-rectangular, ca. 18–24 µm in width, 1–2:1, walls thin to evenly thickened, superficially flat to weakly convex; papillae hollow, simple to bifid, usually 6–8 or more per lumen, occasionally absent; basal cells differentiated across leaf, rectangular, 18–30 µm in width, (2–)4–6:1, walls thin. Autoicous, paroicous, cladautoicous, or dioicous. Perichaetial leaves somewhat larger than the cauline leaves. Perigonia in dioicous species borne terminally on plants somewhat smaller than the perichaetiate. Seta extremely various in size among species, 0.05–2.5 cm in length, 1 per perichaetium, yellow-brown to dull brown, twisted counterclockwise; capsule occasionally systylious, theca 0.8–3.5 µm in length, yellow-brown to dark brown, ovate to cylindrical, occasionally microstomous or macrostomous, exothecial cells rectangular, 25–35 µm in width, 3–5:1, walls thin to irregularly thickened, stomates phaneropore, at base of capsule, annulus of 2–4 rows of vesiculose cells; peristome teeth absent, rudimentary, or of 32 long, filamentous teeth, densely spiculose, when well developed to 1.5 µm in length, with many articulations, twisted counterclockwise, basal membrane occasionally to 500 µm in height, spiculose. Operculum conic to rostrate, often narrowly so, (0.6–)1.2–1.8 µm in length, cells twisted counterclockwise. Calyptra cucullate, occasionally flaring below and not split, smooth, 2.0–4.5 µm in length. Spores sometimes quite large, 8–30 µm in diameter, yellowish, essentially smooth, papillose or tuberculate. Laminal KOH color reaction red. Reported chromosome numbers: n = 25+m, 26, 26+m, 26+2m, 50.

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