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.
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.
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).
New combinations:
Hennediella acletoi (Robins.) Zand., comb. nov. (Tortula acletoi
Robins., Phytologia 12: 390, 1971).
Hennediella acutidentata (Card. & Thér.) Zand., comb. nov. (Pottia
acutidentata Card. & Thér., Bull. Mus. Hist. Nat. Paris 22: 338, 1916).
Hennediella angustifolia (Herz.) Zand., comb. nov. (Calyptopogon
angustifolius Herz., Beih. Bot. Centralbl. 26(2): 63, 1910; Tortula
angustifolia (Herz.) Herz., nom. illeg.; Tortula herzogii
Zand.).
Hennediella arenae (Besch.) Zand., comb. nov. (Barbula
arenae Besch., Bull. Soc. Bot. France 32: 59, 1885; Tortula arenae
(Besch.) Broth.).
Hennediella arenae var. petriei (Broth. ex Beck.) Zand., comb.
et stat. nov. (Tortula petriei Broth. ex Beck., Trans. New Zealand
Inst. 29: 441, 1897; Tortula arenae subsp. petriei (Broth.)
Lightowlers, J. Bryol. 13: 371, 1985), not seen.
Hennediella bellei (Bartr.) Zand., comb. nov. (Desmatodon
bellii Bartr., Bull. Brit. Mus. Nat. Hist. Bot. 2: 56, 1955).
Hennediella densifolia (Hook. f. & Wils.) Zand., comb. nov.
(Barbula densifolia London J. Bot. 3: 543, 1844; Tortula densifolia
(Hook. f. & Wils.) Hook. f. & Wils.).
Hennediella denticulata (Wils. in Mitt.) Zand., comb. nov. (Barbula
denticulata Wils. in Mitt., Kew J. Bot. 3: 50, 1851; Tortula denticulata
(Wils. in Mitt.) Mitt.). This peristomate species is probably the same as Tortula
densifolia (not seen).
Hennediella heimii (Hedw.) Zand., comb. nov. (Gymnostomum
heimii Hedw., Spec. Musc. 32, 1801; Pottia heimii (Hedw.) Hampe).
Hennediella heimii var. alpina (Amann) Zand., comb. nov.
(Pottia heimii var. alpina Amann Bull. Soc. Vaudoise Sc. Nat. 54:
42, 1921), not seen.
Hennediella heimii var. arctica (Lindb.) Zand., comb.
nov. (Pottia heimii var. arctica Oefv. K. Vet. Ak. Foerh. 23:
551, 1867), not seen.
Hennediella heimii var. brachyphylla (Warnst.) Zand., comb.
nov. (Pottia heimii var. brachyphylla Warnst., Hedwigia 58:
100, 1916), not seen.
Hennediella heimii var. brevicuspis (Warnst.) Zand., comb.
nov. (Pottia heimii var. brevicuspis Warnst., Hedwigia 58:
98, 1916), not seen.
Hennediella heimii var. breviseta (Warnst.) Zand., comb.
nov. (Pottia heimii var. breviseta Warnst., Hedwigia 58: 99,
1916), not seen.
Hennediella heimii var. drummondii (Warnst.) Zand., comb.
nov. (Pottia heimii var. drummondii Warnst., Hedwigia 58: 96,
1916), not seen.
Hennediella heimii var. eurystoma (Card. & Broth.)
Zand., comb. nov. (Pottia heimii var. eurystoma Card.
& Broth., K. Svensk. Vet. Ak. Handl. 63(10): 20, 1923), not seen.
Hennediella heimii var. guessfeldtii (Schlieph.) Zand., comb.
nov. (Pottia guessfeldtii Schlieph., Ber. Deutsch. Bot. Ges. 2: 461,
1884; Pottia heimii var. guessfeldtii (Schlieph.) Warnst.), not
seen.
Hennediella heimii var. lanceolata (Warnst.) Zand., comb.
nov. (Pottia heimii var. lanceolata Warnst., Hedwigia 58: 98,
1916), not seen.
Hennediella heimii var. magellanica (Warnst.) Zand., comb.
nov. (Pottia heimii var. magellanica
Warnst., Hedwigia 58: 95, 1916).
Hennediella heimii var. maxima (Card.) Zand., comb. nov.
(Pottia heimii var. maxima Car., Bull. Herb. Boiss. ser. 2, 5:
1002, 1905), not seen.
Hennediella heimii var. spegazzinii (C. Müll.) Zand., comb.
nov. (Pottia spegazzinii C. Müll., Flora 68: 414, 1885; Pottia
heimii var. spegazzinii (C. Müll.) Warnst.), not seen.
Hennediella heimii var. thaxteri (Card. & Thér.) Zand., comb.
nov. (Pottia heimii var. thaxteri Card. & Thér., Rev.
Bryol. n. ser. 2: 165, 1930), not seen.
Hennediella heteroloma (Card.) Zand., comb. nov. (Tortula
heteroloma Card., Rev. Bryol. 37: 127, 1910).
Hennediella heteroloma var. eckeliae (Zand.) Zand., comb.
nov. (Tortula eckeliae Zand., Bryologist 88: 354, 1985 [1986]).
Hennediella kunzeana (C. Müll.) Zand., comb. nov. (Barbula
kunzeana C. Müll., Linnaea 17: 586, 1843; Tortula kunzeana (C.
Müll.) Mont. in Gay).
Hennediella limbata (Mitt.) Zand., comb. nov. (Barbula
limbata Mitt., Kew J. Bot. 3: 354, 1851; Tortula limbata (Mitt.)
Mitt., hom. illeg.).
Hennediella longipedunculata (C. Müll.) Zand., comb. nov. (Barbula
longipedunculata C. Müll., Syn. Musc. 1: 630, 1849).
Hennediella longirostris (Hampe ex C. Müll.) Zand., comb. nov. (Pottia
longirostris Hampe ex C. Müll., Syn. 1: 552, 1849; Tortula longirostris
(Hampe ex C. Müll.) Broth. in Par., hom. illeg.).
Hennediella marginata (Hook. f. & Wils.) Zand., comb. nov.
(Schistidium marginatum Hook. f. & Wils., London J. Bot. 3: 539,
1844; Streptopogon marginatus (Hook. f. & Wils.) Mitt.; Willia
marginata (Hook. f. & Wils.) C. Müll.)."
Hennediella oedipodioides (C. Müll.) Zand., comb. nov. (Pottia
oedipodioides C. Müll., Bot. Jahrb. 5: 79, 1883).
Hennediella polyseta (C. Müll.) Zand., comb. nov. (Barbula
polyseta C. Müll., Nuov. Giorn. Bot. Ital. n. ser. 4: 414, 1897; Tortula
polyseta (C.Müll.) Warnst.).
Hennediella serrulata (Hook. & Grev.) Zand., comb. nov. (Tortula
serrulata Hook. & Grev., Edinburgh J. Sc. 1: 299, 1824).
Hennediella steereana (Zand. & Crum) Zand., comb. nov. (Desmatodon steereanus Zand. & Crum, Bryologist 80: 638, 1977).
