Classification
Class
 Nomenclature
Scientific Name:
Pottiaceae subfamily Barbuloideae Hilp., Beih. Bot. Centralbl. Abt. 2, 50: 612 (1933)
 Taxonomy

Twelve genera are recognised in subfamily Barbuloideae for N.Z. Characteristic substrates are soil or rock, often calcareous; no members are epiphytic.

Perichaetia may be either lateral or terminal, and peristomes are usually well developed, as in Barbula, but may be vestigial or absent, as in Anoectangium, Gymnostomum, and in some species of Geheebia and Tridontium.

In earlier treatments, five or six species of Didymodon were accepted for N.Z. (Zander 1993; Fife 1995; Gibb et al. 2018​​​​​​​). All but one, Didymodon calycinus Dix. (see incerta sedis), are now placed in other genera, namely in Geheebia, Gertrudiella, Trichostomopsis or Tridontium​​​​​​​ (Zander 2013; Jiménez et al. 2022​​​​​​​; Beever et al. 2023​​​​​​​; Allan Herbarium 2023​​​​​​​​​​​​​​​​​​​​​; Zhang et al. 2023​​​​​​​).

 Key
1Moist leaves twisted around stem giving a spiral appearance when viewed from above; leaf margins irregularly dentate above, and lower laminal cells firm-walled and porose; KOH colour reaction of upper laminal cell walls yellowLeptodontium
1'Moist leaves with various stances, but not twisted around stem; leaf margins entire, or, if irregularly dentate above, then lower laminal cells thin-walled and lax; KOH colour reaction of upper laminal cell walls yellow to red2
2Stems prostrate, with shoots forming interwoven mats; leaves arranged in three straight or spiralled ranksTriquetrella
2'Stems more or less erect, with shoots not forming interwoven mats; leaves not arranged in 3 ranks3
3Leaves narrowly triangular; leaf margins revolute from base to apexPseudocrossidium (pro parte) [P. hornschuchianum]
3'Leaves linear, ligulate, lingulate, or lanceolate, not narrowly triangular; leaf margins plane from base to apex or recurved in part, not revolute4
4Plants often brick-red below; leaf margins sometimes dentate above; KOH colour reaction of upper laminal cell walls orange-red to redBryoerythrophyllum
4'Plants green, brown or black below; leaf margins not dentate above; KOH colour reaction of upper laminal cell walls yellow to yellow-orange 5
5Plants pale yellow-green above; shoots slender, forming a dense turf on calcareous substrates; dry leaves appressed and slightly twisted; leaf apices apiculate, acute or rounded, even on the same stemGymnostomum
5'Plants without the above combination of characters6
6Leaf apices rounded; costa fails before the leaf apex (usually by >4 cells)7
6'Leaf apices acuminate, acute or obtuse; costa percurrent to excurrent9
7Plants robust; stems usually 15–80 mm; leaves 2.5–5.0 mm; intramarginal border of elongate cells usually discernible in leaf base; laminal cells smoothTridontium (pro parte) [T. tasmanicum]
7'Plants minute to medium sized; stems 1–8 mm (or if >8 mm then leaves 1.5–2 mm); intramarginal border lacking; laminal cells usually papillose (but papillae may be obscure)8
8Stem central strand absentTridontium (pro parte) [T. cockaynei, T. milleneri, T. novae-zelandiae]
8'Stem central strand presentGeheebia
9Plants with a metallic sheen; shoots filiform, forming a dense turf often banded as if by multiple growth spurts; leaves usually <1 mm. Species known in N.Z. only from Mt Owen, Nelson L.D.Ardeuma
9'Plants lacking a metallic sheen; shoots broader, not filiform, not banded; leaves ≥ 1 mm. Species widespread in South I., or throughout N.Z.10
10Costa excurrent as a long, yellow, weakly flexuose hair-pointPseudocrossidium (pro parte) [P. crinitum]
10'Costa percurrent, or short-excurrent11
11Juxtacostal laminal cells in the leaf base conspicuously differentiated from outer cells, either as thin-walled, smooth, hyaline cells, or as firm-walled, uniseriately papillose, yellow cells.12
11'Juxtacostal laminal cells in the leaf base not conspicuously differentiated from outer cells either by ornamentation or by colour13
12Leaf margins erect above, plane below, unistratose throughout; juxtacostal laminal cells in the leaf base often yellow and with uniseriate papillaeBarbula (pro parte) [B. calycina]
12'Leaf margins variably recurved, bistratose above; juxtacostal laminal cells in the leaf base colourless and smoothTrichostomopsis
13Plants hygrophytic; on damp rock, above 1000 m elevation; stems matted together with dense, red-brown tomentumAnoectangium
13'Plants mesophytic; on soil, rock, or artificial calcareous substrates (e.g., concrete, mortar, shell paths, or midden material), with a wide elevation range from near sea level upwards (but rarely above 1000 m); stems lacking dense tomentum14
14Leaves broadly lanceolate, widest just above base; leaf apices acuminate; stem cross-section with a well-developed sclerodermisGertrudiella
14'Leaves lingulate to oblong-lanceolate, widest at base; leaf apices acute to obtuse; stem cross-section with a weakly developed sclerodermis or sclerodermis lacking15
15Leaf apex acute, often terminated by a conspicuous hyaline conical cell; costa failing a few cells before the apex or percurrent; seta yellow; perichaetial leaves conspicuously sheathing the seta baseStreblotrichum
15'Leaf apex obtuse, lacking a hyaline conical cell; costa excurrent as a cusp; seta orange-red to dark purple-black; perichaetial leaves not sheathing the seta baseBarbula (pro parte) [B. unguiculata]
 Biostatus
Indigenous (Non-endemic)
Number of species in New Zealand within Pottiaceae subfamily Barbuloideae Hilp.
CategoryNumber
Indigenous (Endemic)4
Indigenous (Non-endemic)14
Exotic: Fully Naturalised3
Total21
 Notes

Incertae sedis

The N.Z. endemic Didymodon calycinus (Beever 2024) was described by Dixon (1915) from material collected in the vicinity of Mt Bruce, Wairarapa (W. Gray 176, 18 Sep. 1913 "in a boggy place in grass paddock, among rushes", BM 000845861) with a further specimen seen by Dixon, probably from the same locality (W. Gray s.n., 8 Nov. 1914, BM 000845863) and with duplicates in Sainsbury’s herbarium (WELT M001770, WELT M005813a and b). Sainsbury himself made subsequent collections (G.O.K. Sainsbury 18349, WELT M021900, M010124; G.O.K. Sainsbury 18350, WELT M010125) at a second site 150 km to the NW. Both sites were accurately pin-pointed from extra information on the herbarium packets, but despite considerable search effort the species has not been refound. These two remain the only known collection sites; all other putative specimens of Didymodon calycinus that have been located for the present study appear to be misidentified. Two collected by Sven Berggren, from Napier (S. Berggren 193, AK 123949, BM 000845862) and from Wellington (S. Berggren 388, AK 123948, BM 000845864, CHR 486020,) are referable to Geheebia tophacea and Trichostomopsis australasiae, respectively. A voucher specimen (WELT M008396) for a published report of D. calycinus by Bartlett (1985) is also misidentified.

The correct generic and family placement of Didymodon calycinus Dixon is problematic. It bears a resemblance to the very variable Ceratodon purpureus (Ditrichaceae) in many features (see Burley & Pritchard 1990), namely in the lanceolate leaves with recurved leaf margins, stout costa, smooth and irregularly subquadrate upper laminal cells (with KOH colour reaction yellow), short-rectangular basal laminal cells, and sheathing perichaetial bracts. The stem and leaf cross-sections are likewise very similar in the two taxa (cf. Beever 2024, fig. B–E, O with Seppelt 2004 fig. 57 9–14). There is a similarity in sporophyte characters (as seen in WELT M010124 for D. calycinus): a well-developed annulus, a peristome basal membrane with prominent abaxial trabeculae, rami incurved when dry, straight when moist, papillose and nodose. In its typical form, C. purpureus differs from D. calycinus in having acute leaf apices, with a short-excurrent costa and the leaf margin irregularly and bluntly toothed near the apex (vs a rounded leaf apex, failing costa and entire leaf margins in D. calycinus), theca inclined or suberect, curved, and sulcate at least when old and dry (vs theca erect and cylindric in D. calycinus). Rarely C. purpureus, at least in N.Z., may have "obtuse-rounded" leaf apices and "nearly entire" upper leaf margins (A.J. Fife, pers. comm., July 2023). Elsewhere, in Arctic and Antarctic regions, a form of Ceratodon with failing costae and entire leaf margins (Seppelt & Green 1998; Seppelt 2004; Zander et al. 2007), and thus reminiscent of Didymodon calycinus, has been described as Ceratodon heterophyllus Kindb. There are also resemblances between D. calycinus and species of Geheebia, including obtuse to rounded leaf apices with failing costa, recurved leaf margins, a stem sclerodermis, and well-developed stem central strand. If fresh material of Didymodon calycinus is found, DNA sequencing may help establish its true relationships.

Illustrations: Beever 2024. Dixon 1915, pl. 9, fig. 3; Zander 1993, pl. 50, 1–3.

 Bibliography
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