Radula demissa M.A.M.Renner, 2013

Renner, Matt A. M., Devos, Nicolas, Patino, Jairo, Brown, Elizabeth A., Orme, Andrew, Elgey, Michael, Wilson, Trevor C., Gray, Lindsey J. & Konrat, Matt J. von, 2013, Integrative taxonomy resolves the cryptic and pseudo-cryptic Radula buccinifera complex (Porellales, Jungermanniopsida), including two reinstated and five new species, PhytoKeys 27, pp. 1-113 : 34-41

publication ID

https://dx.doi.org/10.3897/phytokeys.27.5523

persistent identifier

https://treatment.plazi.org/id/AA04F28A-25D9-5706-9D5A-3A4B33FC6622

treatment provided by

PhytoKeys by Pensoft

scientific name

Radula demissa M.A.M.Renner
status

sp. nov.

Radula demissa M.A.M.Renner sp. nov. Figs 14 View Figure 14 -16 View Figure 16

Type.

New Zealand, North Island, Puaiti Bush south of Rotorua, ca 1,600 ft, 20 Jun 1931, K.W. Allison, (holotype: CHR587329, isotypes: AK, F, NSW, WELT).

Radula epiphylla Colenso Transactions and Proceedings of the New Zealand Institute 21: 71. 1888 [1889] nom. illeg. (Art. 53.1) non R. epiphylla Mitt. ex Steph. Hedwigia 23: 151. 1884.

Type. New Zealand: North Island: 'Epiphytical on fronds of Hymenophyllum (sps); damp woods, Dannevirke, County of Waipawa, 1888, W. C [olenso]'. Not located and non vidi, however Colenso a. 2254, on Hymenophyllum leaf, ex herb Steph, received 1897, BM! is possibly a duplicate of Colenso 's type.

Diagnosis.

Similar to Radula buccinifera and Radula strangulata but differs from both species in the leaf insertion attaining the dorsal stem mid-line, leaving no dorsal cortical cell rows leaf-free, the leaf lobes overlapping across the dorsal stem, such that the stem is not visible from above, the falcate leaf lobes, the rhombic lobules with antical margin sloping inwards toward the stem at an angle of c. 45°, and in its typically epiphytic and epiphyllous habit.

Description. [from CHR587329 and AK280339] Forming hanging wefts or patches of interlocking shoots, brown-green when fresh, fading to a glossy tan or brown in herbarium; shoot systems regularly pinnately branched in male plants and sterile female plants, but pseudodichotomous in fertile female plants due to production of pairs of subfloral innovations below gynoecia; dimorphic, 1.0-2.0 mm wide and up to 40 mm long, branches smaller in stature than parent shoot; older shoot sectors retaining leaf-lobes. Stems 115-175 µm diameter, with cortical cells in a single tier of 20-35 rows. Cortical cell walls yellow-brown to brown pigmented; external free cortical cell wall heavily and continuously thickened, radial longitudinal cortical walls thin, inner tangential walls thin or continuously thickened; medulla cells in 22-43 rows; cell walls yellow-brown pigmented, with small to medium triangular trigones, walls between trigones lacking thickenings. Cortical cells on dorsal stem surface arranged in straight longitudinal rows on young and mature shoot sectors. Leaf insertion reaching dorsal stem mid-line, leaving no dorsal cortical cell rows leaf-free, except in instances where stem growth appears to have introduced an additional cortical cell row to the dorsal stem surface, in which case a single row of dorsal cortical cells is leaf-free, but this row is discontinuous between adjacent leaf pairs; leaf insertion not attaining the ventral stem mid-line, leaving two or three ventral cortical cell rows leaf-free. Leaf lobes ovate-falcate, 650-1100 µm long by 430-900 μm wide, larger on leading shoots, contiguous to imbricate, falcate, acroscopic base not sharply deflexed away from stem, flat, obliquely patent, interlocking over the dorsal stem surface, stem not visible between leaf lobes in dorsal view; margins irregularly but minutely repand, entire to weakly minutely crenulate, the interior lobe margin ampliate, covering the dorsal stem surface and reaching or exceeding the opposite stem margin, antical margin curved, exterior margin curved, postical margin curved; angle between postical lobe margin and keel 60-90°. Lobules rhombic when small to widely rhombic, one eighth to one sixth the lobe area, 255-535 µm long by 180-385 μm wide, larger on leading shoots; keel straight to curved, angle between keel and stem 135°, keel apex and postical lobe margin notched; interior lobule margin free for one fifth to one third its length, free portion ampliate, extending at most half way across the ventral stem surface; acroscopic margin S-shaped, apical portion inclined towards stem; apex rounded to apiculate-rounded; free exterior margin curved, margins plane, entire; lobe-lobule junction antical to the acroscopic end of stem insertion in small lobules to level with or below it in larger lobules; attached to stem along 0.66 to 0.8 of the interior margin, stem insertion curved, particularly at basiscopic end, not revolute at acroscopic end; lobule apex bearing a single papilla, with another two papilla situated on the interior lobule margin above the stem insertion. Leaf lobe cells rounded-oblong, not arranged in rows, unequally sized, 14-25 µm long by 9-18μm wide, thin walled with small triangular trigones, medial wall thickenings absent; cells of lobe margin smaller than those of leaf middle, quadrate to rectangular, 9-15 µm long and wide, interior and exterior cell walls not differential thickened, cell lumen not or only slightly bulging medially; leaf lobe cell surface unornamented, smooth or weakly bulging; upper lobe wall differentially thickened over cell lumen, forming a weak papilla. Oil-bodies two or three per cell, ellipsoidal, filling cell lumen, light-brown, surface granular, internally homogeneous or with a hyaline droplet. Asexual reproduction absent. Dioicous. Androecia on indeterminate branches that continue vegetative or reproductive growth, androecial bracts in 4-20 pairs, lobules epistatic, keel deeply curved, bucket-like, free apical portion triangular, apex obtuse, moderately deflexed, lobes rounded, not caducous, antheridia not seen. Gynoecia terminal on leading shoots, subtended by two subfloral innovations that are full-sized and again fertile; archegonia 145-190 µm tall, archegonia neck six cell columns, 6-8 per gynoecium on a small disc of tissue, encompassed by the protoperianth; female bracts in one pair, symmetrical, tightly imbricate, ovate-falcate, lobe 870-940 μm long by 450-560 μm wide, margins entire to repand; lobules ovate to trullate, one third to one half the lobe area, apex rounded to obtuse, keel arched, margins entire; bract insertion lines interlocking dorsally but not ventrally, insertion equitant. Perianths 2300-3900 µm long and 710-760 µm at mouth, mouth entire to irregularly lobed, perianth narrowing gently from slightly flared mouth to approximately one third to one half length above base, where 670-790 µm wide, then tapering to narrow cylindrical base, often inflated independent of sporophyte growth, in which case perianth labia roll inward to overlap one another, rather than laying in plane, the inrolling may cause perianths to appear weakly bicornute, perianth walls unistratose above, with bistratose collar 3 or 4 cell tiers high above the perianth-calyptra junction; basal stem perigynium present, 5-6 stratose, cell walls not thickened or pigmented; perianth-calyptra fusion elevated above female bracts on 9-15 tiers of cells; calyptral perigynium present, base of calyptra bistratose, unistratose above, unfertilised archegonia elevated on surface of calyptra.

Etymology.

From Latin demissa, hanging, in reference to the hanging fan shaped wefts formed by this species when growing on twigs and branches, also in reference to the frequent colonisation of leaves of the fern Hymenophyllum demissum .

Distribution and habitats.

South-eastern Australia in Victoria and Tasmania, and New Zealand. On mainland Australia Radula demissa occurs only on tablelands and highlands in southern and eastern Victoria, but in Tasmania the species is widespread. Radula demissa grows in cool temperate and warm-temperate rainforests and occasionally in wet sclerophyll forests, particularly in the southern part of its range.

In New Zealand Radula demissa occurs throughout all three main islands, and extends east to the Chatham Islands and south to the Auckland and Campbell Islands. It inhabits all forest environments, but is rare in coastal forests with high exposure to salt-laden winds. Radula demissa occupies a broad elevation range, from the coast to around 1600 m asl, where it grows in subalpine scrub above the treeline.

In forests Radula demissa is typically an epiphyte on tree trunks, branches and twigs where it forms characteristic fan-shaped hanging wefts, and is frequently encountered in riparian areas and edge habitats where local light environments are brighter than adjacent forest interiors. Common host species in New Zealand include Melicytus ramiflorus and Beilschmiedia tawa . Radula demissa is also commonly encountered as an epiphyll on the leaves of ferns and broadleaved shrub and tree species. In New Zealand Hymenophyllum species, including Hymenophyllum demissum , Hymenophyllum ferrugineum and Hymenophyllum scabrum are favorite fern hosts, and Beilschmiedia tawa is a frequent broadleaf host. In lowland to montane podocarp-broadleaf forest of tall stature with sparse sub-canopy and shrub layers Radula demissa can be found as a trunk epiphyte and epiphyll on Beilschmiedia tawa , and as an epiphyll on leaves of Hymenophyllum demissum where this species grows in dense carpets on the forest floor. On bark Radula demissa is often associated with Radula allisonii , Radula plicata , Radula strangulata , Metzgeria spp. and various species of Lejeuneaceae including Drepanolejeunea aucklandica and Metalejeunea cucullata , Sematophyllaceae , various Plagiochilaceaespecies including Dinckleria pleurata , Frullania spp. Co-occurrence with phenetically similar species can pose a challenge to identification. On twigs Radula demissa grows with Plagiochila colensoi , Lepidolaena taylorii , Lepidolaena palpebrifolia , Radula plicata , and on leaves Radula demissa grows with Echinolejeunea papillata , Cololejeunea laevigata , and members of the Lejeunea epiphylla Colenso aggregate.

In subalpine scrub Radula demissa occurs on damp rock and adjacent soil, sometimes in highly insolated microsites. Alpine plants exhibit some subtle morphological differences from forest plants, as discussed in the next section.

Variation.

Included within Radula demissa are forms that grow on wet rocks in alpine habitats, typically between 800 and 1600 m asl. Alpine forms exhibit subtle morphological differences from forest plants. The leaves are remote to contiguous rather than imbricate, and the lobes posses a small auricle at the dorsal base of the stem insertion, which is not found in forest plants. The carinal region is typically narrowly inflated along the length of the keel, rather than more broadly inflated across the width of the lobule. Although alpine plants expressing these features are known from several sites in New Zealand, alpine plants are not currently known from Tasmania or mainland Australia.

The existence of subtly different, ecologically distinct, alpine plants of Radula demissa could be explained in several ways. Alpine plants could be an ecotype within a reproductively cohesive species that also occupies forests. They could be a partially reproductively isolated population that retains genetic contact with forest populations. They could be fully reproductively isolated, morphologically cryptic species. These alternatives remain untested. Alpine accessions nest within Radula demissa in our phylogeny, but may warrant investigation through additional genetic, ecological, and physiological data. Given the species straddles what are effectively different biomes (forest and alpine environments: Crisp et al. 2009) the transition from forest to alpine environments concomitant with a change in microsite occupancy could represent a striking example of phenotypic plasticity.

Recognition.

Despite being confused with a range of related and unrelated species, Radula demissa is easy to recognise. The first clue to identity comes from the habitat and microhabitat the plants occupy. Radula demissa is typically an inhabitant of well-lit microsites, and is an epiphyte or epiphyll in forest, and a lithophyte in subalpine shrublands and grasslands. Three other species of the Radula buccinifera complex occupy similar microsites across the range of Radula demissa , and may co-occur with it.

In Australia, Radula buccinifera may co-occur with Radula demissa on tree trunks. Differentiating these two species is only possible on the basis of hydrated, slide-mounted material because deformation associated with drying renders void examination and interpretation of leaf-lobe and lobule shape and orientation in dry material. The most accessible character by which Radula demissa and Radula buccinifera differ is in the orientation and spacing of the leaf lobes, however differences are not as clear cut as between other pairs of species (i.e. between Radula demissa and Radula strangulata ) and this character may not prove reliable in all instances. In Radula demissa the leaves are contiguous to imbricate, and obliquely patent, spreading up and away from the substrate so that they overlap across the dorsal stem surface, such that the dorsal stem surface is not usually visible from above. In Radula buccinifera the leaves are patent to obliquely patent, and while they may spread upward away from the substrate, do not usually overlap completely across the dorsal stem surface so the stem can usually be seen from above, at least in part. However, as there is variation in both leaf lobe orientation and how falcate the lobes are in both species, this character should be used as a guide only. Leaf shape provides another useful clue to identity. In Radula demissa the leaves are falcate, with a distinct notch at the junction of the lobe and keel. In Radula buccinifera the leaves are not or only weakly falcate, and the junction between the lobe and keel forms a simple angle. Lobule shape is also a good source of diagnostic differences, though the differences between these two species are subtle. For convenience, some conceptual re-orientation of the shoot is required to describe these differences clearly. Imagine the shoot is being held vertically with the apex top-most. In this orientation the lobule apex of Radula demissa always lies well above the uppermost point of the ampliate portion of the lobule margin. Between these two points the lobule margin is shallowly S-shaped, and this slopes downward at c. 45° toward the stem. In Radula buccinifera the lobule apex lies variably between the same level as, or slightly above, the uppermost point of the ampliate portion of the lobule margin. Between these two points the lobule margin varies from straight to S-shaped in situ (straight when flattened), but when S-shaped there is a pronounced medial curve. The slope of the antical margin varies between sloping downward toward the stem at up to 45° and remaining level. Several shoots should be examined to gauge the patterns of variation in lobule shape, and comparison with known material is recommended.

The lobule keel differs between Radula demissa and Radula buccinifera . In Radula demissa the keel is curved, on lobules from both main shoots and branches. In Radula buccinifera the keel is typically straight, but may be curved on some lobules, particularly those on branches.

If these more accessible characters prove ambiguous, diagnostic differences can be derived by counting the number of rows of dorsal cortical cells that are not crossed by the leaf insertion lines. In Radula demissa no rows are leaf-free, whereaszero to three rows are leaf-free in Radula buccinifera . To achieve this count, leaves must be removed from the stem in such a way as to preserve the leaf insertion yet make the dorsal stem surface visible, which is a challenging proposition at the best of times.

Radula demissa may co-occur with Radula strangulata on tree trunks and on living leaves close to the forest floor. For guidance on separating these two species on morphological grounds see the recognition section of Radula strangulata .

Radula demissa has been confused with the New Zealand endemic Radula plicata . Radula demissa may co-occur with Radula plicata on tree trunks, branches, and leaves. Although Radula plicata is not closely related to Radula demissa , confusion may arise due to the similarity in size and shape of lobes and lobules, in the curved keel with inflated carinal region, the S-shaped antical lobule margin, and the falcate leaf lobes. When fertile female plants are encountered Radula plicata is readily separated by the plicate perianth surfaces. However, sterile and male herbarium material may cause problems if gross morphological characters are relied on for identification. Radula demissa is readily separable from sterile and male material of Radula plicata by its unornamented leaf-lobe cell surfaces, and bulging, rather than crenulate leaf-lobe margin. In Radula plicata the leaf-lobe cells bear conspicuous coarse granular ornamentation on their surface. This is easily observed in hydrated slide-mounted material on the lobe marginal cells. The cells of the lobe margin are also distinctly crenulate, although comparison with known material is recommended as a guide to the degree of difference between these and the bulging cells of Radula demissa . Differences in stem anatomy also differentiate these two species. In Radula demissa the cortical stem cell walls are brown-pigmented, and the medulla walls are yellow-pigmented. In Radula plicata the cortical and medulla stem cells walls are unpigmented. If living material is being examined, diagnostic differences may be found in the oil-bodies. In Radula demissa there are 2-3 large, light brown, internally homogeneous oil-bodies that have a granular surface and completely fill the cell lumen of each cell. In Radula plicata there are 3-5 small, clear or grayish oil bodies with a smooth surface each containing a hyaline droplet, that are arranged in a loose submarginal ring and do not fill the lumen in each cell.

In alpine environments, Radula demissa may be confused with Radula australiana . However, these two species are readily separated by differences in lobe shape that, with care, can be accessed in the field. In Radula demissa the leaf lobes are falcate, with a notch at the lobe-lobule junction. In Radula australiana the leaf lobes are not falcate, rather the lobule keel runs more or less seamlessly into the lobe outline. Lobule size and shape also differs between these two species. In Radula demissa the lobules are rhombic to widely rhombic and one eighth to one sixth the lobe area. In terms of the same conceptual re-orientation described above, the lobule apex of Radula demissa lies above the ampliate inner lobule margin. In Radula australiana the lobules are quadrate and one sixth to one quarter the lobe area, with the lobule apex lying level with the ampliate inner lobule margin. Furthermore, the ampliate portion of the lobule is typically much larger, and the lobule apex more acute in Radula australiana than in Radula demissa .

Perianth morphology may lead to confusion between Radula demissa and Radula ratkowskiana , particularly given comments by Renner (2005) that inrolling perianth labia and the associated bicornute perianth is unique to Radula ratkowskiana . This is not the case, both features may be developed in young perianths of Radula demissa , which can be distinguished from Radula ratkowskiana by the small rhombic lobules, and the absence of dome-shaped papillae on the leaf lobe cells.

Remarks.

Mitten (1855) was the first to record Radula buccinifera for New Zealand, citing collections by Colenso, Stephenson and Lyall. Hodgson (1944) evidently followed Mitten in accepting Radula buccinifera for New Zealand, but in the absence of an examined type, equated Radula demissa with Radula buccinifera . Hodgson’s (1944) description of Radula buccinifera having glossy dried plants with falcate leaf lobes whose margins extend over the stem and rhomboid lobules agrees with Radula demissa , and on the basis of these characters Hodgson accurately distinguished Radula demissa (as Radula buccinifera ) from Radula strangulata (as Radula levieri for which she had examined type material). Renner (2005) followed Hodgson’s (1944) conception of Radula buccinifera , and the plants presented in his key as Radula buccinifera also correspond to Radula demissa . This error was made despite the fact that he had examined syntype material of Radula buccinifera held in bm in early 2004, apparently the diagnostic differences in leaf lobe and lobule shape were overlooked. As discussed under Radula buccinifera , Mitten’s New Zealand record of that species must be rejected.

In Australia Radula demissa has remained unrecognized, being subsumed within an increasingly broad working circumscription of Radula buccinifera , as discussed under that species.

Colenso’s description of a bipinnately branched plant with dimorphic shoots, ‘peduncled’ perianths with a truncate mouth with slightly uneven labia, and subtended by two divergent subfloral innovations, growing epiphytic on Hymenophyllum is consistent with Radula demissa . However, no definite type material for Colenso’s Radula epiphylla has been located, and our attribution of this invalid name to synonymy under Radula demissa is tentative.

Specimens examined.

New Zealand, North Island: Te Paki Ecological Region and District, Te Paki, Radar Bush, 34°28'03"S, 172°51'15"E, 160 m, 19 Sep 2011, P.J. de Lange 9991 & M.A.M. Renner, NSW970835; Northland Ecological Region, Maungataniwha Ecological District, Mangamuka Range, 35°10'S, 173°24'E, 200 m, 17 Apr 1984, J.E. Braggins 84/26h, AK312107; Maungamuka Stream headwaters, 35°11'S, 173°29'E, 210 m, 30 Nov 1986, J.E. Braggins 86/304, AK259001; Auckland Ecological Region, Hunua Ecological District, Kohukohunui Track c. 1 km west of Kohukohunui summit, 37°02'S, 175°13'E, 630 m, 21 Aug 2000, M.A.M. Renner 00/51, AK280339; Coromandel Ecological Region, Te Aroha Ecological Distric, Mt Te Aroha summit, Dog Kennel Flat track, 37°32'S, 175°45'E, 940 m, 12 Mar 1995, J.E. Braggins 95/213C, AK255288; Rotorua, Blue Lake, May 1953, E.A. Hodgson 10382, BM;Puaiti Bush south of Rotorua, 23 May 1931, K.W. Allison, CHR587333; Near Atiamuri north of Waikato River south of Rotorua, 6 Sep 1929, K.W. Allison, CHR587334; Roto-a-kia Bush, east of Taupo, southern Kaingaroa Plains, 18 Sep 1935, K.W. Allison, CHR587330; near Atiamuri, south of Rotorua, Aug 1930, K.W. Allison, CHR587331; Urewera Ecological Region, Waikaremoana Ecological District, Lake Waikaremoana, Whanganuioparua Inlet, Te Kumi Stream, 38°45'S, 177°9'E, 700 m, 19 Jan 2001, M.A.M. Renner 01/27, AK280333; Central Volcanic Plateau Ecological Region, Taupo Ecological District, Pureora Forest, northern end of Waihora Lagoon, 38°39'S, 175°40'E, 640 m, 27 May 1988, J.E. Braggins 88/020B, AK258496; Eastern Volcanic Plateau Ecological Region, Kaingaroa Ecological District, Poronui Valley, Mangatamingimingi Stream, 38°59'S, 176°16'E, 725 m, 8 Apr 2003, J.E. Braggins & M.A.M. Renner 00/61, AK280574, WELT-H011593, CANB738633.1;Waipapa Ecological Area, Pureora Forest Park, 38°26'S, 175°35'E, 565 m, 26 Jan 1982, J.E. Braggins et al., AK291246; Egmont Ecological Region and District, North Egmont Tourist track and Nature walk, 39°16'S, 174°5'E, 960 m, 12 Sep 1999, J.E. Braggins 99/219C, AK253449;Mt. Ruapehu, off Ohakune Mountain Road, downhill of 7 km marker, 39°22'S, 175°28'E, 890 m, 27 Nov 1992, J.E. Braggins 92/91, AK312261; Hawkes Bay, Morere Hotel, Morere, on tree bark in dense bush, 21 Aug 1964, R.E. Hatcher 2, F;Edge of forest by road, Lake Waikaremoana, 2000 ft, Jan 1955, E.A. Hodgson, ex herb. E.A. Hodgson 11207, S-B89674 as Radula buccinifera ; Egmont Ecological Region and District, Mt Taranaki, Egmont National Park, track to skifield from Pembroke Road, 39°18'S, 174°05'E, 1260 m, 11 Dec 1999, J.E. Braggins 99/317, AK254565; Colenso s.n., ex herb Steph. in FH as Radula plicata ; South Island: Whataroa Ecological Region, Hokitika Ecological District, Lake Kaniere, Slip Bay, 42°52'S, 171°10'E, 200 m, 25 Nov 1995, J.E. Braggins 95/637B, AK285658;Arthur'S, Pass National Park, Nothofagus solanderi Wald am Bridal Veil Fall Nature Walk bei Arthur's Pass Village, auf Fels am Bach, 800-850 m, 5 Feb 1991, Schäfer-Verwimp & Verwimp, Herb. Schäfer-Verwimp 14336; Westland Land District, Otira River, Arthurs Pass, 730 m, 7 Apr 2004, T. Hay, CHR583489; Haast, 3 miles north of bridge, 30 m, 4 Mar 1972, J. Child, CHR453978; Otago Coast Ecological Region, Dunedin Ecological District, Dunedin City, Leith Valley, Morrisons Burn, 45°50'S, 170°30'E, 220 m 20 Nov 1998, J.E. Braggins 98/335; AK253710; Marlborough, Pelorus River catchment, head of Elvy Stream, 41°18'52"S, 173°34'24"E, 270 m, 12 Feb 2012, M.A.M. Renner 6076, NSW895351; South Westland, Haast Pass, Cross Creek, 44°05'50"S, 169°21'31"E, 560 m, 15 Feb 2012, M.A.M. Renner 6127, NSW895397; South Westland, Haast Pass, track to Brewster Hut, 44°04'49"S, 169°23'24"E, 660 m, 15 Feb 2012, M.A.M. Renner 6137, NSW895439; Westland, Chesterfield, Kapitea Creek, 42°37'20"S, 171°07'29"E, 35 m, 17 Feb 2012, M.A.M. Renner 6180, NSW895508; Westland, Chesterfield, Kapitea Creek, 42°37'20"S, 171°07'29"E, 35 m, 17 Feb 2012, M.A.M. Renner 6183, NSW895511; Westland, Paparoa National Park, Fox River, 42°02'26"S, 171°23'58"E, 20 m, 18 Feb 2012, M.A.M. Renner 6227, NSW895686; Nelson, Kahurangi National Park, Cobb Valley, Round Lake cirque, 41°03'08"S, 172°30'03"E, 1410 m, 19 Feb 2012, M.A.M. Renner 6241, NSW896177; Nelson, Kahurangi National Park, Cobb Valley, Round Lake cirque, 41°03'08"S, 172°30'03"E, 1410 m, 19 Feb 2012, M.A.M. Renner 6244, NSW896179;

Chatham Islands: Chatham Island, Waitangi, Crispans Lane, Unnamed Stream, 43°57'S, 176°33'E, 10 m, 9 Jan 2006, P.J. de Lange CH784 & J.W.D. Sawyer, AK299897;

Australia: Victoria: Otway Range, Carslisle State Park, Carlisle-Gellibrand Road, 38°32'08"S, 143°29'16"E, 11 Jun 1996, D.A. Meagher s.n., MEL2053881; Eastern Highlands, Warburton, 37°42'S, 145°42'E, 28 Feb 1902, R.A. Bastow, MEL1037783.

Tasmania: New Norfolk Municipality, tributary of the Styx River 10 miles west of Maydena, 146°32'E, 42°46'S, 4 Dec 1973, D. Norris 28952, F; Kentish Municipality, cliffs above Lake Barrington near Forth Falls, c. 150 m, 41°24'S, 146°11'E, 15 Nov 1973, D. Norris 27306, F; Upper North West Bay River, 42°55'S, 147°12'E, 12 Feb 1980, A.V. Ratkowsky H1137, HO304430, as Radula plicata det M.-L. So; Mt. Wellington, Upper N.W. Bay River, 42°55'S, 147°13'E, 12 Feb 1980, A.V. Ratkowsky, CANB8205585, as Radula plicata det Yamada; South West, South West Conservation Area, Huon River, adjacent Huon campground, 43°02'17"S, 146°18'12"E, 285 m, 23 Jan 2012, M.A.M. Renner 5936 & E.A. Brown, NSW895267; South West, South West National Park, Mount Eliza, track to summit from Scotts Peak Road, 100 meters above Eliza Hut, 42°57'38"S, 146°24'16"E, 1000 m, 22 Jan 2012, M.A.M. Renner 5916 & E.A. Brown, NSW909267; South West, South West Conservation Area, Mount Eliza, Condominium Creek, 42°57'26"S, 146°21'49"E, 350 m, 23 Jan 2012, M.A.M. Renner 5923 & E.A. Brown, NSW895246; South West, South West Conservation Area, Huon River, adjacent Huon campground, 43°02'17"S, 146°18'12"E, 285 m, 23 Jan 2012, M.A.M. Renner 5940 & E.A. Brown, NSW895272; South West, Princess Creek catchment, east side of Lyell Highway between stream and road, 42°08'40"S, 145°29'18"E, 325 m, 26 Jan 2012, M.A.M. Renner 5989 & E.A. Brown, NSW909286; South West, Mount Dundas Regional Reserve, Manuka River, between Lyell Highway and river, 42°08'36"S, 145°22'56"E, 195 m, 26 Jan 2012, M.A.M. Renner 5998 & E.A. Brown, NSW909293; West Coast, Waratah-Savage River Road, Arthur River catchment, unnamed stream, 41°27'52"S, 145°25' 26"E, 490 m, 28 Jan 2012, M.A.M. Renner 6023 & E.A. Brown, NSW909423; West Coast, Waratah-Savage River Road, Arthur River catchment, unnamed stream, 41°27'53"S, 145°25'22"E, 540 m, 28 Jan 2012, M.A.M. Renner 6024 & E.A. Brown, NSW909424; West Coast, Williamsford Road, Ring River, 41°49'33"S, 145°30'41"E, 405 m, 31 Jan 2012, M.A.M. Renner 6036 & E.A. Brown, NSW909452; West Coast, Williamsford Road, Ring River, 41°49'33"S, 145°30'41"E, 405 m, 31 Jan 2012, M.A.M. Renner 6048 & E.A. Brown, NSW909482.