Radula buccinifera (Hook.f. et Taylor) Taylor ex Gottsche, Lindenb. et Nees Synopsis Hepaticarum 2: 261. 1845.

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 : 26-34

Radula buccinifera (Hook.f. et Taylor) Taylor ex Gottsche, Lindenb. et Nees Synopsis Hepaticarum 2: 261. 1845. Figs 11 -13

Jungermannia buccinifera Hook.f. et Taylor. London Journal of Botany 3: 580. 1844.

Type:

Australia: Tasmania: " Van Diemen’s Land ". Syntypes: Voyage of HMS Erebus & Terror. J.D. Hooker s.n. 1840, BM, K, FH00284039! L, NY00831294! ex herb Möller S-B25060! ex herb Lehmann S-B25061! S-B25062! YU; " Van Diemen’s Land ", Gunn, FH00284037! " Van Diemen’s Land", on Sticta glabra , without collector, date or number, FH00284038!

Radula wattsiana Steph. Species Hepaticarum 4: 211. 1910.

Type: Australia: Cambewarra Mountain, 1903, leg. Rev. W.W. Watts, ex herb. Levier No. 4144 in herb. Steph (as Radula plicata ). Lectotype (designated by Castle (1963 p. 46): G00264951! isolectotypes: FH! NSW764184!

Description.

[from NY00831294 and MEL38047] Forming interwoven mats of shoots, brown in herbarium, shoot systems regularly pinnately branched, with additional pseudodichotomous branching due to production of pairs of subfloral innovations below gynoecia; dimorphic, primary shoots 1.2-1.7 mm wide and up to 40 mm long, secondary shoots smaller in stature and either apparently terminating growth after five to seven leaf pairs, or continuing vegetative growth and attaining similar stature to primary shoots by fourth to sixth pair of leaves; older shoot sectors retaining leaf-lobes.

Stems 130-155 µm diameter, with cortical cells in a single tier of 25-31 rows, medulla cells in 20-35 rows, cortical cell walls yellow-brown pigmented, ventral cortical walls occasionally yellow pigmented, external free cortical cell wall continuously thickened, radial longitudinal cortical walls thin or slightly thickened, inner tangential walls continuously thickened; medulla cell walls faintly yellow-pigmented, with small 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 variable within single individuals, reaching the dorsal stem mid-line or not, leaving zero to three dorsal cortical cell rows leaf-free, dorsal leaf-free strip usually present; leaf insertion not attaining the ventral stem mid-line, leaving two to five ventral cortical cell rows leaf-free. Leaf lobes rotund-ovate, 600-845 µm long by 400-655 μm wide, contiguous, not to weakly falcate, acroscopic base not sharply deflexed away from stem, plane, not interlocking over the dorsal stem surface, stem visible between leaf lobes in dorsal view; margins irregularly but minutely repand, otherwise entire, the interior lobe margin shallowly ampliate, not or only just reaching the opposite stem margin, antical margin curved, exterior margin sharply curved through nearly 100°, postical margin shallowly curved or straight; angle between postical lobe margin and keel c. 135°. Lobules quadrate to rhombic when small and large, one eighth to one sixth the lobe area, 310-475 µm long by 215-420 μm wide; keel curved in small stature lobules, to straight, to arched in large stature lobules, angle between keel and stem 135°, keel turning through 90° mostly at keel-lobe junction, keel apex and postical lobe margin with shallow notch; interior lobule margin free for one third its length, free portion weakly ampliate small stature lobules to moderately ampliate on large stature lobules, extending at most half way across the ventral stem surface; acroscopic margin S-shaped (typical in situ) to straight (when flattened), apical portion inclined toward stem; apex obtuse to acute; free exterior margin straight to curved, occasionally with a small knee above the lobe-lobule junction; margins plane, entire or shallowly repand; lobe-lobule junction slightly antical to, or level with, the acroscopic end of stem insertion; attached to stem along 0.66 of the interior margin, stem insertion more or less linear, gently curved at acroscopic and basiscopic ends, not revolute; 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, 10-23 µm long by 11-19 μm wide; thin walled with small triangular trigones, medial wall thickenings absent; cells of lobe margin smaller than those of middle, quadrate to rectangular, 9-15 µm long and wide, interior and exterior cell walls not differential thickened, cell lumen not bulging medially, leaf lobe cell surface unornamented, smooth. Oil-bodies not known. Asexual reproduction absent. Dioicous. Androecia on lateral branches that usually terminate following production of 2-4 pairs of antheridial bracts, occasionally these branches continue vegetative growth; bract lobules epistatic, keel deeply curved, bucket-like, free apical portion triangular, apex obtuse, inner margin ampliate, plane; lobes rounded, not caducous; antheridia 1-2 per bract. Gynoecia terminal on leading shoots, subtended by two subfloral innovations, usually full-sized and again fertile; archegonia 130-155 µm tall, archegonia neck six cell columns, 6-8 per gynoecium, on a small disc of tissue, interspersed with paraphyses of 1-3 moniliform cells capped by a hyaline papilla, not encompassed by a protoperianth. Female bracts in one pair, symmetrical, tightly imbricate, elliptic-obovate, weakly falcate, lobe 725-845 μm long by 390-610 μm wide, margins entire; lobules rectangular, one half the lobe area, apex obtuse to broadly acute, keel arched, margins entire; bract insertion lines interlocking dorsally and ventrally, insertion equitant. Perianths 2670-3650 µm long and 630-930 µm wide at mouth, mouth repand, more or less parallel sided for upper third, then tapering to tubular stem perigynium comprising the lower third to half, faint bulb in basal third, broadest c. one third from mouth where 660-950 µm wide, walls bi- or tri-stratose at junction with perigynium, unistratose above, cell walls with triangular trigones. Long stem perigynium present, 5-6 stratose throughout, cell walls not thickened or brown-pigmented, hyaline, perianth-calyptra junction elevated above female bracts on 9-15 tiers of cells. Calyptral perigynium present, 2-4 stratose at base, unistratose above, unfertilised archegonia elevated on surface of calyptra.

Etymology.

Horn-bearing.

Distribution and ecology.

Radula buccinifera is endemic to Australia, where it is widespread throughout Tasmania, and Victoria but more restricted in New South Wales, being confined to the eastern side of the Great Dividing Range and associated escarpments. In Western Australia it is confined to the far south-west. The northern limit of this species has not been identified, currently the northernmost locality is Point Lookout in New England National Park, in association with Nothofagus moorei . Cool temperate rainforests dominated by Nothofagus also occur in south-east Queensland and Radula buccinifera may be be found in these areas.

Casual field observation suggests the elevational range occupied by Radula buccinifera is correlated with latitude. In Tasmania and Victoria Radula buccinifera occurs across a broad elevational range from sea level to 1500 m,encompassing a range of habitat types from lowland to montane forests, including wet sclerophyll forest and cool temperate rainforest, wetlands, and in alpine scrub and within habitats may occupy a range of microsites from twigs, branches, tree trunks and tree bases, exposed tree roots on the forest floor, to rotting logs, exposed soil on forest banks, dripping rocks adjacent waterfalls, and on rocks within stream beds, sometimes under running water.

The ecological range decreases with latitude; the further north, the more restricted to foothills, escarpments, and mountains in association with cool temperate rainforests. This altitudinal contraction is associated with restriction in the diversity of microhabitats occupied, such that at the northern end of its range, where Radula buccinifera occurs at 1000 m or higher, and is always encountered as a lithophyte most frequently on vertical rock faces associated with bluffs and outcrops.

Radula buccinifera is one of three [ Radula novae-hollandiae , Radula strangulata , Radula buccinifera ] Radula species in south-eastern Australia that may be found growing under running water. On rocks it frequently co-occurs with Radula novae-hollandiae , forming mixed mats. The type specimen of Radula novae-hollandiae comprises such a mixed patch. In north-eastern New South Wales it may co-occur and form mixtures with a number of other Radula species, initially presenting as confusing and exceptionally variable individuals.

Variation.

Individual populations exhibit generous amplitude in shoot stature and lobule shape. Lobules vary from rhombic-quadrate with little ampliation of the interior margin and a straight antical margin, to quadrate with a pronounced ampliate interior margin and an S-shaped antical margin having a distinct, obtuse, lobule apex. Colour of individuals varies, from glaucous yellow-green to dark green, in part this appears correlated to microhabitat, with epiphytic plants tending yellow green, and lithophytic plants being dark green.

The amplitude of morphological variation expressed between individuals appears negatively correlated with latitude. The greatest morphological variation between individuals in plant colour, shoot size and lobule shape occurs in the southern end of the distribution. Individuals vary from brown-green, mid-green and glaucous green and from small to large, with associated differences in lobule shape. In Central and Northern Coast, and Northern Tableland regions in the northern part of Radula buccinifera range, individuals are fairly consistent in morphology being large, mid-green, and having lobules whose antical margin is more or less straight, with an pronounced ampliate free interior margin.

Recognition. Despite the variability exhibited by Radula buccinifera this species is relatively easy to recognize. The first clue to identity comes from the habitat and microhabitat the plants occupy; Radula buccinifera inhabits forest interiors, frequently in microsites on or close to the forest floor that are often well shaded. Other species of the Radula buccinifera complex with which Radula buccinifera could be confused on morphological grounds almost all occupy quite different habitats and microsites.

Radula australiana inhabits subalpine and alpine habitats typically dominated by shrubland, tussockland or grassland, and is almost always found above tree-line. However, Radula australiana and Radula buccinifera may co-occur in forest interiors on forested mountain tops in Victoria and Tasmania, particularly in association with exposed large granite boulders, for example at Mt. Ellery in Errinundra National Park, Victoria. The most accessible morphological character by which Radula buccinifera differs from Radula australiana is the shape of the leaf lobes, where the junction between the lobe and keel forms a simple angle in Radula buccinifera and the leaves are not or weakly falcate as a result. In Radula australiana the keel runs more or less seamlessly into the lobe outline, the two following the same curve. If this character proves ambiguous, lobule shape is a good source of discriminating characters. Lobule shape should always be assessed on the basis of hydrated, slide-mounted material, and at least 50 × magnification. When hydrated the lobules of Radula buccinifera are quadrate to rhombic, and one eighth to one sixth the lobe area. The antical margin is usually (northern plants excepted) S-shaped in situ (straight when flattened), but this is more pronounced with a deep medial curve, and the apex is obtuse. The lobules of Radula australiana are quadrate to rectangular, and one quarter the lobe area. The antical margin is S-shaped in situ and when flattened, but the medial curve is relatively shallow, and the apex varies from obtuse to acute. Further differences exist in the lobe marginal cells that bulge in Radula buccinifera , but are crenulate, due to medial thickening on the external cell wall in Radula australiana . Finally, if all of these 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 buccinifera 0-3 cortical stem cell rows are leaf free, whereasno cell rows are leaf-free in Radula australiana . Obviously multiple counts from a few shoots are required. If none of these characters leads to a satisfactory conclusion, consider the possibility that the material at hand comprises a mixed collection.

Radula buccinifera may be confused with Radula demissa , with which it co-occurs in Victoria and Tasmania where the two species may occupy similar microhabitats. For characters distinguishing these two species, see the recognition section of Radula demissa .

Radula buccinifera could also be confused with Radula strangulata , as the two species occupy similar microhabitats, both have a dorsal leaf-free strip, and are identical in overall appearance. Lobule shape differences are possibly the best, and only, source of discriminating morphological characters. In Radula buccinifera the antical lobule margin is typically straight and perpendicular to the stem and the lobule apex is obtuse at least in large lobules. In contrast the antical margin of Radula strangulata is either inclined toward the stem or arched and the lobule apex is broadly acute in large lobules. Aquatic morphs of Radula strangulata have longitudinally rectangular lobules, whose straight exterior margin runs parallel with the stem and whose interior margin is ampliate over the ventral stem surface. In Radula buccinifera the lobule is trapeziform or quadrate, and not ampliate to the same degree. More substantial characters have not been identified. Some interpretations of morphology would find (and have found) strong evidence for the synonymisation of Radula buccinifera with Radula strangulata from this circumstance. However, here as in other liverwort complexes morphological unity and the existence of morphological continua are immaterial to the phylogenetic structure underpinning the observed morphological diversity, and the existence of populations of both species intermediate in size and shape does not indicate phylogenetic unity ( Renner et al. 2013).

The name Radula buccinifera has been applied to specimens belonging to a wide variety of unrelated species from tropical Queensland, even as far north as Moa Island in the Torres Strait (i.e. CANB9500180). However, with the exception of two specimens collected by Pentzke in 1882, one of which is a mixture of elements from at least two locations (see discussion of the type of Radula mittenii ), all records of known provenance from tropical Queensland are based on misidentifications. Radula buccinifera can be distinguished from virtually every species in Queensland by 1) the presence of a stem perigynium in the perianth, 2) the production of two subfloral innovations, 3) the absence of secondary cell wall thickening in the stem medulla, 4) the sub-dimorphic shoot systems, 5) the rhombic lobules, 6) the presence of two papillae on the interior lobule margin, 7) the smooth leaf-lobe cell surfaces. Radula buccinifera may be confused with Radula notabilis , and Radula imposita ; for distinguishing characters see the recognition section of those species.

Remarks.

Radula buccinifera is among the most widely misunderstood species of Radula , having been confused in herbarium collections with almost every other Radula species in Australia, and several that aren’t. Misidentifications of Radula buccinifera have been the basis for several records of species now excluded from the Australian flora, including Radula physoloba and Radula plicata . Confusion with Radula physoloba seems attributable to R.A. Bastow, all of whose specimens identified as Radula physoloba in MEL ex herb. R.A. Bastow are Radula buccinifera , and confusion with Radula plicata is attributable to Stephani, who misidentified several gatherings by Weymouth and Bastow.

Radula buccinifera was understandably confused with Radula strangulata by Mitten (1855) who cited collections by Colenso, Stephenson, and Lyall from the southern North Island in his treatment for Hooker’s Flora of New Zealand, and this confusion has been perpetuated in various manifestations. Allison and Child (1975) illustrated Radula strangulata as Radula buccinifera , and again in the next figure as Radula levieri Steph. Renner (2005) made a different, and less forgivable, error when he presented a key to New Zealand species that included Radula buccinifera . The plants he referred to were actually Radula demissa M.A.M.Renner, and his error was based on mis-interpretation of type material.

Castle, however, appears to have been hopelessly confused regarding the identity of Radula buccinifera , arbitrarily placing specimens of Radula buccinifera , Radula demissa , and Radula strangulata under the names Radula buccinifera , Radula mittenii , and Radula wattsiana . As an example, the figure illustrating Radula buccinifera in Castle (1967) includes a male shoot of Radula strangulata . Castle maintained that Radula mittenii differed from Radula buccinifera in its more falcate leaf lobes, and the shorter perianths and he may have had Radula demissa in mind in this assessment.

Yamada (1984) synonymised Radula mittenii with Radula buccinifera on the basis of his examination of ‘holotype’ material held in g, and his assertion that the differences observed were environmental. Yamada’s identification of holotype material is incorrect, his interpretation was derived from examination of duplicate material in Geneva and the implications of this are discussed below under Radula mittenii . The synonymisation of Radula mittenii , having a type from tropical Queensland, with Radula buccinifera by Yamada (1984) seems to have encouraged the application of the name Radula buccinifera to a range of specimens from tropical Queensland, to the point where nearly every species occurring in Queensland has, at some point, been identified as Radula buccinifera . Beyond the specimen collected by Pentzke, no populations attributable to Radula buccinifera from tropical Queenslandexists within herbaria, and the species has not been observed during the course of recent fieldwork in the Wet Tropics of Queensland.

Nomenclature.

So (2005) identified a Hooker collection from Van Diemens Land as the holotype of Radula buccinifera . This is inappropriate as Hooker and Taylor (1844) did not identify a single type collection, the only specimen details given in the protologue were "Van Diemen’s Land". Collections from "Van Diemen’s Land" by both R. C. Gunn and J. D. Hooker were made prior to 1844 and were studied by Hooker and Taylor for their contribution to Hepaticae Antarcticae, as made clear in the subtitle to Hooker and Taylor’s (1844) publication, and by the presence of duplicates from both collectors in Taylors own herbarium. At least three gatherings may comprise the syntype series, being the more or less pure gatherings made by Gunn and Hooker, and a gathering on Sticta , whose collector has not been recorded. Furthermore, Castle (1967) had attempted to lectotypify Radula buccinifera , though we are unclear about exactly what Castle intended in his lectotypification. Castle’s (1967) lectotypification of Radula buccinifera is somewhat obscured by his own commentary. In the list of specimens examined he states 'Van Dieman’s Land, the Type (BM K and Y) and Van Dieman’s Land, R.C. Gunn (BM and K)' [italics ours]. Here Castle appears to identify two groups of specimens, one of which was collected by Gunn and was not identified as the lectotype. But Castle then notes 'we may assume that those collections which bear the label Van Diemen’s Land are portions of the collection upon which Thomas Taylor based his Jungermannia buccinifera . Several samples of this type, preserved in the herbaria of the British Museum and of Kew Gardens, also include in the label the name Gunn. Gunn presumably made the type collection as his name, as collector, appears in the title of the article in which the original description of Jungermannia buccinifera was published, without data’. Here Castle implies that the Gunn collection should be regarded as the lectotype. All gatherings comprising the syntype series we have seen, being duplicates of both Hooker and Gunn’s collections, and the specimen sine. coll. are unambiguously assignable to Radula buccinifera , so application of the name Radula buccinifera will remain the same regardless of which gathering and which particular specimen the species is lectotypified on. Castle’s lectotypification at best requires narrowing because he did not identify a single specimen as type, and at worst lectotypification requires repeating because it is not quite clear whether Castle identified a single gathering.

Specimens examined.

Australia: New South Wales: Northern Tablelands, The Cascades, Point Lookout, New England National Park, 30°30'S, 152°24'E, 1010 m, 11 March 1990, A.K. Brooks 204 & E.A. Brown, NSW233743; Northern Tablelands, Washpool National Park, Coombadjha Stream catchment, Washpool walk. Between Bellbird Campground and Coombadjha Stream, 29°28'01"S, 152°19'19"E, 800 m, 11 Apr 2011, M.A.M. Renner 5246, NSW875783; Northern Tablelands, Mount Hyland Nature Reserve, Mount Hyland Circuit Track, southern summit, 30°10'30"S, 152°25'38"E, 1380 m, 13 Apr 2011, M.A.M. Renner 5257, NSW875805; Northern Tablelands, New England National Park, Point Lookout area, Lyrebird Track between Banksia Point and Weeping Rock, 30°29'24"S, 152°24'32"E, 1470 m, 16 Apr 2011, M.A.M. Renner 5288, NSW875835; North Coast, Myall River State Forest, Strike-a-light camping area, 32°17'S, 152°05'E, 210 m, 5 Apr 2002, E.A. Brown 2002/18 & B.J. Conn, NSW491702; Central Tablelands, Blue Mountains National Park, Waterfall Reserve, Waterfall track, 33°31'S, 150°22'E, 860 m, 24 September 2001, N. Klazenga & V. Stajsic 2809, MEL2137146; South Coast, Rutherford Creek near Piper'S, Lookout, c. 12 km WNW of Bemboka, 36°36'S, 149°27'E, 700 m, 17 August 1985, K.R. Thiele 1001, MEL2273913. Central Tablelands, Mount Victoria, track at Thomas Mitchell Monument Hill, 33°34'S, 150°15'E, 13 Mar 1989, E.A. Brown 89/35, NSW436068; Central Coast, Nowra, Cambewarra Mountain, rocks near Cambewarra Lookout, 34°47'56"S, 150°34'36"E, 470 m, 6 Jun 2011, M.A.M. Renner 5303 & E.A. Brown, NSW877190; Northern Tablelands, Barrington Tops National Park, Dilgry River, Devils Hole Campground., 31°54'55"S, 151°28'59"E, 1400 m, 16 Dec 2011, M.A.M. Renner 5868, NSW898654;

Victoria: Wilsons Promontory, headwaters of Blackfish Creek, Wilsons Promentory National Park, 39°02'S, 146°23'E, 27 July 1996, D.A. Meagher s.n., MEL240137; Gippsland Highlands, Tarra Bulga National Park, Bulga Section, Suspension Bridge circuit, 38°26'S, 146°34'E, 650 m, 21 Feb 1997, A.W. Thies FN1626K, MEL241693; Eastern Highlands, Toolangi/Black Range State Forest, Sylvia Creek Road, Wirrawilla rainforest walk, 37°31'S, 145°31'E, 650 m, N. Klazenga & V. Stajsic 2325, MEL2111876; Otway Range, upper Calder River, 8 miles west of Apollo Bay, 38°45'S, 143°31'E, 19 Nov 1995, J.H. Willis s.n., MEL1514803; Otway Range, Grey River Road, Angahook-Lorne State Forest, 28 km NE of Apollo Bay, 38°38'S, 143°46'E, 480 m, 5 Dec 1996, H. Streimann 58978, MEL2300394; Gippsland Plain, Tara Bulga National Park, near Yarram, Gippsland, 38°34'S, 146°40'E, Aug 1960, K. Healey s.n., MEL38047; Grey River Reserve, Angahook - Lorne State Park, 17 km ENE of Apollo Bay, 38°39'S, 143°49'E, 280 m, 5 Dec 1996, H. Streimann 58874, CANB9802545; East Gippsland, Errinundra National Park, Errinundra Road, between Ada River and Errinundra Saddle, 37°20'47"S, 148°51'43"E, 921 m, 04 Mar 2011, M.A.M. Renner 5176 & E.A. Brown, NSW875959; East Gippsland, Errinundra National Park, Errinundra Road, between Ada River and Errinundra Saddle, 37°20'47"S, 148°51'43"E, 921 m, 04 Mar 2011, M.A.M. Renner 5177 & E.A. Brown, NSW875960;

Tasmania: Furneaux Group, Flinders Island, 660 m east of summit of Big Badger Hill, 40°02'S, 148°01'E, 137 m, 29 Jul 2004, J.S. Whinray B1861, MEL2209244; North West, Dismal Swamp Nature Reserve, 41°59'S, 144°51'E, 40 m, 23 Mar 2000, A. Moscal 30979, HO558827; Southwest National Park, Styx Valley, Big Tree Reserve, river walk (7 km SSE of Maydena), 42.81354°S, 146.65559°E, 350 m, 6 Dec 2007, B. Shaw 6463, DUKE; Marriots Falls Track paralleling Tyenna River (4.5 km NE of Maydena, 42.727°S, 146.663°E, 230 m, 6 Dec 2007, B. Shaw 6511, DUKE; Mt Field National Park, Growling Swallet (E of F8 East Road, NNE of Florentine Road), ca. 70 km WNW of Hobart, 42.687°S, 146.496°E, 580 m, 4 Dec 2007, B. Shaw 6287, DUKE; B. Shaw 6351, DUKE; Junee Cave State Park, 3 km NW of Maydena, 42.737°S, 146.596°E, 300 m, 7 Dec 2000, B. Shaw 6619, DUKE; Junee Cave State Park, 3 km NW of Maydena, 42.738°S, 146.597°E, 300 m, 4 Dec 2000, B. Shaw 6361, DUKE; Southwest National Park, Styx Valley, Big Tree Reserve, river walk (7 km SSE of Maydena), 42.812°S, 146.657°E, 350 m, 6 Dec 2007, B. Shaw 6476, DUKE; Scottsdale to St. Helens Highway, 147°40'E, 41°10'S, 800 m, 11 Jan 1974, D. Norris 31903, F; Fingal Municipality, St. Mary's Pass, 148°12'E, 41°34'S, 12 Jan 1974, D. Norris 32308, F; Deloraine Municipality, Lyons Creek 8 miles W of Liena, c. 700 m, 41°33'S, 146°12'E, 15 Jan 1974, D. Norris 32735, F; D. Norris 32744, F; D. Norris 32754, F; Scotts Road, 41°42'S, 146°34'E, 740 m, 10 Nov 1991, J. Jarman s.n., HO310154; Tasmania merid. Mt Wellington, New Town Falls, 17 August 1889, W.A. Weymouth, as Radula plicata det. Stephani, FH00284645; Tasmania, Watts, syntype of Radula wattsiana Steph. ex herb. Steph. FH00284647; South West, Waterfall Creek State Reserve, South Bruny Range, 42°24'S, 147°19'E, 100 m, 28 Apr 1993, A. Moscal 25083, HO558839; Mt. Field National Park, Lake Dobson Road, near Horseshoe Falls, 42°41'S, 146°42'E, 400 m, 7 Dec 1988, J.A. Curnow 2602, HO304660; South West, Deadmans Bay, south coast, 43°32'S, 146°30'E, 5 m, 17 Jan 1987, A. Moscal 14077, HO558830; 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 5939 & E.A. Brown, NSW895271; Central Highlands, Jerusalem Walls National Park, Southern side of Mt Jerusalem, 41°49'23"S, 146°18'02"E, 1315 m, 29 Jan 2012, M.A.M. Renner 6025 & E.A. Brown, NSW909425; Mersey River, below Lake Rowallan, 41°41'56"S, 146°13'08"E, 440 m, 30 Jan 2011, M.A.M. Renner 6027 & E.A. Brown, 30 Jan 2012, NSW909430; Mersey River, below Lake Rowallan, 41°41'56"S, 146°13'08"E, 440 m, 30 Jan 2012, M.A.M. Renner 6032 & E.A. Brown, NSW909436;

Western Australia: Mt Chudalup, 17 km SSE of Northcliffe, 34°46'S, 116°05'E, 185 m, 14 Sep 1994, H. Streimann 54341, CANB9504479, PERTH04957172; Denmark Shire, mid-slopes on the east side of Mt Hallowel on the Bibbulmun track, 35°0'3"S, 117°19'3"E, 14 Aug 2000, B.G. Hammersley 2598, PERTH05803047; Cascades S of Pemberton, 16 Dec 1973, N.G. Marchant 73/45, PERTH01929305; NE of Mount Frankland, 34°49'17"S, 116°47'36"E, 26 Aug 1997, K.A. Redwood 612, PERTH04983963; Stirling Range National Park, Stirling Range National Park, Toolbrinup track, boulder slope at base of hill, 34°23'11"S, 118°02'59"E, 765 m, 25 Aug 2009, E.D. Cooper 09/067 & E.A. Brown, NSW970847; Stirling Range National Park, Stirling Range National Park, Toolbrinup track, boulder slope at base of hill, 34°23'11"S, 118°02'59"E, 765 m, 25 Aug 2009, E.D. Cooper 09/068 & E.A. Brown, NSW970854.

Figures

Figure 11. Radula buccinifera line drawings 1. A Five lobules from secondary shoots showing variation in size and shape B Five lobules from primary shoots showing variation in size and shape C Cellular detail of free interior lobule margin D Cellular detail of lobule apex E Ventral view of male shoot F Ventral view of female shoot G Dorsal view of shoot. Scale bars: A, B: 240 µm. C-D: 60 µm, E-G: 600 µm. A-E from MEL 2054447, F-G from NY 00831294.

Figure 13. Radula buccinifera pictures. A Ventral view of shoot B Mature perianth. C-E: Ventral view of shoots and lobules F Gynoecium G Androecium H Transverse sections of stems from primary shoot I Leaf-lobe marginal cells G NSW 875961, other NSW 909436.