Nephroma australe Rich. var. australe f. homalodes (Nyl.) Murray, comb nov.

Nephroma homalodes Nyl, Lich N.Z, 43 (1888).

Gyeln., Ann. Musei Nat. Hung., Pars Bot., 11 (1935).

Nephroma antarcticum Hellbom, Bihang Kgl. Svensk Vetensk. Akad. Handl. 21, III. (13), 26 (1896).

Nephroma zelandicum Gyelnik, Borbasia, I, 4 (1938).

Nephroma zelandicum f squamicolum Gyelnik, Borbasia, 1, 5 (1938)

Nephroma neozelandicum Gyelnik apud Zahlbruckner, Lich N. Z. 45 (1941), (? lapsus).

Thallus more or less orbicular where substrate allows, 2 to 10 cm dia, variously lobed with lobes ascending at least at tips and sometimes more or less imbricated, 100–150 (-200)μ thick, upper cortex 30–40μ thick (up to 60μ and rugose on backs of apothecia), algal layer 15–20μ thick with algal cells mostly 8μ dia, medulla white (rarely pink in parts of some specimens), K +, P -, C +, pink or -, lower cortex pseudoparenchymatous 6–10μ thick. Hymenium 60–90μ thick including epithecium, hypothecium very pale straw-coloured 20–40μ thick, asci 55–65 × 10–12μ, 8-spored, usually with spores in two series. Spores light reddishbrown, 3-septate, mostly 19–21 × 5½–7μ. Paraphyses 2μ thick.

Habitat. As for var. australe.

Distribution. New Zealand. North Island: Kaipokirikiri (Colenso 820), Ruamahanga R (Colenso 2616); Napier (Colenso 3576 pr.p); Taruarua (Colenso 4733); Manawatu (Colenso 5084), Te Hawera (Colenso 5112); Patangata (Colenso 5268); all in WELT Nelson: Hundalee, Mr. 1264. Marlborough. Tophouse, Mr. 4236 Canterbury. Geraldine, 031; Niger River, 3,500ft, Sc 54; Bealey R, 3,000ft, Mr. 6919; Hook Bush (A. D. Campbell) 3700. Otago: Huxley River 1845, 1846;, Routeburn Valley 0812, 0936, 1041; Greenstone Valley (Holloway) CHR (G 51 sub N. neozelandicum f. squamicolum); Eglinton Valley, Otago University Bot. Dept. 004590 and 004591; vicinity Dunedin, T 157, T 304, T 429, T 580, T 742; T877, T 1085, T 2384, CHR (G4=T 429, G5=T 580, G6=T159, G7=T 304, G54=T 157 + T 742, all sub N. neozelandicum); Maungatua 1, 200ft, Mr. 933; Akatore, Mr. 575, Mr. 576 (pr p), Mr. 573, 1500; Taieri Mouth, Mr. 769, Mt. Charles, 600ft, 3523 (on soil). Southland: Waihopai, Mr. 6912; Wilmot Pass, 2,000ft, 3923 (K nearly-). Stewart Island: Paterson's Inlet, Mr. 697 (K nearly-), Ulva, Mr. 696.

According to Nylander (1888) N. homalodes differs from N. australe in the larger size, more rugose cortex above the apothecia, larger spores, partly red medulla and orange reaction with KOH. N. zelandicum differs from australe in the yellow reaction of the medulla with KOH N. neozelandicum is apparently an error for N. zelandicum N. Homanu, a form of australe with rather closely appressed lobes and marginal squamules was based on specimens from Juan Fernandez and Stewart Island N. zelandicum f. squamicolum is a K + form corresponding to N. Homanii.

Although specimens corresponding to each of these “species” can be found, examination of a considerable range of material shows that there are no clear morphological differences between them. A positive KOH reaction is not associated with larger specimens, and is in any case rather variable, being usually yellow rather than orange. One specimen, Mr. 575, develops a strong orange colour with separation of crystals in a few seconds, but in others the colour is yellow, changing to orange after 15 minutes or more. The reddish medulla described for N. homalodes appears to be an artefact and the condition can be seen in some lobes of specimens T 429, Mr. 573 and 1845. The normal range of spore size for N. australe covers the dimensions given by Nylander for australe and homalodes; I have not been able to associate larger spores with a positive KOH reaction. I have preferred, therefore to treat homalodes as a chemical form of australe. Specimens of N. australe vary considerably in form from large, broad-lobed plants on moss or rock to small lobed plants with marginal lobules or squamules growing on twigs. The production of marginal squamules (“isidia” of Gyelnik) appears to be of no taxonomic significance, it is apparently associated with regeneration following “insect

damage”, and may be found on parts of otherwise normal plants. N. pruinosum (Mont) Zahlbr. (= N. schizocarpum) is apparently known only from the type specimen collected by Hombron in 1839 on Banks Peninsula. Montagne's diagnosis states that the apothecia are white-pruinose and the plant “olivaceo-fuscus”, while Nylander gives the pruina as a doubtful character and renamed the species schizocarpum in reference to the longitudinally divided apothecia; he put it in the subgenus Nephroma (bright green algae). Gyelnik mentions neither pruina nor divided apothecia and describes the algal cells as green but nostocaceous, transferring the species to subgenus Nephromium. Gyelnik apparently used the size of the cells as the most important character for distinguishing between the green and bluegreen symbiotic algae in this group, but this is an unrehable character, and Lamb (1955) for instance has noted that N. analogum has Protococcoid algae, not Nostoc as stated by Gyelnik. On the assumption that Gyelnik made a similar mistake with regard to N. pruinosum I have referred it to N. australe var. australe. The microscopic structure (sec. Gyelnik) does not differ appreciably from that of australe. Although divided apothecia are sometimes seen in australe they seem to be due to drying and cracking since the edges are not, or hardly corticate. Possibly pruinosum is a form resembling the type specimen of var. rigidum with some apothecia entire and some divided. Although pruinosum is said to differ from australe in having the cortex on the backs of the apothecia almost smooth, this is seen also in australe occasionally.

Both forms of australe seem to be widely distributed in New Zealand in forested areas and are particularly common in the beech forests bordering the Southern Alps.

Nephroma australe Rich. var. rigidum Murray, var. nov.

Thallus lobatus, superne viridis, laevis; subtus albus, glaber, laevis, cortex superior hyalinus vel pallide fuscescens, 25–30μ crassus; stratus gonidiale 20–30μ crassum, gonidiis viridibus ad 12μ magnis; stratum medullare ca 200–220μ crassum, K -, hyphis 2μ crassis; cortex inferior 5–8μ crassus, hyalinus, pseudoparenchymatus. Apothecia rotundata, 5–10 mm dia; integra aut lobata vel bis terve divisa; hymenium ca 120μ crassum epithecium includendum, hypothecium pallidum 20–40μ crassum; medulla sub hypothecio 100–250μ crassa et stratum gonidiale illic ad 40μ crassum gonidiis ca 10μ dia; cortex in dorsis apotheclorum continuus, 40μ crassus; paraphyses conglutinati ca 2μ dia; asci cylindrici pro majore parte 75 × 10μ 8-spori, sporae 4-blastae, 15½–17½μ longae et 6–6½μ crassae, pallide refuscentifuscae Pycnidia globosa, brunnea, semi-immersa in margine thallino aut rare subtus, pycnidiosporae non visae.

Habitat. Westland: Kokatahi River, on soil, Scott 142 (Type in my herbarium with isotype in Scott's herbarium).

Var. rigidum differs from var. australe in the thicker, flatter and more rigid thallus, in the higher hymenium and the generally longitudinally divided or lobed apothecia. The spores appear somewhat smaller and the backs of the apothecia smoother than usual for var. australe. N. antarcticum (Jacq.) Nyl. differs in having a foveolate upper surface and in having rather larger spores.

A specimen from Otago, Huxley River (1844) approaches this variety in the thickness of the thallus (180–220μ) and height of hymenium (90–100μ), but has the smaller gonidia (5–8μ dia) and larger spores (19½–22 × 7½–8½μ) of var. australe. The specimen has the general appearance of var. australe also in the ascending lobes and rather dissected margins centrally; the backs of the apothecia are strongly rugose and their margins ragged and lobed apparently due to regeneration following insect damage.

• Nephroma cellulosum (Sm) Ach. var. cellulosum. Lichen cellulosus Sm. apud Ach., Method Lich., 289 (1803).

• Nephroma cellulosum Ach., Lich Univ., 523 (1810).

• Hook, Handb. N.Z. Flora, 566 (1867).

• Nephromium cellulosum Nyl., Synopsis Lich. Vol. I, 321 (1860).

Thallus about 5 cm in diameter, livid chestnut to grey, reticulate-foveolate above, foveolae 2–3 mm wide (smaller towards the periphery), white below darkening to blackish centrally and more or less bullate, glabrous. Apothecia light to dark brown, 5–10 mm broad by 2–5 mm long, with a very narrow entire margin, cortex above apothecia not differentiated from that of the thallus proper, 30–40μ thick; algal layer 50–75μ thick with bluish to olive green algal cells 8μ in dia; medulla 50–250μ thick of loosely woven crystal-encrusted 2 -½ dia hyphae; hypothecium hyaline to faintly brown 25–35μ thick, hymenium 75μ thick, including a thin brownish epithecium. Asci 6–8 spored, cylindrical-clavate with spores in two series; spores pale fuscous, 14–19½ × 7–8μ, 3-septate Immature spores are hyaline, smaller and 1-septate.

Habitat. On trees, rarely mosses.

Distribution, Tasmania, New Zealand, Juan Fernandez, South America. Canterbury: Upper Godley River, 3,000ft, Sc 177, Sc 200, Sc 201; Selwyn Gorge (Beckett 7), Canterbury Museum. Otago: Kaka Point, 0875; Matukituki Valley, 1,700ft (D Scott et al.) 4392 Australia: Mt. Macedon, Victoria (Wilson in Knight coll.) WELT.

There is no definite record in the literature of this species from New Zealand, but it has been attributed to this country on Nylander's statement (1860) “quoque in Tasmania et Nova Zelandia”. This appears to derive from Babington (1855), who says “N. cellulosum may probably occur in New Zealand”. The New Zealand specimens closely match the description (Wilson, 1893) and illustration (Babington and Mitten, 1860) of the species, which is apparently uncommon here.

Nephroma cellulosum (Sm.) Ach. var. isidioferum Murray var. nov.

Thallus 5–15 cm latus, lobatus, castaneo-lividus, superne reticulato-costatus, foveolis ca 3 mm latis, versus centrum isidus squamiformibus in venis; isidia thallo concolora, 0 2 mm crassa et 0 1–0 5 mm dia. Apothecia fusca vel rufo-fusca, 5–10 mm lata et 4–8 mm longa, rotundata bis reniformia, margine integro et tenue. Cortex superior ca 25μ crassus, K—, ex cellulis major minusve oblongis, 2 × 5μ; stratum gonidiale subcontinuum, in rugas excepto, 25–40μ, crassum, gonidis sordide viridibus 5μ dia, medulla K—, 100–25μ crassa; cortex inferior parum distinctus 5–10μ crassus, saepe traversim diffractus. Hymenium ca 65μ altum, hypothecium hyalinum, 25–35μ crassum, asci 55 × 16–19μ, 6–8 spori, sporae lente rufofuscescentes, 4-blastae, (16-) 19 (-22½) × (5½-) 6½ (-8)μ, paraphyses non septati, conglutinati, 1½μ crassi sunt Pycnidia non visa.

Distribution. Otago: Flagstaff, 1,200ft, on Fuchsia excorticata T 1884 (Type). Southland: Sutherland Falls, T 2918.

The variety does not seem to differ very significantly from N. cellulosum except in the presence of isidia. The colour of the thallus varies from a slightly brownishgrey in the shade to a dark reddish brown in the sunlight. It evidently closely resembles N. lepidophyllum (Ras) Gyelnik, a South American species which has finer reticulate markings on the upper surface.

• Nephroma helveticum Ach. var. helveticum

• Nephroma helveticum Ach., Lich Univ., 523 (1810).

• Gyelnik, Ann Muses Nat. Hung. Pars Bot., 11 (1935).

• Du Rietz, Archiv f. Bot., 22A, No. 13, 5 (1929).

• Nephromium helveticum Nyl, Lich N. Z., 43 (1888).

• ?Nephroma resupinata var. papyracea Ach of Hook. Fl. N. Z. II, 272 (1855).

• ?Nephroma resupinata Mont. apud D'Urv., Voy de Decouv Pôle sud Botanique I, 192 (1842–5).

Thallus usually small, 1–5 cm dia, grey to livid-brown, thin, with subcylindrical more or less evenly distributed isidia above, and sometimes with more or less marginally distributed squamules about 1 mm wide; sometimes slightly pubescent above, particularly on the backs of the apothecia; tomentose below with tomentum of erect hyphae 01–1 mm long. Apothecia nearly round to reniform, with narrow, usually dentate thalline margin; hymenium 75–85μ high, hyaline except for pale brownish epithecium; hypothecium 30–40μ, light brownish, medulla and algal layer ca 180μ thick above apothecia, loosely woven centrally, cortex on backs of apothecia up to 40μ thick, slightly brownish; asci 52–60 × 12–14μ, 4–6 spored; spores light reddish-brown, 3-septate, 21–25 × 7½–9½μ, with end cells longer than central pair.

Habitat. Tree trunks, rarely on damp rocks

Distribution. Apparently in most parts of the North and South Temperate Zones. Otago. Huxley River 1847, Routeburn Valley, 0813 Southland Waikaia, Mr. 6913.

The species is frequently classed as a variety or form of N. resupinatum Ach, from which it differs in the constantly shorter tomentum, the absence of pseudocyphellae and probably the presence of isidia on the upper surface (cf. Du Rietz 1924; Gyelnik 1935, p. 11). Possibly this is the best treatment, but I have not seen enough foreign material to judge, true N. resupinatum does not seem to occur in New Zealand. N. helveticum appears to be variable as regards the presence or absence of marginal squamules so far as can be determined from the few specimens so far collected in New Zealand. The New Zealand specimens seem to agree with the type specimens of N. helveticum better than do the normal European plants, although Acharius' specimens were collected in Switzerland (Gyelnik, 1935) It is probable that N. tropicum (Mull Arg) Zahlbr and N. denticulatum (Wain) Gyelnik are synonyms of N. helveticum, the former having the backs of the apothecia pubescent and the latter not In the New Zealand specimens this feature is variable even within the same specimen, so cannot have any taxonomic significance. This pubescence is a microscopic feature, and is only visible macroscopically in a matt appearance of the surface.

Nephroma helveticum Ach. var. rufum (Bab) Murray, comb nov.

• Nephroma resupinatum var. rufa Bab. in Hook., Fl. N. Z., II, 272 (1855).

• Nephromium laevigatum var. rufum Nyl., Synops. Lich. I, 321 (1860).

• Nephromium laevigatum Hook., Handb. N. Z. Fl, 566 (1867).

• Kremph Rcose der “Novara”, Bot. I., 121 (1870).

• Nephromium helveticum var. rufum Nyl. apud Hue, Nouo Archiv. du Museum, ser. 3, Vol II, 310 (1890)

• Nephromium helveticum Hellbom, Bihang Kgl. Svensk. Vetensl. Akad. Handl., 21, III (13), 27 (1896).

Thallus thin, 60–170μ thick, usually dark reddish-brown, mostly appressed to the substratum centrally, 1–10 cm dia; peripheral lobes 3–8 mm wide by 3–12 mm long, ascending, upper surface with scattered isidia singly or in groups, lower surface smooth or slightly wrinkled, shining or matt or microscopically pubescent. Isidia terete, 02 mm dia, by 05 mm high or subsquamulose, 1 mm dia. Upper cortex 12μ thick, slightly brownish and sometimes with minute pubescence of single hyphae 5–15μ, high; algal layer 10–25μ thick, algae (?) Nostoc mostly 3½ × 4½μ oblong; medulla 15–100μ thick, of rather loosely woven hyphae 2½μ in dia; lower cortex light brownish-red, 6½–8μ thick, of 2 or 3 layers of cells about 4μ square, usually with a few protruding hyphae 5–25μ high. Apothecia subrotund to reniform, up to 3 × 5 mm with thin dentate margin, hymenium ca. thick including the pale fuscous to brown epithecium, hypothecium 20–35μ, hyaline to pale brownish, cortex above apothecia obscurely wrinkled, 30–50μ thick of nearly cubic cells 7½μ long, with some protruding hyphae; asci 45–55 × 8–13μ, somewhat clavate, (6-) 8-spored, spores nearly hyaline to pale reddishbrown 3-septate, 15–20 × 5½–7½μ paraphyses conglutinate, 2μ thick aseptate, slightly thickened at tip Pycnidia not seen.

Habitat Trees. damp rocks in shade.

Distribution.? Europe, New Zealand, Australia North Island: Te Kotukutuku (Colenso 5097) WELT Nelson: Aniseed Valley, Mr. 4171, Mr. 4172, (Knight) WELT (sub N. sublaevigatum) Canterbury: Tekapo, Scott 160, Scott 165; Mt. Misery, Philipson 31. Otago Lake Ohau, Mason 192;, Routeburn Valley, 0830; Trotter's Gorge, 3820, 3875; Akatore, 1501, Mr. 574; Kaka Point, 0157, 0439 Southland. Doubtful Sound, 3952 Australia Victoria, Seatoun Creek (F Camphell 51 in Knight coll) WELT sub N. sublaevigatum

The variety differs from var. helveticum in the near absence of tomentum and the rather smaller spores. There is also, at least in the specimens so far examined, a difference in the number of spores in the asci. It appears close to European forms under the names helveticum, subtomentellum, laevigatum, etc, but European.

specimens I have seen lack the characteristic isidia and are rather thicker and bluer than our plants. N. laevigatum appears to be a nomen ambiguum according to Gyelnik (1935), and the correct assignment of plants so identified is uncertain.N. helveticum var. rufum seems close to N. sublaevigatum Nyl. from Mexico, but I have seen no certain specimens of this. It is said to have a slightly reticulatecostate upper surface, a condition sometimes seen in specimens of var. rufum.

Nephroma lyallii Bab.

• Nephroma lyallu Bab. apud Hook., Fl. N. Z., II, 272 (1855), and plate 127a.

• Nyl., Synops. Lich I, 322 (1860).

• Hook Handb N. Z. Fl. 566 (1867).

• Nyl., Lich N. Z. 42 (1888)

• Hellbom, Bihang Kgl. Svensk Akad. Handl 21, III (13), 27 (1896).

Nephroma javanicum Gyelnik, Annal Crypt exot 4, 135 (1931), et apud Zahlbr Lich N. Z. 45 (1941)

Thallus blue or brown or variegated and smooth above, with lacerate to prohiferate margins centrally, brownish and tomentose below centrally, pale and glabrous towards the periphery, 3–8 cm dia; with lobes 3–5 mm wide by (5-) 15 (-25) mm long. Apothecia very pale brown, 5–10 × 3–5 mm, more or less reniform with 1 mm wide entire thalline margin. Hymenium 50–55μ high, hyaline with thin, faintly coloured epithecium; hypothecium 15–25μ thick, nearly hyaline; medulla above hypothecium 60–70μ thick, of loosely woven thin-walled hyphae 2½μ dia; algal layer here 8–15μ thick with Nostoc cells 4–7μ dia; pure blue or blue-green, cortex above apothecia 20–25μ thick of palisade-like cells ca 9 × 6μ. Paraphyses aseptate, 1μ thick, hardly thickened at the tips, asci 45–55 × 6–8 (-12)μ, 6–8 spored, thin-walled, cylmdrical, with spores in two series; spores nearly hyaline to pale reddishbrown, 1-septate, 16–18 × 4½) 5–6½μ sometimes slightly constricted at the septum.

Habitat. On small branches, usually in damp, shady places.

Distribution New Zealand and Java. North Island: Tiritea (G3, Zotov and Allan) CHR; Palmerston North (Zotov) CHR (all under N. javanicum); Hawke's Bay (Colenso) WELT. Westland: Franz Josef (C. K. Boey) 4165. Otago: Green Island, T 1956 Southland: Akatore, Mr. 576 (pr p); Forest Hill, 1011; Waihopai, Mr. 1305. Stewart Island: Oban, Mr. 708.

N. lyallii differs from most other Nephroma species in the uniseptate spores and the commonly blue algae. Gyelnik's brief description of N. javanicum makes no mention of the apothecial characters and is misleading in its reference to “superne isidiatus” since he apparently means merely that the plant has lacerate margins. Gyelnik's description of N. javanicum (1931) is not clearly different from his redescription of the type specimen of N. lyallu (1938).

The Southland and Stewart Island specimens seem a little stouter and with less fragile margins than the North Island specimens identified as N. javanicum by Gyelnik, and have 8-spored asci, whereas (Zotov, Palmerston North) CHR has mostly 6-spored asci. Too few collections have yet been made to decide whether these differences are significant.

The identificaction of Allison 269 as N. lyallu in Zahlbruckner (1941) seems to be an error, since the two portions of the collection retained in New Zealand are clearly N. australe.

Nephroma lyallii Bab. f. isidiatum Murray, f. nov.

A typo differt isidiis squamiformibus ad margines et rimas thalli. Ceteres ut in specie

Distribution. Westland: Runanga, Mr. 6920 Otago: Leith Valley, 3544; Mihiwaka, T 630 Southland: Forest Hill, 0660 (Type); Wilmot Pass, 3935. No locality (Knight) WELT (sub N. schizocarpum).

This form differs from the species in having subsquamulose isidia along the margins and cracks in the thallus. The isidia are mostly about 01 mm thick and 03 mm dia; and do not seem to be associated with damage by insects, etc Some specimens grade into the “typical” form of the species.