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Genus Sphaerophorus Pers.

Thallus fruticose, terete or flattened, or erect foliose, mostly entirely corticate, cortex usually of thick-walled vertical hyphae; algae green (one species with cephalodia containing blue-green algae); medulla of longitudinally arranged close-packed hyphae without distinct strengthening layer. Apothecia terminal or nearly so, with thalline margin (sometimes dehiscent); paraphyses slender, sparse; asci clavate to cylindrical, 8-spored; spores forming a well-developed mazedium, simple, globose, hyaline or brown, with or without encrusting black granules. Pycnidia brown, globose, with simple oblong hyaline pycnidiospores.

The genus has probably less than a dozen species, mostly in the Southern Hemisphere, although some are poorly defined or of uncertain status. Most of the species are rather variable, but undoubtedly the most protean is the Sph. melanocarpus-Sph. australis complex. Although the extreme forms in this group are very different in appearance, there seems to be a nearly complete series of intermediates, so that I have thought it most convenient to place all the New Zealand variants as varieties and forms of Sph. melanocarpus pending examination of specimens from other parts of the world.

Sphaerophorus melanocarpus (sensu latiore)

This species appears to be of nearly world-wide distribution, but with the greatest development of forms in Australasia, where it is also particularly abundant. Despite the very different appearance of the various varieties and forms, the morphological and chemical differences between them are hardly significant. A moderate proportion of the specimens examined are K + faint yellow, but this does not seem to be correlated with any other feature, and since the reaction is often doubtful I have not taken it into account in separating the subspecific taxa. It is largely a matter of convenience how many varieties and forms should be recognised, and it is unlikely that any two lichenologists would agree completely about their number and exact status, so the present scheme cannot be regarded as final. It might be equally satisfactory to treat the three varieties as separate species, particularly since they have different geographical ranges. Some of the forms are probably ecotypes, and this could probably be confirmed by experiment, but others must be more constant since two forms can sometimes be found growing together but easily separable into different populations. The most distinct taxa seem to be var. australis f subteres and var. scrobiculatus. The former sometimes looks like Sph. fragilis but flattened stems can usually be found near the periphery of the clump. The average spore diameter for var. scrobiculatus is a little greater than for the others, although the ranges overlap almost completely.

A curious feature of several specimens is the presence of fruiting structures additional to apothecia and the normal pycnidia; these are not visible macroscopically and sections showing them have been obtained only by chance. They are apparently randomly distributed over the upper surface and consist of irregular spaces in the algal layer and lower part of the cortex, without definite wall but lined with hyphae producing hyaline ellipsoid spores I have not clearly seen either the opening of these structures to the outside or the mode of attachment of the spores.

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Key to New Zealand Sphaerophorus Species
1. Thallus more or less compressed, dorsiventral 2
Thallus terete 3
2. Apothecia subterminal on ventral surface, over 1 mm dia., with thalline margin melanocarpus
Apothecia terminal on lobules, up to 1 mm dia., thalline margin absent in mature fruits cuneatus
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3. Thallus stout, with bluish cephalodia stereocauloides
Thallus not over 2 mm dia., without cephalodia 4
4. Apothecia in irregularly dehiscent globose receptacles globosus
Apothecia hemispherical, thalline margin lost in mature fruits tener
Sphaerophorus melanocarpus varieties and forms
1 Main stems and branches up to 1.5 times as broad as thick, branchlets mostly subterete var. melanocarpus 2
Main stems at least twice as broad as thick, branchlets compressed or subterete 3
2. Stems to 2 mm dia., with small sympodial branches or several times di- or trichotomously divided normal form
Sympodial axis to 1 mm dia., almost terete, branches not much narrower than axis and repeatedly di- or trichotomously divided in more than one plane f. ramosissimus
3. Fruiting stems up to 3.5 mm broad, more or less branched or dissected, backs of apothecia smooth or nearly so, apothecial margin narrow var. australis 4
Stems more than 3 mm wide, subentire to marginally laciniate, backs of apothecia strongly rugose-scrobiculate, margins usually broad var. scrobiculatus 7
4. Thallus several times dichotomously divided into branches 2–3 mm wide, without main stem f. palmatus
Thallus with evident main stem and final branches less than 2 mm wide 5
5. Margin of fruiting fronds laciniate f. proliferus
Margins entire or nearly so 6
6. Upper branches closely clustered, subterete f subteres
Small, repeatedly branched with final branches 0.05–0.3 mm wide f delicatus
More or less sympodially branched, final branches 0.2–0.8 mm wide f. angustior
Repeatedly sympodially and di- or trichotomously divided, branches widely spreading, final branchlets mostly 1 mm wide f vividulus
Branches linear, sparingly subdivided, 5–15 × 0 5–1 2 mm, main stems rather wide f insignis
Stems usually strongly flattened, with relatively few small branches “normal” form
7. Fruiting stems 3–8 mm wide normal form Fruiting stems 8–25 mm wide, thin f. macrophyllus
Sphaerophorus tener forms
Branching mainly dichotomous except for fruiting stems; plant usually on bark normal form
Sympodial axis clearly evident; plant usually on soil among debris f globosoides

Sphaerophorus cuneatus (Stirt.) Murray, comb. nov. Pl. 13, Fig. 1.

Calycidium cuneatum Stirton, Proc. Philos. Soc. Glasgow 10, 292 (1877).

Coniophyllum colensoi Müll Arg, Bull. Soc. Bot. Belg., 31, 23 (1892)

(?) Sphaerophoron polycarpum Colenso, Trans. N. Z. Inst., 16, 361, 1883 (1884).

Thallus of erect fronds 5–10 mm long by 3–6 mm wide, entire, K-; upper surface smooth, brownish (originally grey-green) lower surface pale brownish (originally white) smooth to somewhat longitudinally ribbed and wrinkled; upper cortex 50μ thick, hyaline, of vertical septate hyphae sometimes appearing almost pseudoparenchymatous with cells up to 20μ long; algal layer 25–40μ thick, with light green thinly halonate gonidia 7–13μ dia.; medulla at least 150μ thick, of moderately loosely woven anastomosing hyphae about 2½μ dia.; lower cortex discontinuous, brownish, up to 25μ thick, similar to upper cortex. Apothecia on marginal lobules about 1 mm long, 1 to 15 per frond, mostly slightly displaced to ventral surface, hemispherical, 0.5 mm dia., with thin thalline margin at base (like mature Sph. tener fruits), K + brownish, hymenium up to 100μ high, more or less hyaline except for brown epithecium and mazedium; hypothecium dark brown, mostly 200–250μ high; faintly brownish proper margin 12–15μ thick continuous beneath hypothecium; paraphyses very thin, rather numerous, entangled, apparently simple, asci cylindrical, 8-spored; spores monostichous,

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globose, brown, without pigment granules (4-) 5 (-5-½)μ dia., thin walled, forming a well-developed mazedium. Pycnidia not seen. (Description from isotype specimens.)

Habitat. On bark.

Distribution. Chatham Islands. Southland: Secretary Island, 3980, 3997. Chatham Island: (Colenso 7), (Colenso 30), WELT (apparently isotype specimens from a single collection).

This species was till very recently known only from the type specimens acquired by Colenso (probably collected by Travers) about 90 years ago. The Secretary Island plants are rather larger (to 15 × 8 mm) with less entire margins than the isotypes, have mostly longer fertile lobules with dark brown fruits, a more continuous lower cortex and a less cellular upper cortex 65μ thick, but are clearly conspecific. The structure of sterile fronds is very similar to that of Sph.. melanocarpus var. scrobiculatus and indeed the plant would be taken for this in the absence of fruits. There is no justification, then, for retaining the genus Calycidium and I have reduced it accordingly. The apothecia are similar to, and develop in the same manner as, those of Sphaerophorus tener. The present discontinuous distribution suggests that the species will be found elsewhere in New Zealand, although it is almost certainly not present on the East side of the Southern Alps.

Sphaerophorus globosus (Huds.) Vainio.

Lichen globosus Huds., Fl. Angl. 460 (1762).

Sphaerophorus globosus Vainio, Lich. Ant. 35 (1903).

Sphaerophorus coralloides Pers., Usteri N. Ann. Bot. 1st, 23 (1794).

? Hook., Handb. N. Z. Fl., 559 (1867)

? Hook., Fl. N. Z., 304 (1855).

? Buchanan, Trans. N. Z. Inst., 6, 231 (1873) (1874).

Thallus up to 10 cm high, forming loose clumps, primary axes to 2 mm dia., sparingly branched, smooth or slightly impressed, terete or sometimes slightly compressed, with short terete phyllocladial branches 0.2–0.3 mm thick, whitish or greenish-white or brownish, cortex 90–110μ thick, hyaline, of conglutinate more or less vertical branched hyphae, not clearly delimited from medulla., algae Cystococcus; medulla of longitudinal 4–5μ dia. hyphae, I + blue. Apothecia rather rare, in tips of thicker branches, globose, irregularly dehiscent above; hypothecium globular-columnar, brownish above, hymenium nearly hyaline, paraphyses few; thin, simple, asci cylindrical, spores globose or subglobose, becoming bluish then black, 7–11μ dia. with papillate wall. Pycnidia immersed in tips of phyllocladia. (Description from Vainio. 1927).

Habitat. On mossy rocks, soil, etc.

Distribution. Northern Hemisphere, South America and adjacent Islands,? Australia and? New Zealand.

Exsiccata seen: Fl Suecica 19 (Henriksson).

Although this species has been reported several times from New Zealand and the Subantarctic Islands, I have seen no undoubted specimens. Possibly the plant referred to is Sphaerophorus tener f. globosoides, a form which is hardly distinguishable from Sph. globosus when sterile, as it usually is. Hooker records the species as being collected by Colenso, but there are no specimens of it in the portion now in the Dominion Museum. A short, caespitose form of the species in Europe is known under the name Sph. globosus f. curtus (Hook.) Zahlbr. based on Sphaerophoron curtum Hook. from Campbell Island. This is evidently a misidentification, as the Hooker species is conspecific with Sph. tener. Although the records of Sph. globosus for New Zealand are probably incorrect, I have included a description of it as the species may well be present. Two specimens identified as “Sph. coraloides” in the Knight collection are a small form of Sph. melanocarpus and a very isidiose specimen of Menegazzia nothofagi Zahlbr.

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Sphaerophorus melanocarpus D. C. var. melanocarpus.

Sphaerophoron melanocarpon D. C., Fl. Fr. 6, 178 (1805).

Sphaerophoron compressum (pr. p.) Hook., Fl. Antarctica, 1, 196 (1844).

Fl. N. Z., 305 (1855).

Handb. N. Z. Fl., 559 (1867).

Lindsay, Trans. Linn. Soc., 25, 530 (1866).

Kirk, Trans. N. Z. Inst., 4, 235 (1871).

Buchanan, Trans. N. Z. Inst., 6, 231 (1873).

Nyl, Compt rend Paris, 83, 87 (1876)

Lich., N. Z., 13 (1888).

Müll Arg., J. Linn. Soc. Bot., 32, 198 (1896).

Hellb, Bihang Kgl. Svensk Vetensk. Handl., 21, III (13), 129 (1896).

(?) Sphaerophoron compressum var. candidum Müll Arg, J. Linn. Soc. Bot., 32, 198 (1896).

Thallus fruticose, more or less erect, sometimes forming clumps up to 15 cm across, 2–10 cm high, either with evident sympodial axis or several times di- or trichotomously branched; main axis up to 2 mm broad and 1.2 mm thick, sometimes impressed or foveolate on ventral surface and with cortex up to 220μ thick but usually about 90μ, of vertical conglutinate thick-walled hyphae 12–20μ dia., thinner on lower surfaces and on branches; algal layer 40–65μ thick but not clearly distinct and thinner and less continuous on lower surface, algae green, 5–10μ dia.; medulla of longitudinal thick-walled hyphae up to 7μ dia., often encrusted with colourless crystals, K- or f. yellow, P-, I- Phyllocladial branches down to 50μ dia. with 10μ thick cortex but usually considerably larger, terete or flattened. Apothecia subterminal with narrow margin or on ventral surface near tip of main stem and larger branches; irregularly lentiform and finally hemispherical, 1–3 mm dia., hypothecium hyaline, hemispherical, not clearly distinct from medulla; asci cylindrical, paraphyses few, simple, slender, spores globose, 6–7½ (−12) μ, smoky-grey and encrusted with black pigment granules which ae weakly K + purplish. Pycnidia black, globose, 100–300μ dia., pycnidiospores hyaline. ellipsoid-oblong, 3 × 1μ.

Habitat. On soil and debris, rarely on trees.

Distribution. More or less cosmopolitan. North Island: Herekino (Poole) CHR; Ruahines (Colenso) WELT: Marlborough. Mt. Stokes, CHR: Canterbury. Poulter Valley, Sc 70, Sc 71. Otago. Haast Pass, 3902; Mt. Watkins, T 2064 Southland: Doubtful Sound, T 2839, T 2840; Lake Hauroko, Mr. 7304. Campbell Island: (Oliver) WELT 18.

Exsiccata seen: Arnold 873 (WELT), Gyelnik's Lich. (CHR).

Sphaerophorus melanocarpus var. melanocarpus f. ramosissimus Murray f. nov. Axis ramesque subteretes, repetito ramosi ramulis teretibus.

Thallus erect, greenish, usually forming a loose hemispherical clump to 10 cm high, main axis and branches about 1 mm dia., terete or nearly so, repeatedly branched, sometimes in more than one plane, final branches almost terete with continuous, even algal layer. Fruiting stems rare, distinctly flattened, apothecia as for the variety but somewhat scrobiculate on backs. Spores 10 (-13) μ dia.

Habitat. Usually on branches.

Distribution. North Island: Little Barrier Island (Hamilton) CHR; Hawke's Bay (Colenso) WELT, Ramahanga R. (Colenso 2171), Tarawera, (Colenso 3843), Ruahines summit (Colenso 2711, sub “Sph. album” W. C. nom. nud.) WELT. Southland: Secretary Island, 2,800ft, 4052, 4053, 4054 (Type).

This seems a distinct form with something of the appearance of a stout Sph. tener, but the fruit clearly demonstrates its affinity to Sph. melanocarpus. The apparently discontinuous distribution is probably due to insufficient collecting, although it does seem to be rare.

Sphaerophorus melanocarpus var. australis (Laur.) Murray comb. nov. Pl. 13, Fig. 2.

Sphaerophoron compressum (pr. p.) Hook., Handb. N. Z. Fl. 559, 1867, et. auct. al. Sphaerophoron australe Laurrer, Linnaea II, 44 (1827).

Hook, Fl. Antarctica 1, 195 (1844).

Fl. N. Z., 304 (1855).

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Nyl., Synops. Lich., I, 170 (1860).

Compt. Rend. Paris 83, (1876).

Krmph., Reise der Novara, Bot. I., 127 (1870).

Chilton, Subant. Islands N. Z., 2, 530 (1909).

Cockayne, Trans. N. Z. Inst., 42, 320 (1909).

Müll Arg., J. Linn. Soc. Bot., 32, 198 (1896).

Hellb., Bihang Kgl. Svensk Vetensk. Akad. Handl., 21, III (13), 129 (1896).

Sphaerophoron compressum var. australe, Linds., Trans. Linn. Soc., 25, 530 (1866).

Thallus fruticose (often growing horizontally from tree trunks), strongly compressed, (5-) 30–50 (-100) mm high, main axis ½–3 mm wide, 0.2–1 mm thick, at least in part more than 1.5 times as broad as thick, sparingly to considerably branched in one plane, branchlets more or less flattened (except in f. subteres). Cortex of main stems hyaline, 50–150μ thick, of conglutinate very thick-walled hyphae, similar on ventral surface but thinner; algal layer continuous above, 15–35μ thick, mostly absent below, algae green or sometimes yellowish-green, mostly 5–10μ dia., medulla of densely packed more or less longitudinally arranged thick-walled hyaline hyphae; K- or f. y, P-, I-. Tips of fertile branches expanded, apothecia on ventral surface, subterminal or with narrow margin, initially usually lentiform becoming round when mature, smooth to weakly scrobiculate on back, 1–4 mm dia, traces of brown excipulum sometimes present, hypothecium hemispherical or flattened, to 0.8 mm thick, brown, in saucer-shaped depression, hymenium hyaline in lower parts, paraphyses very few, sparingly branched, to 1μ thick with small swellings, asci 40 × 5μ cylindrical to irregularly clavate, spores mostly in one series, globose, 8–10½ (-13)μ, faintly brownish with encrusting black granules (K + purplish or brownish), forming thick mazedium. Pycnidia black, globose, 100–300μ dia., partly immersed in tips of final branches and less commonly on ventral surface, pycniospores hyaline, oblong, 3½ × 1μ. In some specimens ellipsoid spores, 5 × 2½μ are present in hyaline irregular (?) pycnidia.

Habitat. On bark, less commonly on soil or mosses.

Distribution. Australia, New Zealand, South America, Philippines, Hawaii, Pacific Islands. North Island: Ruamahanga R. (Colenso 2617, 2621 and 2714), Te Hawera (Colenso 2718), Toruarau (Colenso 4721), Te Kotukutuku (Colenso 5094), Ruahines (Colenso 1509) WELT; Taupo, Allison 264; Westland: Rununga, Mr. 6879; Toaroha River, 3,200ft, Sc 150; Greymouth, Mr. 7095. Otago: Haast Pass (Smith) 0953; Mt. Cargill, T 556; Mihiwaka, T 659; Dunedin, 3548; Kaka Point, 0390. Southland; Purakanui Falls, 0648, Wilmot Pass, 3930, 3932 Doubtful Sound, T 2841; Secretary Island, 4055. Stewart Island: T 3009; T 3013; T 3014; T 3015; T 3016. Chatham Island: (Colenso 23) WELT. (Colenso 5) WELT. Campbell Island: (Oliver) WELT. 29.

The specimens left under the variety name only are not very uniform; apart from immature material they include mostly smaller specimens with a few irregular branches. It is doubtful whether many of the forms have real taxonomic significance, but it is convenient to use them with a species so variable as Sph. melanocarpus. Several attempts were made to arrange the specimens in natural groups, and the one given here seems the most successful, although the scheme would doubtless require modification to accommodate non-New Zealand specimens satisfactorily. It is interesting that some of the forms appear to be restricted to certain parts of the country, often to one side or the other of the 60in isohyet.

The apothecia develop initially beneath the lower cortex, and break through it as they mature, the cortical covering splitting and eventually disappearing. The black pigment granules which coat the mature spores are present in a thick layer in the immature fruits before any spores are discharged from the asci; the spores while still in the asci are smooth and hyaline, and when mature are only very faintly coloured.

The entities to which Colenso applied the names vividulum and polycarpum must remain in some doubt, since a careful search of his specimens in the Dominion Museum failed to disclose any bearing these names, although three carried other apparently unpublished names. It is possible that Sph. polycarpum Col. is actually Sph. cuneatus but this cannot be determined in the absence of the type specimen.

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Sphaerophorus melanocarpus var. australis f. angustior (Reinke) Murray comb. nov. Pl 13, Fig. 3.

Sphaerophoron australe f angustior Reinke, Pringsheim's Jahrb. f. wiss. Bot., 28, 85 (1895) and Figs. 20, 21.

Sphaerophorus australis var. angustior Zahlbr., Lich. nov. Zel., 11 (1941). (reprint) Plant to 8 cm high, main axis 1–2 mm wide to ½ mm thick, more or less sympodially and repeatedly branched with final branches 0.2–0.8 mm wide; apothecia usually nearly terminal and rather small, 1–2 mm dia.

Habitat. On bark.

Distribution. Australia and New Zealand. North Island: Makakahi R. (Colenso 2713, sub. Sph. umbilicatum W. C. nom. herb), Cape Kidnapper (Colenso 3570) WELT; Puketitiri (M Clark) 4500. Otago. Huxley River 1835; Mt. Cargill T 524, T 568; Silver Peaks T 1185, T 1197; Saddle Hill T 184; Routeburn Valley 0787, Dunedin 3547 Southland. Milford Track T 2887, T 2915; Tautuku T 1163; Wilmot Pass 3937; Doubtful Sound T 2822; Secretary Island: 3978, 3995. Stewart Island Freshwater River T 3065.

Sphaerophorus melanocarpus var. australis f. delicatus Murray, f. nov. Pl. 13, Fig. 4.

Thallus parvus, delicatus, flabelliformis, ramibus ultimis parvissimis; apothecia parva. Plant 1–3 cm high, fan-shaped, with stems up to 2 mm wide at the base, but mostly 0.8 mm, repeatedly sympodially branched with final branchlets 0.05–0.2 mm wide, distinctly flattened, cortex 20μ thick. Fruiting stems 1 mm wide, apothecia 1 mm dia. with very narrow margin; spores hyaline, 8μ dia. heavily encrusted with black pigment granules.

Habitat. Probably on wood.

Distribution. North Island. Puketitiri (M. Clark) 4295 (Type). Otago: Matukituki Valley, 3,000ft (R. F. Smith) 0958; Saddle Hill T 178.

Although this form is unusually delicate for the species and only the broadening at the very base of a few fronds in the type specimen places it in var. australis, it is approached by a few specimens of f. angustior.

Sphaerophorus melanocarpus var. australis f. insignis (Laur.) Murray. comb. nov. Pl 13, Fig. 5

Sphaerophoron insigne Laur, Linnaea, II, 45 (1827).

Sphaerophorus australis f. insignis Müll. Arg. Flora. 66, 17 (1883).

Sphaerophorus australis var. insignis Zahlbr, Lich Nov Zel, 11 (1941) (reprint) Main stems short, up to 3 mm wide, sparingly branched, branches more or less linear, 5–15 mm long by 0.5–1.2 mm wide, apothecia as in the variety.

Habitat. Mostly on soil among mosses, etc.

Distribution South America, New Zealand. Nelson: Lead Hills T 1975. Canterbury Lewis Pass, Mr., 6870. Otago: Routeburn Valley, 0815. Southland: Secretary Island, 3994; Lake Hauroko, Mr. 7303. Stewart Island. Freshwater River, T 3034.

One of these specimens, Mr. 7303, has hyaline irregular? pycnidia scattered beneath the upper surface, with ellipsoid spores 5 × 2½μ

Sphaerophorus melanocarpus var. australis f. palmatus Murray. f. nov. Pl 13, Fig. 6

Thallus ad 2 cm latitudine et altitudine bis terve dichotome divisus, laciniis (1-) 2–3 mm latis.

Thallus 2 cm high, several times dichotomously divided into branches of nearly uniform width, 2–3 mm wide and 0.3–0.5 mm thick, ends rounded or with terminal laciniae 1 mm long and 1 mm wide, lower surface weakly impressed, white pruinose near the ends of branches; upper cortex hyaline of thick-walled vertical highly gelified hyphae, 75μ thick, lower cortex 30–70μ thick, of more or less vertical thick-walled hyphae appearing distinct in KOH (resembling lower cortex of Sph. cuneatus); algal layer 25–30μ thick, of numerous closely packed cells 5–8μ dia., medulla looser than usual for the species, hyphae 5–8μ dia., not regularly longitudinally arranged, K-, P-, I- The pruinose appearance of most of the lacinae is due to irregular proliferations of the lower cortex and some medullary hyphae protruding. Pycnidia immersed, brown, globose with wide ostiole, 300μ dia., pycnidiospores oblong 3 × 1μ.

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Habitat. On bark.

Distribution. Otago: Dunedin, T 2132 (a doubtful juvenile specimen). Southland: Doubtful Sound (Thomson & Simpson), CHR (Type).

Although there is only one good collection of several individual plants, they are readily separable from any other specimens of the variety I have seen; they resemble in shape f. vividulus, which is, however, considerably narrower and more branched. There are no apothecia on the specimens.

Sphaerophorus melanocarpus var. australis f. proliferus (Wilson) Murray, comb. nov.

Sphaerophoron australe var. proliferus Wilson, J. Linn. Soc. Bot., 28, 370 (1891). A small form of var. australis in which the margin of the apothecium is fimbriate with branches linear and several mm long, and rarely carrying small apothecia at their tips.

Habitat. On bark.

Distribution. Tasmania North Island: Orongorongo River (Healy) CHR. Otago. Mt. Cargill, T 540.

Specimens of var. australis rarely have apothecial margins with one or two short processes, but these carry a regular corona of 4 to 10 branchlets.

Sphaerophorus melanocarpus var. australis f. subteres (Zahlbr.) Murray, comb. nov. Pl. 14, Fig. 7.

Sphaerophorus australis f. subberes Zahlbr. apud Magn., Arkh. f. Bot., 31 A, (1), 24 (1944).

? Sphaerophoron fragilis, Chilton, “Subant. Ids. N. Z.”, 2, 530 (1909).

Forming dense mats or clumps, 5–25 mm high, main stems short, commonly subterete, with short vertical closely-clustered subterete often simple branches 0.5–0.8 mm dia. flattened stems and branchlets commonly only present near periphery of clumps; cortex nearly even round main stems and 100μ thick, algal layer thin and discontinuous; cortex and algal layer of uniform thickness round upper branchlets. Apothecia rare, appearing terminal in globose receptacles like those of Sph. globosus and opening irregularly circular. Structure of apothecia and pycnidia as for the variety. Thallus K-, P-, I-.

Habitat. On soil in rock crevices in subalpine situations.

Distribution. Australia, South America, Hawaii North Island: Napier (Colenso, 1510) WELT. Canterbury. Arthur's Pass (D. Billings, NZL 46) 4499. Otago: Trotter's Gorge, T 1934, 3849; Mt. Watkin, T 1766, T 2010; Silver Peaks, T 248, T 249, T 250, T 280, T 232, T 1196, 4285, 4286; Maungatua, 2,000ft, 1170, T 2600, T 370, Akatore Gorge, 3724. Campbell Island: Mt. Beeman (1958 party), 3652.

This form has much the appearance of Sph. fragilis, from which it is distinguished by fruit characters and chemical reactions, or a stunted Sph. melanocarpus var. melanocarpus from which the few clearly flattened stems present in most specimens separate it. Sections of an immature apothecium show it to develop slightly to one side of a thickened stem, and to resemble that of melanocarpus rather than fragilis or globosus.

Sphaerophorus melanocarpus var. australis f. vividulus (Colenso) Murray, comb. nov. Pl. 14. Fig. 8.

Sphaerophoron vividulum Colenso. Trans. N. Z. Inst. 17, 263 (1884 iss. 1885).

Thallus to 10 cm high, main axis not strongly marked above, branching regularly sympodial then di- and trichotomous, forming fan-shaped branches, branchlets mostly 1 mm wide and 0.5 mm thick, smooth or impressed beneath; apothecia usually small and obliquely inserted in tips of slightly expanded branchlets.

Habitat. Usually on moist soil among bryophytes.

Distribution. New Zealand. North Island. (Colenso) WELT. (Attwood V2) CHR; Otara (Colenso 2712, sub. Sph. lacunosum W. C. nom. nud.) WELT (doubtful), Tararuas, (Zotov) CHR. Otago: Mt. Cargill, T 528, T 530. Southland: Hokanui, 0390; Doubtful Sound (Simpson & Thomson) CHR; Secretary Island, 3996.

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Colenso's type specimen was collected at Norsewood, but no collection from there in the Dominion Museum material matches his description which, however, fairly clearly indicates this form. The specimen Colenso 2712, from the Rangitikei River, has prominent apothecia with rugose receptacles, and may be a distinct form.

Sphaerophorus melanocarpus var. scrobiculatus (Bab.) Murray, comb. nov. Pl. 14, Fig. 9. Sphaerophoron australe var. scrobiculatum Bab. apud Hook. Fl. Nov. Zel., II, 304 and Tab. CXXX, c (1855).

Zahlbr., Lich. nov. Zel., II (1941) (reprint).

Thallus of erect, sometimes imbricating foliose fronds (0.2-) 0.4–0.8 mm thick and 3–8 mm wide by 1–3 cm (sterile) or 2–9 cm (fruiting) long, subentre or once or twice divided or divided at margin into laciniae about 1 mm wide and up to 10 mm long; cortex as for var. australis but mostly 60–70μ thick above and 40–50μ thick below (sometimes much thinner); medullary hyphae mostly near 5½μ dia. Apothecia on narrower lobules, large, 3–8 mm dia., round or broadly lentiform, margin very variable in width, (1-) 2–3 (-8) mm wide, backs of apothecia deeply rugose-scrobiculate; spores globose, 8–11 (- 14)μ dia., nearly hyaline with black pigment granules which are K + dark purplish. Pyenidia as in var. australis.

Habitat. On tree-trunks and branches among bryophytes.

Distribution. New Zealand. North Island: Pirongia CHR; Tararuas, 3,000ft (Zotov) CHR. Marlborough; Pelorus Bridge, Mr. 1221. Westland: Tokaroha River, Sc 146. Styx River, Sc 135. Otago: Mt. Cargill, T 537: Southland. Doubtful Sound, T 2819–21, T 2880; Wilmot Pass, 3931; Manapouri, 3918; Secretary Island, 3991; Orepuke (Sorensen) CHR Stewait Island: Freshwater River, T 3064.

Sphaerophorus melanocarpus var. scrobiculatus f. macrophyllus (Zahlbr.) Murray, comb. nov. Pl. 14, Fig. 10 Sphaerophorus australis var. macrophyllus Zahlbr., Lich. Nov. Zel., 11 (1941) (reprint).

Thallus as for the variety but with fronds (particularly fruiting ones) up to 25 mm broad and 0.25–0.45 mm thick, mostly entire or coarsely crenate. The margin of the apothecia may be very broad and with secondary apothecia on proliferations or only 1 mm wide, even in the same specimens. Frequently there are narrow fronds like those of var. australis f. insignis mixed with the broad ones. Structural details of thallus and apothecia are as for the variety.

Habitat. As for the variety.

Distribution. New Zealand. North Island: Tararuas, Mt. Dora (Chamberlain) CHR (? isotype), Mt. Dennan (Zotov) CHR; Tiritea (Allan) CHR Canterbury: Cass (Allan) CHR. Southland. Secretary Island, 3979, 3990, 3993, 4051. Stewart Island: Table Hill, T 3005, T 3122.

The specimens listed under this form are not particularly homogenous, and nonfruiting specimens may be difficult to separate from the typical form of the variety. However, field studies in the Doubtful Sound area where most taxa in Sphaerophorus are very common indicate that at least 80% of specimens can be assigned to one form or the other without difficult.

Sphaerophorus stereocauloides Nyl.

Sphaerophoron stereocauloides Nyl., Flora. 69 (1869).

Lich. Nov. Zel., 12 (1888). Hellb., Bihang Kgl. Svensk Vetensk. Aka. Handl. 21 III (13), 129 (1896).

Colenso, Trans. N.Z. Inst., 17, 264 (1884).

(?) Chilton, “Subant. Ids. of N.Z.” 2, 530 (1909).

Sphaerophorus nobilis Zahlbr., Lich. Nov. Zel., 10 (1941).

Thysanophoron Pinkertonii Stirt., Trans. Bot. Soc. Edinb., 14, 359 (1882).

Thallus white or grey (greenish-grey when fresh), 7–15 cm high, fastened to substratum with brown branching rhizoids, sometimes polytomously oranched at base and usually several times dichotomously divided with many clustered phyillocladial branchlets. Main stems and branches terete, 2–3 mm dia., usually with annular cracks, cortex of vertical gelified thickwalled

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hyphae, algal layer discontinuous, medulla of longitudinal very thick-walled hyaline hyphae, phyllocladial branchlets mostly 1–5 mm × 0.2 mm dia., terete or compressed near tips, simple or branched with cortex 12–15μ thick, algal pale greenish, 5–8μ dia., medullary hyphae 5½-8½μ dia., cephalodia rare to frequent among phyllocladia, pale blue to greenish blue, mostly irregularly clavate, 500μ long by 150–400μ dia., cortex like that of phyllocladia, smooth, containing both discrete hyphae and structureless hyaline material with scattered coiled chains (up to at least 80μ long and 5–7μ wide) of Scytonema, cells bright greenish blue and about 3μ long. Apothecial terminal on branches, in globose receptacles 1½-3 mm dia., usually stellate dehiscent, hypothecium brown, columnar-globose, mazedium black, thick, spores finally dark brown, more or less globose, 9–12μ dia, without encrusting granules; pycnidia not seen.

Habitat. On trees and logs.

Distribution. New Zealand. North Island. Summit of Ruahines (Colenso, 2202 WELT. Canterbury: Lewis Pass, T 2430; Andrew River, Sc 47; Sc 51; Waimakariri River (Allan, L9) CHR, Arthur's Pass (Martin, 13) CHR; Cass (Philipson 35 and 77), Canterbury Univ. Bot. Dept. Otago. Howden, 0800, 0841. Southland: Stuart Range (W. A. Thomson, ZA 424) CHR (isotype of Sph. nobilis Zahlbr.); Lake Manapouri (Billings, NZL 8) Duke Univ. Bot. Dept.; Lake Monowai (D. Hamilton), 089. Stewart Island: (J. D. Smith) CHR.

This handsome species is the largest in the genus, and the only one to have cephalodia. In fresh material these cephalodia are almost indiistinguishable from phyllocladia, but are readily seen in herbarium material after a few years when the rest of the plant bleaches to white. Evidently Zahlbruckner had fresh material when he erected Sph. nobilis as the cephalodia are now clearly visible on the isotype specimens; these are also unusually large and stout for the species. In one of the above specimens the cephalodia seem to be absent. Like Sph. melanocarpus var. scrobiculatus, Sph. stereocauloides is apparently confined to parts of New Zealand with high rainfall—i.e., to the western side of the South Island and some parts of the North Island.

Sphaerophorus tener Laur.

Sphaerophoron tenerum Laur., Linnaea II, 45 (1827).

Hook., Fl. Antarct., I, 195 (1844).

II, 530 (1847).

Fl. N. Z., II, 304 (1855).

Handb. N. Z. Fl., 559 (1867).

Mont., Voy. Astrolobe Pôle sud, Bot. I, 170 (1845).

Nyl., Synops. Lich., I, 171 (1860).

J. Linn. Soc. Bot., 9, 244 (1865).

Lich. N. Z., 13 (1888).

Linds., Trans. Linn. Soc., 25, 530 (1863).

Krmph., Reise der Novara, Bot. I, 127 (1870).

Kirk, Trans. N. Z. Inst., 4, 235 (1871).

Buch., Trans. N. Z. Inst., 6, 231 (1873).

Nyl., Comptes rend. Paris, 83, 87 (1876).

Hellb., Bihang Kgl. Svensk Vetensk Akad. Handl., 21, III (13) 129 (1896).

Müller, J. Linn. Soc. Bot., 32, 198 (1896).

Chilton, “Subant Ids. N. Z.”, II, 530 (1909).

Szat., Borbasia, 1, 55 (1939).

Zahlbr., Lich. N. Z., 12 (1941).

Sphaerophoron australe Tayl. & Hook, London J. Bot., 3, 654 (1844).

Sphaerophoron tener var. stereocauloides Nyl., Synops. Lich., I, 171 (1860).

Sphaerophoron curtum Hook. & Tayl., London J. Bot. 3, 654 (1844).

Sphaerophoron tenerum var. curtum Tayl. & Hook, Fl. Antarct. I, 195 (1844).

Thallus terete, fruticose, white or pale greenish, smooth to somewhat shining, forming clumps (0.2-) 2–10.(-50) cm high by repeated dichotomous branching; lower stems mostly 1 mm dia., final branches 0.15 mm dia., K-, P-, I-. Cortex of primary stems hyaline, even, 25–30μ thick (down to 12μ in final branches), of gelified vertical thick-walled hyphae; algal layer usually discontinuous, 20–30μ thick, algae pale green, 8μ dia., medulla of closely packed

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longitudinally arranged very thick-walled hyphae (4-) 7 (-10)μ dia., with rough surface (clearing in KOH). Fruiting stems thicker and longer than sterile, sometimes scrobiculate below apothecia; apothecia more or less globose, without thalline or proper margin at maturity, 0.8–1.2 mm dia.; hypothecium dark brown, cushion-shaped to hemispherical, about 400μ wide, K-, composed of small entangled hyphae, hymenium hyaline below; asci cylindrical, 8-spored, spores smoky-grey with few or no pigment granules, globose, (6-) 7–8μ dia.

Habitat. On trees or moss cushions on soil.

Distribution. Australia, New Zealand, South America. North Island: Rangitoto (Allan L 7) CHR; Tongariro (Allan) CHR; Maungapohatu (Cranwell & Moore) CHR; Pirongia CHR; Raetihi (Attwood) CHR; Moehau (Moore) CHR; Tararuas (Allan, Chamberlain) CHR; Waikaremoana, Auckland Univ. Bot. Dept., Ruahines (Colenso, 2715, sub Sph. excelsum) WELT; Te Hawera (Colenso 2716, 2717 and 2.822) WELT; Ruamahanga R. (Colenso, 2124) WELT; Tararuas (Colenso, 2183) WELT; Mt. Holdsworth (Zotov) CHR. Nelson: Lead Hills, T 1978: Westland: Greymouth (Mackay). CHR; Styx River, Sc 148, Sc 149. Canterbury: Andrew River, Sc 74; Arthur's Pass (Allan, 21) CHR. Otago: Huxley River, 1854., Mt. Cargill, T 531, T 536, T 539, T 564; Mihiwaka, T 660, T 661, T 669, T 670; Silver Peaks, T 1191, 4287; Cave Hill, T 236; Leith Valley, T 2134; Flagstaff, T 81, 1167, Saddle Hill, T 154; Howden, 0835. Key Summit (R. E. Corbett) 3641. Maungatua, T 2898. Southland: Pahia Point, T 2251. Tautuku, T 1161; Doubtful Sound, T 2842; Secretary Island, 4056, 4057; Ben Bolt, 1074, Bluff Hill, T 831; Riverton, T 795; Orepuke (Sorensen) CHR. Stewart Island; T 3018, T 3109; Freshwater River, T 3063; Port Pegasus, 077, 0416. Auckland Island: Adam's Is. (Turbott & Easton) CHR. Campbell Island: (Bayley) 1630, (Rae) 3653.

Sphaerophorus tener is probably universally distributed in New Zealand but is commoner in the wetter areas, in the Fiordland subalpine beech forest it is the commonest epiphytic lichen. It varies greatly in size, the smallest fruiting plant I have seen being only 4 mm high, and the largest being a clump 120 × 50 cm and up to 40 cm thick; nevertheless the structure is very uniform throughout. The fruiting stems are extended and usually simple, but sometimes furcate or with several fertile branches. As in other species of the genus the apothecia develop under the cortex, which is later thrown off or remains as a thin flat rim at the base of the fruit.

Sphaerophoron curtum Hook. & Tayl. is a small, rather densely caespitose form not uncommon in exposed situations. Some specimens from Campbell Island form a turf a few mm high and 30 cm or more in diameter Pl. 14, Fig. 14.

Sphaerophorus tener f. globosoides Murray f. nov. Pl. 14, Fig. 14.

? Sphaerophoron coralloides Hook. Fl. 559 (1867).

A forma typicale differt axilibus sympodialibus, ramulis brevibus tenuibusque et KOH reactione normaliter flavescente.

Thallus caespitose, white or brownish, often shining, 1–6 cm high branching mostly sympodial from terete main stems (which may be dichotomously divided) 0.7–0.8 (-1.6) mm dia., branchlets mostly short and often clustered. (100-) 150–300 μ dia., sometimes breaking up into soredia at tips. Cortex completely gelified even in KOH. 15–30μ thick on main stems and 10–15μ on branches; algae in scattered or contiguous colonies about 50μ dia., beneath cortex, algal cells pale yellowish-green, Trebouxia, about 8μ dia.; medullary hyphae thick walled, rough (clearing in KOH), 4½-8μ dia., algal layer near tips of branchlets is usually K + yellow, P-, I-; apothecia as in typical form but very rare, spores pale greyish, 7–8μ dia. Pycnidia not seen.

Habitat. Subalpine grass and scrubland.

Distribution. North Island: Ruahines (Colenso 2730) WELT; Tongariro.

(Allan) CHR. Otago. Mt. Watkins, T 1570., Silver Peaks, T 1190, T 1192, 4288 (Type); Cave Hill, T 233; Maungatua, 3,000ft, T 1804. Key Summit, T 2927; Southland: Tautuku, T 1662; Secretary Island; 3,400ft, 4058. Stewart Island

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Table Hill, T 3010, T 3017, Mt. Anglem, 3,200ft, Mr., 30, Mr. 42. Campbell Island: (Bayley) 1631, (A. F. Rae) 4326, (Oliver) WELT 23 and 31.

Extreme examples of this form have a very distinct appearance, but they are connected with the typical form of the species by intermediates, and the taxon is probably an ecotype. It usually differs from the normal form of the species in the positive reaction with KOH, although this is absent in shaded specimens. It is hardly distinguished macroscopically from Sph. globosus, but has a thinner cortex and a different reaction with iodine. The Stewart and Campbell Island specimens have very stout main stems with relatively few phyllocladial branchlets, and may possibly prove to be separable as another form; they also have a more matt surface than usual for f. globosoides.

I am much indebted to Mr. W. Martin, Mr. D. Scott, Mr. T. A. Thomson, and the Directors of Botany Division, D.S.I.R. and the Dominion Museum for gifts and loans of specimens, and to several collectors whose names appear in the species distribution records above.