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their growth is dense other cryptogams are absent save for Sphagnum australe. Where the growth is short or more open, and especially on bare patches of peat the following cryptogams were collected:—Ferns: Schizaea fistulosa and Lindsaya linearis. Lycopods: Lycopodium ramulosum. Mosses: Polytrichum juniperinum, Campylopus torquatus, C. introflexus, Campylopodium euphorcladum and Dicranoloma billiardieri. Hepatics: Cephaloziella exiliflora, Chiloscyphus coalitus, Chiloscyphus normalis, Lophocolea heterophylloides, L. insularis, and on the sides of a large drain Marchantia berteroana and Riccardia sp. Fungi were not uncommon, all being agarics, but lichens were few. These were the three Cladoniae of the subgenus Clathrina—viz., Cladonia aggregata, C. retipora, and C. sullivani, together with C. cornutoradiata and C. verticillata var. evoluta.

4. Bogs with “Red Tussock” as the Dominant Species.

The grass Danthonia rigida var. rubra, generally known as the Red Tussock, occurs throughout eastern and southern Southland on open hillsides, by stream courses, in swamps, and on peat bogs, but only on such peaty areas as are not continously water-logged. In swamps it may attain an overall height of four feet or more, but normally it is less than three feet tall. In swampy areas cryptogams are few, but on drier peaty soils the three species of the sub-genus Clathrina are common associates on the ground, with several red fruited Cladonias, and the brown-fruited C. cornutoradiata, C. carassensis, C. pityrea and C. verticillata common on the mounds left by dead tussocks or less commonly on the lower “trunks” of the largest tussocks and even on the peat itself, where C. aggregata in several forms is the commonest lichen. Two agarics, one yellow with decurrent gills and the other dull red, and both fairly small, were common. Campylopus introflexus and C. torquatus cover patches of some extent, the latter almost always sterile, and with the same two Sphagna noted elsewhere comprise the moss flora. Marchantia berteroana is present but rare.

5. “Manuka” Swamp-bog.

This is the driest type of bog, and in summer the surface may be quite dry. After rain the area is really swamp rather than bog, though the soil is peat several feet deep. As areas are drained, manuka (Leptospermum scoparium) speedily spreads to areas formerly occupied by Hypolaena, Phormium or Danthonia, and rapidly overtops and kills these by exclusion of the light. On these soils it is on the shady side of the manuka groves or for a yard or so within the margin that the cryptogamic flora is most abundant. Many groves are separated from neighbouring groves by open spaces only a few yards wide. These areas often have epacrids and heaths or perhaps Hypolaena forming an open shrubbery in which cryptogamic plants are exceedingly numerous.

Where the manuka groves reach up to ten to fifteen feet in height the interior is commonly open and diffused light penetrates freely. In such stations the ferns Blechnum procerum, B. penna-marina, Histiopteris incisa, and more rarely Polystichum vestitum may be found, or even the scrambling Lycopodium volubile, seen but once by the writer. In other areas the presence in the manuka groves of dead Danthonia tussocks and decaying “flax” bushes is evidence of aggression by the manuka following the drying of the soil consequent on drainage operations. Where the manuka is under ten feet tall the penetration of light is difficult and no crytogams are encountered save for a distance of two or three feet within the margin of the grove. This is a very common location for Cladonia retipora—the Coral Lichen, and for C. leptoclada and C. alpestris; less commonly C. crispata, C. Boryi, and C. cornutoradiata occur in this station. C. aggregata in two principal forms is also found here but is commoner in areas just clear of the manuka.

as A.47, T.1912, and S.150: det. W. M.): Kelly Range D. S. I. R. No. 2396; det. W. M.).

4. Cladonia pycnoclada (Pers.) Nyl.

Hitherto this has been regarded as the commonest member of the New Zealand Cladinae. As mentioned above, Santesson has stated that the New Zealand plants so named by Vainio were not that species. Through the courtesy of the director of the D. S. I. R. I. have examined the extensive collections housed in their herbarium and find that not one is P+, as all plants of the South American species are, and all lack perlatolic acid. Morphologically the two species are much alike, but it is clear that if important chemical distinctions are to be maintained as specific characters then “f. flavida”, as many are labelled, must be replaced by C. leptoclada, and “f. exalbescens” by C. impexa. However, these two “forms” merge in so many cases so that it is difficult to relegate many specimens to the correct group. The difference in colour is the best guide, but intermediate shades exist so that no clearly defined colour boundary separates C. leptoclada from C. impexa. Indeed, C. leptoclada and C. impexa, but for the presence of both in New Zealand, might have been regarded as mere geographic races of a single species—namely, C. impexa.

5. Cladonia leptoclada Des Abbayes.

This species was segregated from C. impexa by Des Abbayes to incorporate plants from Central Africa, New Caledonia and New Zealand which in the mass had a different aspect and a yellower colour. The South American equivalent was similarly segregated as C. confusa by Santesson. The differences between these three species (?) are so slight that it would often be necessary to know the source from which a specimen had been obtained before it could be named (4). All three contain usnic and perlatolic acids; all three show dichotomies and trichotomies round open or closed axils; all tend to produce sympodia in their lower parts; while their surfaces and ultimate branchlets are very similar. Apart from slight differences in density of branching and in the colour, there is little to differentiate this species from C. impexa C. pycnoclada lacks perlatolic acid and is P+, and though C. leptoclada may sometimes be lacking in perlatolic acid, it is invariably P− What has formerly been regarded in New Zealand as C. pycnoclada var. flavida must now be labelled C. leptoclada, though with the reservation that this species may ultimately be merged into C. impexa, from which it had been segregated, as a colour form.

Localities previously recorded include Kaiteriteri, in Nelson; Cleddau Valley, in West Otago; Awarua, in Southland; Lake Manapouri; and Stewart Island. The following specimens in Herb. D. S. I. R. are transferred from C. pycnoclada to C. leptoclada Mt: Egmont (A. 2); Herekino (A. 3); Whangarei (A. 12 and A. 20); Little Barrier Is. (A. 9), Hen and Chicken Islands (A. 15), Poor Knights Islands (A. 26); Mt. Moehau (A. 24, A. 27)., Rangitoto Is. (A. 28, A 30, A 45); Kaihere (A. 42), Kaingawa (A. 7); Matauhi Bay (A. 8), Ruakura (A. 21), Rotorua (A. 14); Hutt Valley (A. 18); Bull Mound (A. 19); Chelsea (A. 22); Kirita Bay (A. 31); Mt. Maungatua (A. 6, A. 41); Lee Stream (A. 32); Verterburn (A. 34); Greymouth(A. 33, A. 36, A. 37).

6. Cladonia impexa Harm.

This species was deemed almost cosmopolitan by Des Abbayes, who included both New Zealand and Hawaii in its range as well as South America. Magnusson soon showed that Hawaiian plants were distinct and named them C. Skottsbergii. Similarly, Santesson segregated South American plants as C. confusa, and Des Abbayes himself segregated C. leptoclada, leaving C. impexa in its restricted connotation as a plant possibly confined to the Northern Hemisphere. Evans (4) who

absence of algal cells in the thallus, the septation of the spores and whether parasitic or not on other lichens. Some of Vainio's genera are classed as fungi, but it seems that in a few species some specimens have algal cells and others not. The division between species with simple and with 1-septate spores is also not clearly marked, although the majority fall readily into one group or the other. I have included all these variations within the genus, but divided the New Zealand species among three subgenera, one of which is new. All the New Zealand species have whitish cvanescent or obsolete thalli.

Calicium abietinum Pers. var. abietinum

Calicium abietinum Pers, Dispos. Meth. Fungor., 59 (1797).

Calicium curtum Borr, Bab. in Hook Fl. N. Z. Vol. II, 304 (1855).

Nyl. Synops. Lich. I, 156 (1860).

Hook Handb N. Z. Fl., 558 (1867).

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

Hellbom, Bihang Kgl. Sv. Vetensk. Akad. Handl., 21, 111, 130 (1896).

Calicium abietinum var. denigratum (Vain) Zahlbr. apud Szatala, Borbasia, 1, 55 (1939).

Thallus whitish, very thin or evanescent, more or less homoiomerous with scattered bright green algal cells; apothecia (including stipe) up to 2 mm high but usually 0.8 to 1 mm, stipe 0.1–0.2 mm thick, black with hyaline outer layer; capitulum more or less turbinate (topshaped), 0.2–0.5 mm wide, white pruinose beneath or marginally only, or epruinose; spores dark, ellipsoid, 2-celled, distinctly constricted at septum, 7–12 × 5–6μ with roundish loculae. (Description adapted from Vainio, 1927, p. 41.)

Habitat. On old wood.

Distribution. More or less cosmopolitan. Otago: Flagstaff, 4,600; Merton, 1649.

This is apparently the only species previously found in New Zealand, having been reported from the North Island by Babington, Szatala and Kirk, and from Otago by Hellbom. The species is evidently a variable one in Europe, and several varieties and forms have been reported from there. I am not quite certain which should be regarded as the typical variety, but have taken it to be Vainio's var. glaucellum (C. glauccllum Ach.) since this is apparently much the commonest form in the Northern Hemisphere. According to Vainio (1972), C. abietinum f. denigratum is an inconstant form differing from the typical variety in the less developed thallus and total absence of pruina on the apothecia; actually it is ranked as a form also by Zahlbruckner.

Calicium abietinum var. australe Murray, var. nov.

A C. abietino Pers. var. abietino differt stipite tenuiore, capitulo parviore et nudo, et sporis majoribus. Thallus evanescens.

Thallus whitish or greyish, very thin or obsolete, containing a few trebouxioid algae among the hyphae; apothecia about 0.8 mm high, stipe tapering upwards from about 0. 18 mm at base to 0.07–0.09 below capitulum; capitulum subcylindrical to rather turbinate, black and somewhat shining, epruinose, K+ pale brown, about 0.2 mm dia. and 0.18 mm high with mazedium forming a nearly flat top to 0.3 mm dia. Apothecia dark brown in section with hyaline outer layer of ill-defined structure 8–9μ thick; paraphyses few, very slender, simple and aseptate; asci cylindrical ca. 55 × 5μ; spores in one series, dark brownish or smoky grey, oblong to ovate, 1-septate, definitely constricted at septum, (7½−) 12–15 (−18) × (5−) 6–7½ (−8)μ wall and septum 1½μ thick (spores still in the ascus are hyaline, 1-septate, 7½ × 4½μ). Pycnidia not seen.

Habitat. Old wood.

Distribution. Otago: Taieri Beach, 3718; Akatore Gorge, 3725. Southland. Forest Hill, 3474 (Type).

The variety differs from typical abietinum principally in the larger spores and total absence of pruina. Var. meizopus Vain. has similar large spores but the apothecia are 1.5–2mm high and the capitulum is broader and more turbinate; only the type specimen of this variety seems to have been reported.

Calicium sphaerocephalum (L) Ach.

Mucor sphaerocephalus L., Sp. Pl. 1185 (1753).

Calicium sphaerocephalum Ach., Method. Lich. 91 (1803).

Thallus white to pale cream, very thin or evanescent, penetrating within the substrate and containing scattered trebouxioid algal cells (and occasional Nostoc cells which are probably chance contaminants); apothecia about 2 mm high, stipe black, epruinose, tapering from 150μ at base to 100μ at top, capitula dark brown or nearly black, cup-shaped, 300–350μ diameter and 300μ high; excipulum without hyaline layer but reddish pruinose and K+ reddish-brown, mazedium convex, spores dark brown, all 1-septate, rather strongly constricted at septum, oblong to ovoid with nearly spherical cells, 6½-9 × 5μ.

Habitat. On old wood.

Distribution. Northern Hemisphere, (?) South America, Australia (as varieties). Southland: Forest Hill, 0265a (pr p. with Calicium subnigricans).

The single small specimen seen (consisting of six apothecia) has slightly smaller capitula and spores than usual for European material (Vainio, 1927, gives the capitula as 350–800μ dia. and the spores (6-) 10–13 × (3-) 5–7μ for Finnish specimens), but in other respects it agrees closely. There are pycnidia on the specimen, but I am uncertain with which species they are associated. The Australian C. trachelinum var. elattosporum Wilson is similar but is said to have still smaller spores (3–8 × 2–4μ), although Wilson's microscopic measurements on other lichens are often rather small, presumably due to a calibration error. I have not used this variety name without seeing material of it, in any case the name could well be dropped because of possible confusion with Calicium elassoporum Nyl.

Calicium floerkei Zahlbr.

Calicium floerkei Zahlbrucknei, Cat. Lich. Universalis 1, 598 (1923–40) (nom. nov. for Calicium pusillum Hepp., non Flk.)

Thallus very thin. up to 30μ thick on surface of substrate and penetrating beneath, whitish to pale creamy-white or partly evanescent, green globose algal cells scattered in surface layer; apothecia about 350μ high. stipe light brown, 75–100μ thick at base tapering to 35–60μ at top, without any hyaline layer; capitula top-shaped, 100–200μ dia, and same in height, dark brown, epruinose, K+ pale brownish; hymenium 75–80μ thick, hypothecium pale brown, conical; paraphyses more or less conglutinate, 1μ thick; asci cylindrical, 45–60 × 3μ, 8-spored, spores smoky grey (tuining brown with K), oblong to almost cylindrical, 1-septate with some simple thin-walled, 6–8 (-12) × 2.5–2.8μ not constricted at septum. Pycnidia not seen.

Habitat. On old bark.

Distribution. Europe. Otago: Mt. Cargill. 1, 500ft, T 2165 (on Libocedius bidicillii).

The specimens consist of several pieces of bark with an unrecognisable lichen of which little remains but disintegrating lecideine apothecia, and which is covered with the minute fruits. It agrees very closely with Vainio's description (1927) of Calicium subpusillum Vain. and Embolidium italicum Sacc. (=Calicium pusillum Hepp), which apparently differ significantly only in that the former has algal cells in the thallus and the latter not. In the New Zealand specimen some of the apothecia seem to have no gonidia within 100μ of their bases, while others are clearly associated with a lichen thallus, although the visible thallus, particularly the white portions, may in part belong to the decomposing lichen In some parts of the thallus the lichen hyphae were observed to produce thin-walled globose (?) chlamydospores, 5–8μ dia.

Calicium subnigricans Murray, sp. nov.

Thallus tenuissimus aut vix ullus, subalbidus, laevigatus, K-, parcibus gonidiis viridibus; apothecia altitudine 0.5–0.7mm, stipite 60–80μ crasso, dilute fuscescente, pellucido, intus distincte pallidiore; capitulo turbinato, major minusve albido-pruinoso, 200μ diametro et 100μ altitudine, K-; asci cylindrici, 8-sporae; sporae monostichae, pallidae vel griseae, semper simplices, oblongae, 4½ × 2.2μ, mazedium bene evolutum, nigricans, hemisphaericum; paraphyses 1½μ crassi, subsimplici, irregulares, pycnidia non certe visa.

Thallus marked by a whitish area containing a relatively small proportion of? trebouxioid cells 8μ dia. scattered singly and in groups among the thin hyphae and decomposing substrate cells; stipes 80μ dia. at base tapering to 60μ below capitula, structure not clearly seen but outer 15μ a slightly deeper brown than the centre, which is a pale straw colour. The whole capitulum is nearly globular, the upper two-thirds of which is the dark grey mazedium; the excipulum in young apothecia is white pruinose, the pruina disappearing in older fruits; the spores are all simple, oblong to somewhat ellipsoid, hyaline but darkening in the mazedium to a smoky grey.

Habitat. Old wood.

Distribution. Southland; Forest Hill, 0265b (pr. p. with Calicium sphaerocephalum) on dead Griselinia littoralis (Type).

This species with its simple spores falls in Mycocalicium, a group which has been variously interpreted in the past as including Calicium species without algae, in the thalli (i.e., fungi), lignicolous non-lichenized species with brown simple spores, and Calicium species with all simple spores whether lichenized or not. I have preferred to follow the last of these interpretations (Santesson, 1943) since it is not always possible to be sure whether algae are truly associated with the fungus hyphae in certain species, or indeed whether the apothecia are certainly derived from the thallus on which they are growing. C. subnigricans resembles in the nearly colourless spores C. arenarium Hampe (Coniocybopsis arenarium (Hampe) Vainio), a species of fungus parasitic on lecideine lichens, and differs from it principally in the smaller capitula with more developed mazedia and smaller spores, as well as in the habitat. The white pruina may not be a constant character.

There is evidently another New Zealand species in the Mycocalicium subgenus, judging by a North Island specimen from Hawke's Bay (Colenso, 3596, WELT). It is not complete, the capitula being reduced to the narrow excipula on dark brown stipes of uniform structure, K+ purplish; thallus none remaining; spores still clinging to the stipes are brown, simple, broadly ellipsoid, 7½ × 3–5½μ It is no longer completely identifiable, but is not referable to any European or South American species. The Australian Mycocalicium australicum Räsänen appears very similar but for slightly longer spores.

Crassistipitum Murray, subgenus Calicii novum.

Thallus tenuis vel obsoletus; stipites apotheciorum magis crassi, a capitulis non bene distincti, totum apothecium subcylindricum vel infundabiliforme; sporae pro majore parte nigricantes, uniseptatac.

The new subgenus is separated from Calicium proper by the relatively broad stipes which are not clearly delimited from the capitula. The apothecial margin and

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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,

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.

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).

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.

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.

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.

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

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

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

.

Xanthoria novozelandica Hillm.

Xanthoria novozelandica Hillmann. f edde, Repert. spec. nov. 45, 176 (1938).

Zahlbr., Lich. N. Z., 123 (1941).

Thallus orbicular, appressed, 5–30 (-50) mm dia., yellow or greenish-yellow or greygreen, K- or pink in places, with smooth marginally imbricate lobes 1–3 mm long, 0.5–1.2 (-2) mm wide, 50–90μ thick and minutely crenulate, white beneath, shortly rhizinose, with submarginal horizontal white fibrils projecting about 0.2 mm beyond margins (occasionally very few or none)., upper cortex 12–25μ thick, plectenchymatous with isodiametric 4–6μ dia. cells, algal layer 25–50μ thick, filling all the space between the cortices, algae Trebouxia 8–14μ dia., lower cortex 15–30μ thick, similar to upper cortex. Apothecia covering most of the plant in larger specimens, 1–1.8 (-3) mm dia., shortly pedicellate on pedicels 0.2 mm high, excipulum plane, with a few white fibrils 0.2–0.3 mm long projecting downwards; disc plane or slightly convex with thin entire and finally obsolete margin; hypothecium hyaline, obscurely cellular, 25–30μ thick; hymenium (65-) 75–80μ high including granular epithecium. paraphyses 1½μ thick, simple or furcate, asci clavate-cylindrical, 45–50 × 13–15μ, spores rather variously shaped, polaribilocular, (11-) 13 × 6½-7½μ, r = 0.2–0.4^* Pycnidia globose, 200μ dia, singly in prominent orange warts on upper surface, pycnidiospores cylindrical, not constricted, 3 × 0.8μ.

Habitat. On bark, or rarely twigs.

Distribution. New Zealand Canterbury: Hurunui (A. J. Healy, 58/499) CHR (uncertain), Waitohi R (A. J. Healy 58/520) CHR;, Cheviot (A. J. Healy) CHR;, Godley Valley (H. H. Allan, H64) CHR (Isotype), Sc 170. Otago: Lake Ohau (R. Jones) 1721, Matukituki R. (R. F. Smith) 1192;, Stewart's Gully, T 1993 (and in CHR as X. parietina f. virescens), Dunedin 1226; Flagstaff, 1,200ft, 3633; Brighton, 021b (uncertain) Southland Kaiwera, Mr; 717, Gore, Mr. 899.

The species seems to be rare, although it is evidently quite widely distributed. The material is not very uniform and further collecting may confirm the existence of subspecific taxa. A few specimens from the coastal regions have scarcely any fibrils or even none, and then only differ from small forms of X. parietina in the

thinner, narrower crenulate marginal lobes with prominent pycnidia. One specimen, A. J. Healy 58/499, has many large apothecia as well as lacking fibrils, and looks very like Xanthoria polycarpa except that the lobes are uniformly flat.

Xanthoria parietina (I) Th. Fr. var. parietina.

Lichen parietinas L., Spec. pl. 1143 (1773).

Xanthoria parietina Th. Fr., Lich. Arctoi, 67 (1860).

Müll. Arg., Bull. Herb. Boissier II, 40 (1894).

Hellb., Bihang Kgl. Svensk Vetensk. Akad. Handl., 21, (13), 51 (1896).

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

Cranw. & Moore, Rec. Auck. Mus., 1, 314 (1935).

Zahlbr., Lich. N. Z., 122 (1941).

Physcia parietina D. Notr.

Nyl., Synops. Lich., I, 410 (1860).

J. Linn. Soc. Bot., 9, 249 (1865).

Lich. N. Z., 45 (1888).

Linds., Trans. Linn. Soc., 25, 518 (1866).

Hook., Handb. N. Z. Fl., 573 (1867).

Physcia parietina var. platyphylla Krmph., Reise d. Novara, Bot. I, 116 (1870).

Physcia ligulata Krb., Reliq. Hochstett., 30 (186?) (sec. Nyl., Lich. N. Z., 45 (1888).

Parmelia parietina Ach.

Hook., Fl. N. Z., 287 (1855).

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

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

Thallus 2–5 (-10) cm dia., yellow or orange-yellow or greenish yellow, more or less orbicular, lobed at circumference, lobes 3–6 mm long × 2–5 mm wide, smooth to irregularly wrinkled or weakly folded or pitted with raised, often sinuate margins, white beneath or yellowish peripherally, with a few short simple white rhizines; upper cortex 10–25μ thick, plectenchymatous of vertical hyphae forming cells about 5μ dia., algal layer 30–40μ thick, algae Trebouxia, 8–15μ dia;. medulla 10–80μ thick of moderately thick-walled 3μ dia. hyphae varying from fairly compact to loosely woven. lower cortex similar to upper cortex, 8–15μ thick. Apothecia sessile, well constricted at base, 1–5 mm dia., margin thin, persistent, often inrolled; disc concave, usually darker than thallus, smooth; hypothecium and excipulum hyaline, 20μ thick; hymenium hyaline except for the orange granular epithecium, 75–90μ high;, asci 45–55 × 15μ, spores hyaline, polaribilocular, ellipsoid, 11–14 (-16) × 6½-9μ, r = 0.4–0.6 Pycnidia immersed in orange warts, globose, pycnidiospores 2–3 × 1–1 2μ.

Habitat. On trees, rarely on rock or other substrates.

Distribution. Apparently cosmopolitan in temperate regions. North Island:

Thames, CHR; Kaitake (O. J. Oates) 021c; Feilding (H. H. Allan) CHR, (A. J. Healy 468) CHR; Bulls (B. J. Sweetman) 4413; Netherby (R. Mason and N. T. Moar) CHR; Hawke's Bay (Colenso, 6219) WELT. Canterbury: Port Hills, CHR; Temuka, Mason, 251 and 252. Otago: Saddle Hill, Mr., 485, Mr., 7349; Mosgiel, 058; Lee Stream, T 207; Hindon, T 247; Kaiwera, Mr., 1318; Pukerau, Mr., 490 (pr. p.), Mr., 5418. Southland: Forest Hill, 021e, 049.

The typical variety of the species may be introduced, since I have rarely seen it far from settlements, and it seems to favour exotic trees, usually fruit-trees. It seems to be common in parts of the North Island, although I have seen only a few North Island specimens in herbaria. Like many other wide-ranging species, X. parietina is a variable plant and more than 30 distinct varieties and forms have been described, mostly on European material. I have not been able to see all of these as descriptions or specimens, and furthermore some of the named specimens have not agreed with the descriptions; consequently the identification of the New Zealand specimens with European subspecific taxa is sometimes open to doubt. In certain cases the local material is close to but not clearly identical with the European forms although the differences are not easily expressed in quantitative, terms Although the genus has been surveyed twice by Hillman (1922, 1935) the

separation and nomenclature of these subspecific taxa still show unsatisfactory features, and for this reason I have included several distinct forms under var. ectanea without attempting to name them.

Xanthoria parietina (L) Th. Fr. var. parietina f. albicans (Müll.) Hillm.

Teloschistes parietinus f. albicans Müll. Arg., Flora 338 (1890).

Xanthoria parietina f. albicans Hillm., Ann. Mycol., 14 (1920).

Zahlbr, Lich. N. Z., 122 (1941).

As for the typical form but thallus fairly thin, flat, whitish, K-;. apothecial disc yellow.

Distribution. South America, South Africa, New Zealand and probably elsewhere. North Island: Kaipara (L. B. Moore, ZA125) CHR. Southland: Forest Hill, 046.

This form does not seem significantly different from f. cinerascens (Light) Sandst., and the two plants seen are hardly distinct from the normal form of the species or from f. chlorina.

Xanthoria parietina var. parietina f. chlorina (Chev.) Oliv.

Imbricaria chlorina Chev., Fl. Envir. Par. I, 62 (1826).

Xanthoria parietina f. chlorina Oliv., Mem Soc. nat. Sciences natur. Cherbourg, 36, 228 (1906–7).

Thallus thickish, grey-green centrally with pale yellow raised sinuate margins; apothecia concave with grey margins and yellow discs.

Habitat. On shaded tree-trunks, etc.

Distribution. Europe, and presumably elsewhere. North Island: Hawke's Bay.

(Colenso, 6219 pr. p.) WELT. Otago Mosgiel 021.

Xanthoria parietina var. parietina f. virescens (Wedd.) Sandst.

Physcia parietina f. virescens Wedd., Bull. Soc. bot. Fr., 16, 198 (1869).

Xanthoria parietina f. virescens Sandst., Abh. Naturw. Vereins Bremen, 223 (1911–2).

Thallus rather small and thin, green with yellow apothecial discs; otherwise as in the typical form. A shade form.

Distribution. Europe, but probably cosmopolitan Canterbury. Temuka, Mason, 265.

Xanthoria parietina (L) Th. Fr. var. adpressa Mereshk.

Xanthoria parietina var. adpressa Mereshk., Hedwig. 61, 209 (1919).

Lobes shorter, thinner and narrower (1–2 mm) than in the typical form of var. parietina, smooth, adpressed, margins not sinuate; medulla very thin; apothecia (½-) 1–3 (-4) mm dia., sessile, nearly plane, becoming convex with excluded margin; hypothecium 25μ, hymenium 52–60μ, spores 11–15 × 5–7μ.

Habitat. On trees, often in exposed situations.

Distribution. Asia, Europe. Canterbury: Leithfield (A. J. Healy, 58/530) CHR (pr. p.); Hurunui (A. J. Healy, 58/505) CHR; Lake Tekapo, 1,500–2,000ft, Mason, 8, 245, 246 and 247. Otago. The Kaik, Mr., 603, Saddle Hill, Mr., 488, Mr., 7349. Chatham Islands: (Colenso 9) WELT.

Some of the above specimens are not well separarted from var. parietina and may not truly belong to var. adpressa, of which I have seen no authentic specimens. They seem to differ significantly from var. parietina in the narrower spores as well as in thallus features.

Xanthoria parietina (L) Th. Fr. var. aureola Rostr.

Parmelia aureola Ach, Lich univ, 487 (1910).

Xanthoria parietina var. aureola Rostr; Bot. Tidsskr, 4, 96 (1871).

Hellb., Bihang Ggl. Svensk Vetensk. Akad. Handl., 21 (13). 51 (1896).

Müll. Arg., Bull. Herb. Boissier, II, app. 1, 40 (1894).

“Thallus stout, rigid, dark golden-yellow or orange-yellow, knotted-warty and wrinkled centrally; apothecia with crenate margin.” (Description translated from Anders “Die Strauch —und Laubflechten Mittelseuropas” p. 192 (1928)).

Habitat. Mostly on rocks.

Distribution. Eurasia, New Zealand.

I have not seen specimens which clearly belong here, although most large specimens of var. parietina are more or less wrinkled and some have a few apothecia with subscrenulate margins.

Xanthoria parietina (L) Th. Fr. var. ectanea (Ach) Grönl.

Parmelia parietinaβectanea Ach., Lich. univ. 464 (1810).

Xanthoria parietina var. ectanea Grönl., Bot. Tidsskr. 4, 161 (1871).

Nyl., Synops. Lich., 410 (1860).

Zahlbr., Lich. N. Z. 122 (1941).

Physcia parietina Oliver, Trans. N. Z. Inst., 44, 87 (1911).

Parmelia elegans Hook, Fl. N. Z., 291 (1855).

Placodium elegans Hook., Handb. N. Z. Fl., 577 (1867).

Nyl., Lich. N. Z., 58 (1888).

Caloplaca elegans Hellb, Bohang Kgl. Svensk Vetensk. Akad. Handl., 21 (13), 66 (1896).

Allan, Tuatara 1, 21 (1948) (pr. p.).

Caloplaca elegans var. tenuis Cranw. & Moore, Rec. Auck. Museum, 1, 314 (1935).

Zahlbr., Lich. N. Z. 121 (1941) (pr. p.)

Thallus yellow to orange, white or pinkish beneath, lobes more or less radiate, smooth, plane or slightly swollen or channelled, 2–10 mm long by (½-) 0. 8–1. 5 mm wide, linear, sparingly branched, separated, contiguous or imbricate, adpressed or with raised edges; upper cortex 12–25μ thick, outer part interspersed with yellow granules; algal layer 25–40μ thick, medulla usually loosely woven, 70–140μ thick; lower cortex 15–20μ thick; apothecia concolorous with the thallus, ½-1-½ (-2½) mm dia., plane to convex, margin usually persistent; hypothecium and proper margin 40μ thick, hymenium (55-) 65μ high, paraphyses 2μ thick, asci clavate to irregularly saccate, about 40 × 15μ, spores ellipsoid, (?8-) 13 (-16) × (5½-) 6–87½μ, r = about 0. 5.

Habitat. Usually on coastal rocks.

Distribution. More or less cosmopolitan.

(a) Yellow, forming a mass of imbricated and epithalline lobes 2–4 mm long × 1 mm wide, not appressed, weakly concave or channelled, medulla 70–100μ thick, spores about 14 × 7½μ.

Otago: Port Chalmers, T 703 (ZA 53 in CHR). Southland: Doubtful Sound, 3963.

(b) Forming circular patches of contiguous and/or imbricate lobes, medulla 90–120μ thick, spores about 13 × 8μ.

North Island: Hen and Chickens Ids. (L. M. Cranwell) CHR; North Wairoa River, CHR; Rangitoto (H. H. Allan) CHR; Coromandel Peninsula (L. B. Moore, ZA 126) CHR; Whangarei (W. A. Given) CHR; Hutt Valley, on bone (H. H. Allan) CHR; Wellington, Mr., 584. Marlborough: Goose Bay, Mr., 7335. Nelson: Karteriteri, Mr., 1248 (pr. p.). Otago: Cornish Head, 3883 (contorted from with large apothecia); Harbour Cone, 500ft, Mr., 5419; Waikouaiti, T 30 (on bark), T 2928; Howell's Point, T 818 (uncertain); Portobello, 3513, 3514; Blackhead, T 403; Mosgiel (on tree) 058 (pr. p.); Brighton, 021a. Southland: Riverton, T 787. Stewart Island: (Cockayne, 0836) CHR.

(c) Lobes stellate-radiate, discrete, sometimes plano-convex, appressed, spores 13 × 8μ (perhaps f. ratilans (Ach.) Zahlbr.).

North Island. Ahuriri (Colenso, 2964) WELT; (?) Hawke's Bay (Colenso, 6244) WELT. Westland Greymouth (W. A. Mackay) CHR Nelson; Kaiteriteri, Mr., 1248 (pr. p.); Maitai Valley, Mr., 7326 Otago: Akatore Estuary, 3730, 3731.

(d) Thallus 4 cm dia., orange-red, lobes stellate-radiate, 10–20 mm long × 0.5 mm wide, 0. 2–0. 5 mm apart, medulla about 100μ thick, spores 5½-6½μ wide.

North Island: Whangarei (W. A. Given) CHR (pr. p., on bark).

(e) Lobes irregularly humped 1-1½ mm wide, contiguous, medulla loose, 120–150μ thick, apothecia very convex, hymenium 55μ high, spores 8½-10½ × 6-½-8μ.

Otago: Kyeburn, 1,000ft, T 41 (and in CHR). Southland: Ben Bolt, 1,500ft, 1191.

Except in the few cases where a height or substrate is given, all the above specimens were collected from coastal rock, where the variety is common, often invading the Verrucaria zone; elsewhere it seems to be very rare. The distribution pattern is quite similar to that recorded in Britain for var. ectanea (Smith, 1922). I have put the local specimens into five groups for which names may be already available. Groups (c) and (d) look deceptively like Caloplaca elegans from which they are only distinguished by the rather flatter less regular lobes which are never partly hollow; group (c) usually occurs near high-tide mark or in exposed situations, and seems only a trivial form of the variety, but the plant in group (d) may be distinct although it does not appear to belong to any of the recorded varieties of Xanthoria parietina with such narrow lobes. The small specimens in group (e) from subalpine situations are hardly separable from the plants in group (b) superficially, but are certainly distinct in spore measurements. Although the usual specimens of var. ectanea are easily distinguished from var. parietina, intermediate specimens are not uncommon—e. g., 021a, Mr. 584, T 787 above.

Much more collecting and field studies are needed, particularly in the North Island, to decide the status of the forms grouped together under var. ectanea here.

Xanthoria parietina. (L) Th. Fr. var. incavata (Stirt.) Murray, comb. nov. Physcia incavata Stirt., Trans. N. Z. Inst., 32, 82 (1899).

Xanthoria incavata Zahlbr., Lich. N. Z., 123 (1941).

Thallus orbicular, yellow, flat, more or less continous centrally with marginal lobes 3–5 mm wide × 5–8 mm long and up to 0.2 mm thick, contiguous, broadly rounded, entire, matt to rather arachnoid or minutely pitted, without longitudinal wrinkles, appressed, white beneath; medulla arachnoid, up to 150 thick; apothecia brodly sessile, initially plane with thin margin becoming convex with excluded margin, orange, 2–3 mm dia hymenium 75μ high, spores ellipsoid to reniform 13–15 × 8μ pycnidia. immature.

Habitat. On bark.

Distribution. New Zealand. Marlborough: Molesworth (A. J. Healy) CHR; Mt. Tapuaenuku, 4,000ft, T 1526. Canterbury: Selwyn Gorge (A. K. Beckett), Canterbury Museum. (Type, or perhaps isotype specimen).

The specimens I have seen are not very uniform, and the plant is evidently not specifically distinct from X. parietina. In var. parietina specimens with lobes as wide as 5 mm are always ascending at the margin, somewhat imbricate and more or less wrinkled, with the apothecia larger and hardly convex, but the microscopic details are probably not significantly different. The type (? or isotype) specimen is only a fragment with two lobes and has been damaged. Var. incavata seems close to the South African var. macrophylla (Stizb.) Hillm. which, however, has pale apothecia with shining waxy margins.

Caloplaca elegans Th. Fr. var. elegans.

Thallus orbicular, orange-red, lobes radiate, contiguous or partly imbricate and loosely attached, about 1 mm wide, plano-convex, sparingly branched, surface often minutely pitted centrally, medulla arachnoid or or lobes partly hollow, apothecia 06–12 mm dia., broadly sessile, concolorous, margin persistent; hymenium 60μ high, spores 10½-13 × 6½-8½7mu; r=0.2–0.3.

Habitat. On rock in alpine, arctic or antarctic regions.

Distribution. Northern and alpine Europe, Himalayas, North America, New Zealand, Polar regions Canterbury: Mistake Peak (Tekapo), 6,200ft, Sc 39; Godley Valley, over 3,000ft, Sc 168 (pr. p.), Sc 217, Sc 218; Ben Ohau Range, 6,000ft, Mason, 153.

Physcia chrysophthalma Nyl., Synops. Lich., 410 (1860).

J. Linn. Soc. Bot., 9, 249 (1865).

Lich. N. Z., 45 (1888).

Linds., Trans. Linn. Soc., 25, 518 (1866).

Hook, Handb N.Z. Fl., 572 (1867).

Parmelia chrysophthalma DC., Buch; Trans. N.Z. Inst., 6, 231 (1873).

Hook, Fl. N.Z., II, 287 (1855).

Thallus small foliose-fruticose, in small clumps to 4 cm dia. and ½–2 cm high, usually formed from single plants. Thallus initially of stellate-radiate laciniae ½ mm wide, eventually forming erect, branched, rigid, dorsiventral lobes up to 2½ mm wide with almost spinous marginal fibrils, golden or orange yellow above particularly near ends of lacinae, matt or slightly shining, smooth or weakly longitudinally ridged; white or partly yellow below and usually longitudinally veined and wrinkled; cortex 20–50 (-100)μ thick, very uneven, of horizontal or periclinal thick-walled hyphae; algal layer mostly on upper side, discontinuous, algal cells Trebouxia, 10–15μ dia.; medulla of loosely woven 3μ dia. hyphae or lobes partly hollow; lower cortex like upper but more uneven. Apothecia on the ends or margins of small laciniae, 1–6 mm wide, nearly plane or older ones convex, disc orange-yellow, margin thin, slightly elevated, with up to 150 concolorous fibrils ½-2 mm long; hypothecium hyaline, 10–25μ thick, hymenium hyaline except for granular yellow epithecium, 40–55μ high; paraphyses simple or furcate, 1½μ thick, thickened at tip; asci almost ellipsoid, about 35 × 13μ; spores ellipsoid, (10-) 13–15 × (3½-) 5–6 (-8) μ, r= 0. 25–0. 5 Pyenidia not seen on New Zealand specimens.

Habitat. On bark.

Distribution. Probably cosmopolitan. North Island: New Lynn (L. M. Cranwell) CHR; Atiamuri, Allison—, Taupo (Colenso, 998) WELT; Waitangi (Hawke's Bay, Colenso, 3008, 3540 and 3555) WELT; Napier (Colenso) WELT; Canterbury; Waipara (A. J. Healy, 58/493) CHR; Leithfield (A. J. Healy, 58/530) CHR; Temuka, Mason, 253; Timaru (H. H. Allan) CHR; Otago: Waikouaiti, T 27, T 334; Macrae's Hill, T 2481, Mt. Royal, T 322 (pr p); the Kaik, Mr., 579, Taiaroa Head, T 1023, Saddle Hill, 1,000ft, Mr., 852; Taieri, 3559; Akatore R., Mr., 583; Balclutha (H. H. Allan) CHR. Southland Waikaia, Mr., 5414; Kaiwera, Mr., 1317; (pr. p.), Gore, Mr., 606; Forest Hill, 018.

Teloschistes chrysophthalmus var. chrysophthalmus f. cinereus (Müll. Arg.) Zahlbr.

Teloschistes chrysophthalmus var. cinereus Müll. Arg., Flora 63, 265 (1880).

Teloschistes chrysophthalmus f. cinereus Zahlbr, Cat. Lich. univ., 7, 316.

Lich, N.Z., 123 (1941).

Thallus and apothecial margins greyish, K-, fibrils grey, or yellowish grey, thallus sometimes pale yellow tinted at ends.

Distribution. South America, New Zealand. Otago: Dunedin, 1220, 1221; Saddle Hill, Mr., 484. Southland: Kaiwera, Mr., 725, Mr., 5415; Invercargill, Mr., 5416.

Teloschistes chrysophthalmus var. chrysophthalmus f. denudatus (Hoffm.) Mull. Arg.

Platisma denudatum Hoffm; Descr Adumbr Pl. Lich., II, 23 (1794).

Teloschistes chrysophthalmus f. denudatus Müll. Arg. apud Zahlbi., Lich. rar. exs. no. 40.

Teloschistes chrysophthalmus var. denudatus Müll Arg., Bull Herb Boissier, II, app. 1, 29 (1894).

Thallus yellow or partly grey, apothecia usually larger than in the common form, margins without fibrils or with a few very short ones.

Distribution. Europe, New Zealand. North Island: Tauranga CHR; Kakariki (A. J. Healy) CHR (very small plants); Wellington Heads (A. J. Healy) CHR. Otago: Brighton, Mr., 776. Southland: Te Anau (W. G. H. Edwards) 1183, 1184.

Teloschistes chrysophthalmus var. chrysophthalmus f. subinermis (Müll. Arg.) Zahlbr.

Teloschistes chrysophthalmus var. subinermis Müll. Arg., Flora, 63, 265 (1880).

Teloschistes chrysophthalmus f. subinermis Zahlbr, Cat. Lich. Univ. 7, 318.

Lich. N.Z., 123 (1941).

Thallus grey, including apothecial margin which lacks fibrils; fibrils on laciniae few and short.

Distribution. Temperate Southern Hemisphere. Otago: Mt. Royal Station(Thomson, H 57=T 322 pr. p.) CHR.

The species is probably common throughout New Zealand, at least in coastal areas and on exotic trees, particularly in orchards. Like Xanthoria parietina var. parietina, it may be partly or entirely introduced, although it was certainly recorded by 1845. The above forms are not well separated from the typical form, which I take to be that called f. armatus by Hillmann (1930). Certainly these forms can hardly be maintained as distinct varieties as was done by Hillman. The North Island material is generally rather smaller and more delicate than typical Otago specimens, but both differ notably from Hillmann's description of European plants in the considerably lower hymenium and narrower spores (Hillmann gives the hymenium as 70–82μ high and the spores as broadly ellipsoid, 10–17 × 6–8μ). In the local specimens I have examined the spores are rarely more than 6½μ wide; in the Tauranga specimen under f. denudatus many of the spores measure 15 × 3½μ. If these differences prove to be significant taxonomically, presumably a complete set of new forms and varieties would be required.

In the Knight Collection in WELT are a number of specimens without locality which apparently belong to var. expallens Müll. Arg., with small yellow rosulate thalli with small entire margined apothecia. I have not seen any undoubted New Zealand specimens, and Knight's material probably came from Australia. However, in view of the discovery of the Australian T. sieberianus and spinosus in New Zealand, the occurrence of T. chrysophthalmus var. expallens here also cannot be discounted.

Teloschistes chrysophthalmus var. flavoalbidus (Krmphb.) Malme.

Physcia chrysophthalma f. flavo-albida Krmph, Verh K. K. zool-bot Ges Wien, 18, 322 (1868)

Teloschistes chrysophthalmus var. flavoalbidus Malme, Ark. f. Bot., 20A, 46 (1926).

Plant up to 5 cm dia, 3 cm high, forming an open cushion of relatively long, erect, linear, sparingly branched lobes ½–-1 (1½) mm wide with fibrils mostly terminal, pale yellow near tips of lobes, elsewhere grey-white, cortex 35–40μ thick, more even and with more horizontal hyphae than in var. chrysophalmus, apothecia small, yellow, with fibrillose margins, hymenium 50μ high, spores ellipsoid, 12–15 × 5. 2–6 5μ, r=0 3–0 5.

Distribution. Chile. Otago; Portobello, 3905.

Further collections may show this to be no more than a form of the species, although the few specimens in this collection seem distinct. Again the New Zealand material I have put here has narrower spores than described for the Chilean specimens, although there seem no macroscopic differences (Hillmann gives the spores as 10–14 × 7–8 .5μ).

Teloschistes fasciculatus Hillm var. fasciculatus.

Teloschistes fasciculatus Hillm, Fedde's Report Seec nov, 49, 176 (1938).

Zahlbr, Lich NZ, 123 (1941)

Zahlbr, Lich NZ, 123 (1941)

Thallus forming clumps ½–3 cm dia and up to 1 cm high of interlaced erect orange-red laciniae 3–10 mm long by 0. 15–0. 5 mm wide, sparingly branched, smooth and whitish beneath except in the upper portions, flat or caniculate above, matt or reddish pruinose, with margins granular sorediate or with whitish eroded spots; soredia concolorous, in minute elliptical soralia or sometimes a few patches of farinose yellow soredia present also, sparingly fibrillate on margins, fibrils concolorous, ½-1 mm long. Cortex of periclinal to longitudinal hyphae, the outer part heavily interspersed with orange granules, 10–50μ thick, algae very few, scattered at base of cortex, algal cells Trebouxia, 14–18μ, with many zoosporangia up to 30μ dia. present, lower cortex similar to the upper but hyphae more vertical. Neither apothecia nor pycnidia seen.

Habitat. On rock debris in subalpine situations.

Distribution. New Zealand. Otago: Mt. Sutton (Lake Ohau), 2,300ft, 1767;

Mt. Ida, T 2742; Alexandra (W. Martin) 1664, Mr. 7351; Kyeburn, T 24 (and in

CHR); Maungatua 2,900ft, 0339a, 1962, T 357 (and as H56 in CHR, isotype specimens).

A very distinct species, which is very rare in the type locality and elsewhere, although it is probably to be found throughout inland subalpine Otago. There is evidently another anthraquinone pigment present besides physcion, since application of KOH gives red needles and purple prisms of the potassium salts.

Teloschistes fasciculatus Hillm. var. nodulosus Murray, var. nov.

A varietate typicale differt lacinus brevioribus, subimbricatis ad terminalia nodulososorediatis, plantam plus caespitosam formantibus; apothecia pycnidiaque non visa.

Thallus forming patches 5–20 mm dia. and 2–5 mm high, of ascending more or less imbricate sparingly branched laciniae 1–4 mm long × 0.1–0.8 mm wide, mostly plane, pruinose or not, yellow or orange above, mostly white below, with a few concolorous fibrils on margins and sometimes also on upper surface; margins entire towards the base, irregularly nodulose and sorediate towards the ends; soredia mostly granular, but a few elongated patches of greenish-yellow farinose soredia often present also: structure as for the typical but medulla more loosely woven.

Habitat. On mosses in alpine localities.

Distribution. Canterbury: Cass (W. R. Philipson, β11) Canterbury University Bot. Dept.; Mistake Peak, 5,000ft, Sc 69; Cass River, 4,000ft, Sc 55, Sc 1070; Tekapo (H. H. Allan V 181) CHR (pr. p.); Ben Ohau Range, 5,000ft, Mason, 130 (pr. p.); 6,000ft, Mason, 154 (Type). Southland: Ben Bolt, 1,500ft, 0339 (doubtful).

The variety is superficially similar to Antarctic forms of Xanthoria candelaria, from which it is distinguished by the fibrils and the different structure of the cortex, although the lower cortex in a few specimens approaches a plectenchymatous appearance. The Southland specimen is very small and has no soredia, and possibly belongs to some other species.

Teloschistes flavicans (Sw) Norm var. flavicans

Lichen flavicans Sw., Nov. Gen. Spec. Plant., 147 (1788).

Teloschistes flavicans Norm., Nyt. Mag. f. Naturvid, 7, 229 (1853)

Physcia flavicans Hook; Handb. N.Z. Fl., 572 (1867).

Teloschistes flavicans f. glaber Wain. Cranw. & Moore, Rec. Auck. Mus., l, 314 (1935).

Zahlbr; Lich; NZ; 123 (1941)

Thallus pale yellow, forming loose clumps 2–10 cm dia., 4–10 cm high, of terete or slightly angled matt or minutely tomentose laciniae 0.1–0.4 mm dia., dichotomously branched with nodes 1–5 mm apart, with scattered 1 mm long black-tipped fibrils and a few pale sorediate patches; cortex 80μ thick, of longitudinal hyphae with some vertical hyphae extending to form a thin tomentum; algae scattered under cortex, 8–12μ dia.; medulla hardly evident and lobes mostly hollow. (Apothecia 2–4 mm dia., lateral, sessile, plane to convex, yellow to brown, hymenium 70–100μ high, spores 12–18 × 6–12μ. Adapted from Hillmann. 1930).

Habitat. On trees or rarely mossy rocks.

Distribution. Cosmopolitan in warm climates. North Island: Hen Island (L. M. Cranwell, ZA 17) CHR (several collections).

The specimens seen by Zahlbruckner have black-tipped fibrils, whereas f. glaber is described as having all yellow fibrils. The New Zealand specimens thus correspond to var. intermedius Müll. Arg., an African form of the species, but except for this feature they are very like North American specimens.

Teloschistes flavicans var. compressus Murray, var. nov.

Evernia flavicans Hook., Fl. N.Z., II, 269 (1855).

(?) Physcia flavicans Linds., Trans. Linn. Soc., 25, 519 (1866).

A varietate typicale differt laciniis brevioribus, glabris, pro majore parte compressus, ad 0.8 mm latis et supra major minusve plants, solum ramulis ultimis teretibus, fibrillis concoloribus dispersis.

Thallus forming an orange-red mat to 20 cm dia. and 1–1½ cm high, of several times di- and trichotomously branched laciniae: laciniae mostly flat and dorsiventral or weakly

caniculate, up to 0.8 mm wide and 0.15 mm thick, final branches terete to angular, 0.1–0.15 mm dia., with scattered concolorous or rarely black-tipped fibrils and granular sorediate patches on margins and upper surface, cortex about 50μ thick of conglutinate horizontal hyphae without protruding hyphae, algae scattered under cortex, Trebouxia, 10–14μ dia.; medulla thin and obsolete or arachnoid and up to 80μ thick, lobes not hollow, sterile.

Habitat. Uncertain, apparently on subalpine rocks or logs.

Distribution. Australia, New Zealand. North Island: Waiariki (near Wellington) (Colenso 2170) WELT; Colonial Knob (Wellington) (H. H. Allan) CHR.

The identification of these specimens with the Australian variety is not certain, although they match the meagre description very well. The Australian plant is apparently known only from the type specimen, and this was not found by Hillmann (1930). According to Hillmann var. croceus differs in being a larger plant with nearly terete lobes, and var. exilis in being usually grey-green and forming much smaller clumps than Colenso's plant. The plant doubtfully identified as T. flavicans by Lindsay was from a subalpine situation, and may also belong here. The variety looks intermediate between var. flavicans and T. fasciculatus.

Teloschistes spinosus (Hook. f. & Tayl.) Murray, comb nov.

Parmelia spinosa Hook. f. & Tayl., Lond. J. Bot, 3,644 (1844).

Teloschistes sieberianus Hillm, Hedwigia, 69, 315 (1930) (pr. p.).

Thallus yellow to orange-yellow, initially squamulose on twigs, finally forming small cushions to 3 cm dia. and 2–5 mm high of more or less stellate-radiate laciniae 1–3 (-5) mm long × 0.3–0.8 mm wide and 0.1–0.15 mm thick, sparingly branched, flat to canicullate above, loosely adnate to substrate or ascending or erect, with scattered 1-½ mm long concolorous marginal fibrils and a few whitish rhizines on lower surface; esorediate, upper cortex fibrous, 20–50μ thick of 4μ dia. hyphae, algal layer irregular, with Trebouxia cells 10–18μ dia.; medulla loosely woven of 3μ dia. thin-walled hyphae, lower cortex up to 50μ thick, between fibrous and plectenchymatous. Apothecia at first sessile on upper surface, eventually appearing stipitate with 2–4 mm long grooved stalks (due to the thallus thickening and becoming erect), 3–5 mm dia., plane or slightly convex and sinuate, margin thin and disappearing, disc slightly darker than thallus, matt; hypothecium obscurely cellular, 40–45μ high; hymenium 65μ high, hyaline except for orange granular epithecium, paraphyses conglutinate, simple or furcate, 1½μ thick, clavate at tip; asci 40 × 13μ, thickened in upper part; spores broadly ellipsoid, 13–15 × 6–7½μ, r=0.4, pycnidia semi-immersed near margins of lobes, globose, 150μ dia., orange, pycnidiospores cylindrical (?) pseudoseptate, 3 × 1μ.

Habitat. On debris.

Distribution. Eastern Australia. Westland: Toaroha River, 3,000ft, Sc. 159.

Canterbury: Temuka, Mason, 256.

The species may be very local, as it has not been found in several other likely places in the South Island. The Temuka specimen is only a very small scrap, and could be an aberrant specimen of the following species, but the other collection of several plants agree exactly with Hillmann's description; the plant has much the apperance of a small delicate T. chrysophthalmus.

Teloschistes velifer Wilson.

Teloschistes velifer Wils., Proc. Roy. Soc. Tasmania 1892, 176 (1893).

(?) Physcia parietina var. lychnea Linds., Trans. Linn. Soc., 25, 518 (1866).

Xanthoria parietina var. spinulosa Mull. Arg. Bull. Herb. Boissier, II, app. 1, 40.

(1894) (not of Krempelhubei, 1868).

J. Linn. Soc. Bot. 32, 203 (1896).

(?) Teloschistes chrysophthalmus var. fornicatus Müll Arg., Bull. Herb. Boissier, 4, 89 (1896).

Zahlbr., Lich. N.Z., 123 (1941).

Xanthoria spinulosa Hillm., Ann. Mycol., 10 (1922).

Zahlbr., Lich. N.Z. 122 (1941).

(?) Xanthoria parietina f. rutilans. Zahlbi. Lich. N.Z. 122 (1941).

Thallus ascending, erect to subfruticose, orange-yellow to orange-red, varying from individual plants 3 mm dia. and 2–3 mm high to small cushions up to 3 cm dia. and 1 cm high. Juvenile lobes yellow, stellate-radiate, more or less linear ascending at ends, flat (0.2-) 0.5 (-0.8) mm wide, di- or trichotomously divided white and smooth beneath with

yellow marginal (or rarely ventral) fibrils 0 5–1.5 mm long; mature lobes orange-red, erect, weakly bullate, hollow, 2–5 (-8) mm long with ends expanded to 2–3 mm wide, hooded and open-ended to expose the algal layer as greenish soredia, with few or no basal fibrils; upper cortex 15–35 (-50)μ thick, uneven, of more or less longitudinally arranged 1μ dia. conglutinate hyphae; algal layer discontinuous with scattered clumps of Trebouxia cells up to 15×13μ, medulla arachnoid, of 3μ dia hyphae, absent near hooded ends of lobes; lower cortex about 40μ thick, fibrous or hyphae forming a loose few called plectenchyma. Apothecia very rare, one per frond, 1–2 mm dia, at first broadly sessile on upper surface with prominent inrolled thalline margin, finally appearing terminal on 1–2 mm high striate pedicels and then moderately convex with excluded margin, disc orange-red, matt; hypothecium 5–15μ high, hyaline, not well delimited; hymenium 55–60μ high, paraphyses simple or furcate, 2μ dia., asci clavate, thickened in upper part, 50×12–14μ; spores ellipsoid with narrow ends, 10–13 x5½-6½μ, r=0.3–0.5; pycnidia not seen.

Habitat. On twigs, bark, wood, rocks, rarely dry soil.

Distribution. Tasmania, New Zealand. North Island: Waiotapu, Allison, 305 and 334; Hutt Valley (H. H. Allan) CHR; Marlborough: Molesworth, CHR. Westland: Kumara, Mr., 1277; Greymouth (Mackay, W50) CHR. Canterbury: Waipara (A. J. Healy, 58/491) CHR, Hurunui (A. J. Healy, 58/502) CHR; Cheviot (A. J. Healy) CHR; Castle Hill (Z 33) CHR; Godley Valley, 2,500ft, Sc 223 and 224, (H. H. Allan) CHR; Esk River, 2,300ft, Sc 42 (plants unusually large and finely laciniate); Lake Tekapo, Mason, 248 and 250: Otago. Lake Ohau (R. Jones) 1722; Pigroot, T 1421; Shag Valley, T 1422. (and as H4 in CHR under T. chrysophthalmus var. fornicatus); Patearoa T 1378 (=H2 in CHR as T. chrys var. fornicatus); Lamb Hill, T 263 (and in CHR as X. parietina var. rutilans; Mihiwaka 1,000ft, Mr., 481; Waikouaiti, T 1849, Dunedin, T 898 (=H3 in CHR); Mr., 1129, 1223, Abbot's Hill, 1,200ft, 1605; Taieri, T 124 (=ZA 55 in CHR), T 1007 (=H61 in CHR); Maungatua, Mr., 1153; Taieri Mouth, 1294, 1372, 1446, 1699; Conical Hills (G. B. Rawlings) CHR. Southland. Forest Hill, 027, 033; Invercargill, Mr., 6917; Waihopai, Mr., 5415, Mr., 5417; Stewart Island Table Hill, Mr., 770.

There is evident confusion over the correct name for this very characteristic species. Zahlbruckner, in his Cat Lichenum Universalis, seems to have followed Du Rietz in identifying the New Zealand plant with Parmelia spinosa Hook. f. & Tayl., but Hooker himself later stated (1855, p. 287) that their species was identical with P. sieberiana Laur (i. e., Teloschistes sieberianus). Assuming this to be correct (compare Hillmann, 1922, 1930), Krempelhuber's is the first valid name for the species. Zahlbruckner in his Lich. N.Z. incorrectly cited the species as Xanthoria spinulosa (Hook. f. & Tayl.) D. Rietz, although the reference specimens in CHR are labelled X. spinosa Du Rietz. Apparently when Hillmann monographed the Xanthoriae he did not find Krempelhuber's material and depended on a specimen identified by Müller Argau. Although Hillmanns, description of Müller Argau's specimen confirms that it belongs to the present species, it does not clearly agree with Krempelhuber's account, which could equally apply to T. fasciculatus and perhaps other species.

I have not seen Müller Argau's description of T. chrysophthalmus var. fornicatus, but in any case this name would be predated by Teloschistes velifer Wils. (1892). Wilson's description of Tasmanian material is unmistakeable, although he gives details of the apothecia despite the statement that the specimens were sterile! The near absence of the species from older New Zealand collections is surprising, but may be due to its apparent rarity in the North Island, it is quite common in Otago and Southland. Du Rietz suggested that this species might belong in Teloschistes., although he had apparently seen no specimens, and in fact the structure of the cortex clearly relates it to this genus. Juvenile specimens of velifer, sieberianus fasciculatus and even chrysophithalmus are not easily separated. Fruiting specimens

for the whole of the North Island thermal region.Much of its area is occupied by the crater, near which no vegetation can survive owing to frequent ground movements and dense sulphurous fumes. The corrosive nature of the atmosphere is demonstrated by the marked decomposition of the rock surfaces and the equipment left in the abandoned fertiliser works.

Oliver (1915) suggested that the chief factor determining the peculiarities of the vegetation was the poisonous steam charged with HCl fumes which rose from the surface of the crater lake. This lake disappeared in the eruption of 1914, actually before his paper appeared in print, but there are still plenty of active steam vents.

Fleming (1948) remarked that probably no part of the island was quite free from the effect of the permeation of acid gases.Such vegetation as survives is also subjected to a constant rain of fine volcanic dust which showers from leaves and branches at the slightest touch.

The plants persisting under these condition are few and, like seabird vegetation, are predominantly halophytic although ascending 5–600 feet above sea level.In most cases their leaves are protected from salt spray, wind desiccation, guano, corrosive fumes and dust alike by a thick epidermis (e.g., Coprosma repens, Disphyma australe) a woolly tomentum (e.g., Metrosideros excelsa, which is exceptionally resistant in other areas to the guano of shags nesting in its branches) or scale-like hairs (e.g., Chenopodium allanii = C triandrum). These 4 species and the notably guano-tolerant Poa anceps v. condensata were those of chief importance, and only 4 others were recorded—viz., a few clumps of Phormium tenax on eastern clifftops, patches of Histiopteris incisa in dry shady creeks beneath trees in the west, a few clumps of the man-induced weed Phytolacca octandra in the vicinity of the fertiliser workers' old camp at Ohaurora and a single plant of Sonchus oleraceus in a gull-frequented part of one of the gannet colonies.

Oliver (1915) recorded a total of 12 species, his list including Metrosideros robusta (habitat unstated), Deyeuxia forsteri and Gnaphalium luteo-album (“rare, shingle beaches”) and Solanum nigrum (“near gannets”). Of the 9 species recorded in 1957, only Phytolacca did not appear in his list.

No vegetation was recorded on Captain Cook's chart, but in 1863 Captain Thomas reported that the island was covered on the N.and W.sides with a scrubby growth of timber and peopled by thousands of gannets and mutton birds. In 1924 “ there was at least 100 acres of pohutukawa forest capped by a cone of rose pink rock and broken up by white patches covered by thousands of gannets and seagulls” (White I. Agri.Chem. Co. Ltd. Bulletin, 1924). The “seagulls” may actually have been terns, as the only nesting gulls seen in 1957 were about 30 pairs of red-bills among driftwood in Crater Bay.Oliver (1915) makes no mention of either gulls or terns, only gannets and mutton birds.

Pohutukawa (Metrosideros excelsa) was still the dominant vegetation in 1957, forming a double, horsehoe-shaped belt around the lower part of the cone; broken where the crater extended to the E coast at Crater Bay and Wilson's Bay.

The lower part of the belt consisted of more mature forest, made up of trees 3–8 m.high. This was a fairly even-aged stand forming a closed community with practically no undergrowth and terminating abruptly at its upper margin. The upper part of the belt was again an even-aged stand but very open and of individuals only ½–¾ m.high protruding from loose rubble at 2–4 m. intervals.

Contrary to appearance these young pohutukawas were not the pioneers of a primary sere but of a secondary sere, having become established on ground which had previously borne plants. The surface of the intervening pink and grey “boulders” could be chipped off to reveal that the rock-like appearance was only

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on David Rocks there was little evidence that they were killing back the marginal bush and spreading on to the vacated ground as gannets and shags do.

White fronted terns were seen to be quite abundant on the Stewart Island waters of south New Zealand in February, 1957, but no nesting colonies were visited.Large gatherings of birds congregated on rocks beneath the Half Moon Bay lighthouse and on Ulva Island and Goat Island in Paterson Inlet and the following plants, none of which were found in the north New Zealand tern colonies, were recorded:—

The absence of plants such as Disphyma, Salicornia, Senecio and Spergularia was largely due to the paucity of blown sea spray in the sheltered Stewart Island habitats; the absence of many of the other northern species was due to latitudinal differences between the flora of the “sub-antarctic” south and the “sub-tropical” north.

A nesting colony of the slightly larger crested tern (Sterna bergii cristata (Stephens, 1826)) on a reef off Little Dog Island, in the Eastern Bass Strait, Tasmania, approximately 1,550 miles W.S.W.of the Hauraki Gulf colonies was, like them, dominated by Disphyma australe. Almost as common was Tetragonia implexicoma^* and Salicornia australis was quite frequent. These three species were the only members of the tern flora, although two others occurred on a part of the reef unaflected by the birds.

The reef was visited towards the end of the nesting season (22.1.58) when the chicks were almost fully grown and the plants very worn, a considerable number having been killed. The quantity of freshly dead stems in even the foulest areas indicated either that the vegetation regenerated seasonally from portions which had survived the breeding season in the partial protection of crevices or that the reef had been recently colonised by the birds (Terns had not previously been recorded here by C.S.I.R.O.ornithologists doing intensive studies in the area during the previous 12 years.)

“Disphyma australe was an important constituent of three other crested tern colonies in the Furneaux Group, Bass Strait (Scott's Reefs, Tuck's Reef and Nett's Reef.)”

A much larger colony of crested terns nested on the N. coast of Cat Island on the Tasman Sea fringe of the E.Bass Strait Group in 1954–5, and its effects on the vegetation were still very apparent three years later, in February, 1958.

There was no Disphyma, but the rocks previously occupied by terns were dominated by the related Carpobrotus rossii (= Mesembryanthemum aequilaterale) (28% ground cover), adjacent unoccupied rocks by Poa poaeformis (=Poa billardieri) (22%) and Pelargonium australe (16%) with Carpobrotus occupying only 1%.

On the deeper soil of the more inland part of the colony Bromus unioloides (= B.catharticus) had invaded the old nest site to give a 30% cover.This species.

when viewed from a distance, the difference being accentuated by the higher proportion of glaucous Chenopodium near the nests and the scarcity of flowers which might be attributable to the upsetting of the carbon: nitrogen ratio.

There was plenty of room for further expansion free from tree cover, and it seemed that the position of some of the nests varied from year to year, some previously occupied sites remaining empty and allowing plants not completely killed to regenerate. This ensured a more uniform appearance in all but the oldest established parts of the colony, but the ground seemed not to remain free from gannets for long enough to permit the entry of shrubs.

(b) On Gannet Point, the largest colony, there was also room for expansion, the Metrosideros excelsa forest being well back from the coast and much of the Coprosma repens belt being replaced by a luxuriant growth of Poa anceps. Some of the bare patches were steeply sloping, heavily burrowed by mutton birds and subject to erosion, but the presence of burrows alone was insufficient to deter the gannets and quite a few pairs were found sitting in the hollows formed by burrow entrances. In some cases the burrows had been effectively blocked by the accumulated nest material, and it is unlikely in any case that they could have remained occupied. This was most commonly seen on the Poa of the inland margin of the most westerly section of the gannetry where the birds were extending their range towards the trees at the expense of the mutton bird rookery.

Naked and downy chicks and eggs were present, but many pairs were still building, plucking Disphyma, Poa and Chenopodium from around the nest and so hastening the degradation of the indigenous flora.(In the Rocky Point woodland colonies, where little or none of this material was available on the site, the nests were of seaweed, the species having been identified by Miss Moore (Moore & Wodzicki, 1950). Some birds were building on Disphyma-covered c.1 m. in diameter but many preferred natural hollows.

The plants were surviving well, although photosynthesis must have been severely curtailed by the coating of guano. Disphyma and Poa were flowering, Chenopodium was in the vegetative condition, dark green and very large leaved.

(c) On Rocky Point the gannet colonies were much nearer sea level than were the other two more open colonies and had eaten back into the Metrosideros bush in three places.Usually the coastal fringe of Coprosma was tolerant of more guano as well as more exposure than was Metrosideros, though it still protected the taller trees behind from wind-borne sea spray. Neither approached the gannets much closer than 2 m. and a fringe of dead wood separated the living parts of the trees from the birds. The death of the branches only on the side of the tree where the roots were subjected to guano suggests that there may be little lateral translocation of solutes from the excreta. These trees were more stunted than those suffering greater exposure but no guano deposition.

The low cliffs below the gannets as well as the bush behind were tunnelled by mutton birds which might conceivably have occupied the entire area had they not been in competition with the larger birds. The Disphyma, Poa and Chenopodium occupying the seaward mutton bird rookery were creeping up into vacated parts of the gannetry.

(d) General “Gannet Succession.” Disphyma was the pioneer, advancing closest to the gannets and trailing up over the marginal, wind-trimmed Coprosma for distances of 2 m. and to heights of 1½ m. It did not, however, penetrate far beneath their shade Chenopodium and Poa pioneered the areas beneath the dead branches and Poa penetrated furthest beneath the living bushes where light was sufficient. Coprosma seedlings were scattered in these herb mats.

The part of the colony on the cape proper is the oldest and is thought to have been established about 1850. In 1946–47 nearly 3,000 nests were counted there, but 600 of these were unoccupied. A hundred years of occupation destroyed the vegetation on most of the cape, and the only nearby survivors seen in 1957 were plants of Disphyma australe over the brink of the vertical northern cliffs.

It is only at the junction of cape and plateau that birds and plants come into close proximity and this end of the gannet colony has been populated only since 1920, a large patch on the S. side of the W. slope even more recently (1932). The marginal vegetation may thus be regarded as having been influenced by gannets for approximately 30 years.

Figure 2
Plant Zonation in Relation to Proximity of Gannets Cape Kidnappers plateau colony
8. 1. 57.

the nests of the larger southern group had been re-occupied by birds, so it was not possible to see whether there had been any tentative colonisation by Poa during the winter there.

In British bird colonies some of the Poa annua which invades in winter, survives through the summer, but in the drier, warmer Hawke's Bay climate it apparently dies seasonally, and it formed a large proportion of the peripheral belt of dead grass recorded in January, 1957 (Fig. 2).

Lolium perenne is common in the “medium fertility zone” in Britain (where not grazed out by rabbits) and Agrostis tenuis, Bromus mollis, Vulpia bromoides, etc., on the poorer ground beyond (Gillham, 1955 a, b, 1956). Most of the other ornithocoprophilous Kidnapper's species, Urtica urens, Hordeum murinum, Geranium molle, etc., are also found in British seabird colonies.

The winter plant zonation adjacent to the Kidnappers plateau colony is shown diagrammatically in Fig. 3.

Figure 3
Relationship of Gannets and the Dominant Vegetation
Cape Kidnappers plateau colony. 8. 9. 57.

Another resemblance of the two similar habitats on opposite sides of the world is in the algal flora, Prasiola, much of it in the Hormidium stage, increasing in both British and New Zealand nesting grounds after the birds depart in autumn. P. crispa has been found more commonly in Britain, P. stipitata in New Zealand, and specimens collected by Mr. Beauglehole from the Lawrence Rock Gannetry off the south coast of Victoria proved to be the first record of P. stipitata for Australia. The author has since had Prasiola specimens determined from bird islands off North Tasmania and S.E. Victoria and on Macquarie I. in the sub-Antarctic.

An analysis of the macroscopic summer vegetation bordering the Kidnappers plateau colony is compared with that 20 m. away in Table III.

Table IV shows that species growing in the vicinity of both the cape and plateau gannets approach nearer to the latter by an average of just over 2 m. This is due.

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of all the more remote gannetries visited; Poa anceps, the other, of the White Island gannetries.

Species recorded within 15 m. of the gannets but not closer than 5 m. gave a narrower native: alien ratio of 1:1.75. Species of the cliff and plateau vegetation not found within 15 m. of the gannets gave a native: alien ratio of 1. 2:1.

The ratios of annuals to perennials in these three zones were 4.2:1, 1.75:1 and 1.2 respectively (Fig. 4..)

Figure 4
Diagram Showing the Increase of Alien and Annual Plants in Vegetation Affected by Birds
Cape Kidnappers gannet colony January 1957

These two sequences illustrate how short-lived alien species, the majority of them weeds of cultivation or pasture, are stimulated where birds disturb and enrich the soil, even when the birds are not themselves bringing seed as gulls do. Conditions suitable to plant growth may be restricted to as little as 4–5 winter months each year, sufficient only for the maturation of ephemeral plants, and the perennials which are killed out by the birds have little opportunity to become re-established.

The boundary of the gannetry acts in effect as a nursery for the weed species, ensuring an adequate supply of seed from which migration back to the pasture could take place in the event of the deterioration of the sward due to over-grazing or drought.

The mesophytic Kidnapper's perennials compete less adequately with the annual invaders than do the halo-coprophilous perennials of the island gannetries. It seems possible that the lack of guano-tolerant halopytes is due, not only to the less exposed situation of the mainland habitat, but also to the incidence of grazing. The only succulent halophytes recorded, Disphyma australe, Spergularia media and

The 1849 and 1851 specimens are usually said to have come from Resolution Island and Secretary Island respectively (e. g., Buller, 1888 and Oliver, 1955). As the species is flightless and the distance to be covered by swimming considerable, this seems most unlikely, and Mantell says the first specimen was “caught in the gully of a sound behind Resolution Island” (my italics). Hector (1863: 460) says he was told by one of the captors of the second specimen that it came from Deas Cove, on the mainland, and R. Henry (in Dollimore unpubl; p. 81–2) gives a verbatim account of its taking as well as that of two others (one from near Resolution Island) whose remains have apparently not been preserved. Only one complete adult specimen has been found since 1898. It was found dead in Takahe Valley in 1958.

Though the takahe first came to European attention through Mantell's discovery, it had been known to the Maori well before this in the southern part of the South Island for according to tradition, it was long hunted in parts of Fiordland (Beattie, 1949). In 1949 tradition was confirmed when a midden was found in Takahe Valley which contained takahe bones as well as those of other birds, including a species of bush moa (Duff, 1956). Radiocarbon dating of some of the associated material sets it in a period the middle of which is 1720 A. D. ± 60 years (Fergusson & Rafter, 1957). There seems to be no Maori legend explicitly referring to takahe in other parts of New Zealand, though subfossil and midden remains have been recorded from places mainly in the southern part of the North Island (Owen, 1872; Oliver, 1955; Greenway, 1958; Yaldwin, 1956; Falla, pers. comm.), and the eastern part of the South Island (Gurr, 1952). Reliable records for both islands are shown in Fig. 1.^* As some of the costal sites are middens they are only approximate guides to true distribution, for food was often carried for long distances.

Classification. On the North Island bones Owen founded Notornis mantelli. Meyer (1883) considered the skeleton from the 1879 specimen sufficiently different from the North Island bones to merit the founding of a new species, N. hochstetteri. But Forbes (1923) made public Parker's suggestion that the differences did not justify so great a separation. This opinion is in accord with modern ideas on taxonomy and has been endorsed by Peters (1934) and the New Zealand Checklist Committee (Fleming, et al. 1953), though Oliver remains staunch to the older view. Mayr (in Fleming, 1950) has suggested that Notornis and Porphyrio are congeneric, and Greenway accepts this opinion. In describing a new species of ectoparasite from the takahe, Rallicola takahe, Holloway (1955) remarks upon its close resemblance to the feather louse R. lugens, from pukeko, and suggests that this resemblance indicates that “N. mantelli and P. porphyrio melanotus are more closely related than at present indicated in the literature.” Although this view has much to commend it, the genus has been retained in the present paper. Notornis, then, is a genus endemic to New Zealand comprising two subspecies: N. m. mantelli, in the North Island, now extinct, and N. m. hochstetteri, in the South.

General Description, Weights, Sex Differences, Moults, Etc. Oliver (1955) and Buller give a detailed description of the adult N. m. hochstetteri. The following general account will thus suffice: The plumage has a loose and silky texture, that of the head, neck, breast, abdomen and thighs being indigo where exposed, but charcoal brown otherwise. Scapulars are peacock blue, changing to a metallic sage-green on the mantle Back, rump and upper tail coverts are olivegreen and the under tail coverts white. The primaries are indigo on the distal vanes but charcoal brown proximally. A striking contrast to the plumage is provided by the beak and legs—the former scarlet at the base and on the large frontal shield

Fig. No. 1

but becoming wax-pink distally; the legs and feet bright red. Eyes are reddish-brown. Though the species is flightless the wings are of a moderate size (length 243 mm, range 221–260 mm), and there is a spur at the carpal flexure. Colour plates appear in Buller—of these that in the earlier edition is the better—and in Rothschild (1907) and Oliver (1955). The last is a photograph of a living bird, but the reproduction of colour is poor. Rothschild's plate is the best in this respect.

Apart from its brighter and more varied plumage, the takahe is similar in general appearance to the pukeko, though much heavier (2–3 times), more robustly built and with shorter legs. Fleming (1951, 1958) has compared the general body structure of the adults of the two species particularly in regard to the differences in physiology, structure and function that arise through that of weight. There is no obvious sexual difference in the plumage and the bird stands about 20 inches (50 cm) high.

Fig. No. 3.—Scatter diagram showing sexual dimorphism of adults of N. m. hochstetteri.

Thirty-six living adults have been weighed and measured during the investigation and it has been assumed that—as in other of the Rallidae— the larger in a pair of birds is the male. By using either culmen measurements or weight the sexes apparently can be separated fairly accurately (Williams & Miers, 1958a) with but three exceptions, assumed males have culmen lengths greater than 87 mm and, with four exceptions, maximum weights greater than 2 · 60 kg. A scatter diagram

of culmen lengths and maximum weights shows only one male falling entirely within the assumed-female range (see Fig. 3). However, as this bird was mated with an even smaller one of culmen length 82 mm and weight 2 · 10 kg the assumption that it is a male seems justified. (The 1898 specimen—a sexed female (Benham, 1899a)—has a culmen length of 82 mm, and recently the bird found dead in 1958 was sexed by dissection and found to be a female, as presumed). The not very numerous records available of seasonal weight variations indicate for both sexes, over the time for which there are records, that weight is at a minimum about December, when the birds are busiest with nests and young (see Table No. 2). As the four assumed males with weights less than 2 · 60 kg have been recorded only at this time, this may be the reason for their lightness. In the absence of any other simple method for sexing takahe in the field, the present one seems satisfactory— that is, any yearling or adult whose weight is not greater than 2·60 kg, and whose culmen length is not in excess of 86 mm is a female, all others are males. It has been tested by seeing if such changes of mate as are known to have occurred involve the right sexes. No contradictions have yet appeared. A method for sexing the pukeko also dependent on culmen and weight measurements was checked by dissection and found to be 93.7% reliable (Williams & Miers, 1958b).

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Weights taken to nearest 0.05 kg; lengths to nearest mm.

The plumage sequences, now being studied in captive birds (Williams & Welch unpubl.) may be summarized as follows: The nestling is closely covered in black fur-like down except in the position of the future frontal shield where it is sparse. The bill is white from the external nares to the tip but black elsewhere. Legs are dark horn with a purplish tinge. At about one month down begins to be replaced by juvenile feathers to the tips of which it clings, and at about three months this moult is complete giving the juvenile plumage. Though similar to the adult, it is duller on head, neck, breast, mantle and back. The bill and developing frontal shield are almost black except for a paler tip to the upper mandible and the legs and feet are horn-coloured. A partial moult into a first winter plumage soon follows, the new feathers being mainly confined to the back and mantle. Though brighter than the juvenile they still lack the richness of those of the adult Legs and bill are now beginning to turn red, the former uniformly, the latter at the base and frontal shield only, elsewhere it is bluish-horn. The first summer plumage is still slightly browner on back and mantle than the adult, and the vanes of the distal primaries are greenish-indigo instead of the clear indigo of the adult, and although the rest of the bill has become pink there is still a bluish cast that is quite distinct. Yearlings and adults undergo a general post-nuptial moult which begins about mid-January and lasts throughout February and March. At this time takahe are quiet and secretive and large numbers of feathers are found under the shelter of tussocks, bushes or rocks where the birds have rested for long spells and aided the progress of the moult by plucking out feathers themselves.

Approximate weight at hatching is known from an embryo which died a few hours beforehand and was 61 grams. Twin chicks less than a week old weighed 96 grams and another set judged less than a fortnight old weighed 103 and 131 grams respectively. In captivity, three survivors of these four chicks reached the final stages of their body-weight growth curves at approximately seven months; wild young are in the adult weight range before they are twelve months old.

Internal anatomy has not recently been discussed. The skeleton of the nineteenth century specimens received considerable attention and detailed accounts with comparisons with related species may be found in Owen (1848b), Meyer (1833), and Parker (1882, 1886). Benham (1899b & c) studied the viscera— mainly the alimentary canal, larynx and syrinx. A recent observation of interest has been the great size and muscular development of the gizzard in both chick and adult. Benham, too, remarked on this in the adult he examined. He also published a paper (1899a) on the skin of the 1898 specimen. Recently Verheyen (1957) has given a general account of the characteristic morphology of the sub-family Porphyriinae to which Notornis, together with Porphyrio and Porphyrula, belongs.

N. m. mantelli is known only from bones. Those examined so far (Parker 1886, Yaldwin) suggest, surprisingly, that this race was larger than that from the colder South Island, but a more critical examination is necessary to take account of age and sex classes.

Since the above was written I have been able to compare the bones of the recent South Island adult female referred to in this paper with the skeletal material of extant and extinct forms held in New Zealand museums and with the full-scale plates of the Waingongoro bones described in Owen's original papers. There can now be no doubt whatsoever that the North Island subspecies was considerably larger than that from the South Island (as seemed fanly likely in any case from Oliver, 1955, and Falla in Greenway). However, leg bones from what is possibly

Fig. No. 1.— The Murchison Mountains area west of Lake. To Anau — the present range of
N. m. hochstetteri.

a single mature bird from the Martinborough Cave, North Island, fall within the size range of the South Island race and are smaller than any other North Island material. (The bones are held jointly by the Canterbury Museum, Christchurch, and the Dominion Museum, Wellington). The simplest explanation is that these belonged to an abnormally small adult N. m. mantelli.

Present Range. This includes the Murchison Mountains which lie between the Middle and South Fiords of Lake Te Anau, and part of the Kepler Mountains situated between the South Fiord of Lake Te Anau and Lake Manapouri (see Fig. 4). Though there have been occasional reports of takahe being seen elsewhere none has yet been confirmed.

Habitat. Most of the range is rough and mountainous. Peaks rise steeply to between five and six thousand feet and ridges and divides are kept narrow and jagged by the action of frost and snow. Streams are numerous and swift-flowing, there are many lakes and tarns and, in areas of poor drainage bogs. Rainfall is probably at least 100 inches a year with a sharply-decreasing gradient of intensity eastward and there is a heavy winter snowfall (Turbott, 1951). Below the treeline the dominant vegetation is evergreen beech forest, mainly Nothofagus menziesii (silver beech) and N. cliffortioides (mountain beech) with the degree of dominance of each varying from place to place. Above the forest is a band of subalipine scrub made up mainly of Dracophyllum uniflorum, Hebe buxifolia, and Olearia moschata. Above this again the tall tussocks are dominant—Danthonia flavescens and D. teretifolia on well-drained slopes and D. crassiuscula and D. oreophila elsewhere (Baylis, 1956, unpubl.). Rarely, but importantly as far as the takahe is concerned, tall tussocks are dominant well below the tree-line. When this is so the species most commonly occurring is D. rigida.

Takahe are generally to be found at about 3,300 to 3,800 feet where the forest has given way to scrub and snow tussocks—mainly D. flavescens. Occasionally tussock and scrub tongues run down through the forest to lower altitudes and takahe may then be found on these tongues. In winter and early spring, when snow lies thick on the tussock-covered mountain tops, the birds move into the forest until a thaw occurs.

Takahe Valley, where the species was rediscovered and where numbers are far more concentrated than elsewhere, is unusual in that the three-mile-long valley floor is fairly flat, wide and treeless (see Fig. 5); and although about five hundred feet below the tree-line (which here lies between 3,400 and 3,500 feet) it is, for the most part, covered in a luxuriant growth of red tussock, Danthonia rigida, with Festuca novaezcalandiae and Poa colensoi prominent among the grasses. In boggy places sedges (especially Cares spp.) are common. On each side of the valley floor forest-covered slopes rise steeply to the subalpine scrub and the tall-tussock meadows above. A stream collects much of the drainage of the upper valley and of the cuque at the head and meanders through occasional patches of bog to empty into a lake about 1,200 yards long and 250 yards wide situated at the lower end of the valley. The lake is drained by a stream which discharges through a gap in some limestone bluffs beneath which the takahe often shelter in the winter, and after a series of water-falls and caverns it finally reaches Lake Te Anau nearly 2,500 feet below.

The Point Burn, the area of next highest takahe concentration, lies immediately to the south and is separated from Takahe Valley for the most part by a low forest-covered ridge that rises between 300 and 400 feet. The valley of the Point Burn is typical of those in the Murchison Mountains and the rest of Fiordland in that it is narrow with its floor covered in beech forest except for an occasional clearing where mainly red tussock occurs. The lowest clearing in which takahe nest in this.

valley is at an altitude of about 2,400 feet and this is the lowest altitude at which the birds are known to continuously live and breed.

There is little doubt that distribution is correlated with that of the two species of tall tussock, D. rigida and D. flavescens, upon which the birds mainly depend for food, cover and nesting sites, and that the relatively dense population in the Takahe Valley—Point Burn area exists there because of the unusually favourable habitat. Baylis has made the general comment that “the wet Frordland mountains are botanically the least favourable part of the South Island for survival of the takahe since this is the region in which the large Danthonias are most extensively replaced at comparatively low altitudes by smaller species such as D. crassiuscula and D. teretifolia”. This then poses the problem of why takahe now exist only in this area and not also in apparently favourable ones elsewhere where the preferred species of Danthonia do occur.

Changes in Distribution and Possible Causes. Notwithstanding a tentative identification (Phillipps, 1959) of the legendary North Island “moho” or “mohoau” with the takahe, there is no clear reference to the species in Maori legends of the North Island and in those of the northern part of the South Island. This implies that it has long been rare or extinct there, and the fact that only subfossil or very old midden remains have been found supports this view. None of these have been accurately dated, but the midden material at Wairau Bar and Lake Grassmere has been assigned by Dufl to the Moa-hunter culture of 950–1550 A. D. and the rest is unlikely to be any more recent. Bones found at Pyramid Valley are associated with those of the large moas which became extinct before 1450 A. D., according to Duff. Inland deposits further south, for example, those at Earnscleugh and Castle Rock, show a similar association. Some, at least, of the Otago and Southland records are from middens—the one in Takahe Valley being dated, as we have seen, at approximately 1720 A. D. In none of the deposits so far critically examined have Notornis remains been abundant which implies that the species has not been common during the thousand or so years man has been in New Zealand. Only in western Otago are there reliable records of sightings in European and pre-European times, though Gurr (1952) accepts records given by Park (1888) implying a notable further shrinkage in range in this area within the last century. It is clear from present knowledge that Park's claims to have heard takahe are wrong. The acceptable records are Maori traditions which tell of hunting the birds in the mountains between Lakes Monowar are Te Anau, the evidence, gathered over the last eleven years, of the species' distribution in the Murchison and Kepler Ranges and the five specimens found before 1900. A rough parallelogram including Dusky Sound, Doubtful Sound, the Middle Frord of Lake Te Anau and the junction of the Whitestone and Mararoa Rivers delimits this recent range.

So early a scarcity of remains coupled with a major shrinkage in distribution exonerates all those suggested causes resulting from European settlement, such as habitat destruction, predation by—or competition with—introduced animals or epidemics from newly-introduced diseases. Any part played by the early Polynesians in hunting the birds or unfavourably altering then habitat has probably only accelerated a natural trend towards extinction. In this regard it should be remembered that the present population has survived in spite of being hunted until at least the beginning of the European era; and that even the presence during the last half century or so of possibly inimical exotic mammals such as stoats, red deer and Australian opossums does not seem to have had a detrimental effect. This may not be so in the future if the numbers of any of these mammals should greatly increase. Competition for food with any surviving native species is highly improbable because of the takahe's very specialized feeding habits. Whether any competion

earlier age I have seen a chick taking Viola filicaulis from a parent's bill. When the chick is being fed, one parent picks up the food and passes it on to the chick or accepts that collected by the other parent and then passes it on. Pukekos and wekas, Gallirallus australis, behave similarly. Chicks ingest grit at an early age—I have found quartz grains in the droppings of a bird less than a fortnight old.

Kean (1956) has corapared the availability of foods in two Takahe Valley territories, using results from an analysis of droppings to support a contention that food resources of a territory are reflected in the degree of success attained in raising chicks. However, his analysis is only of plant foods and covers only one season.

Droppings are a sure sign of active occupation of an area. When fresh they are most commonly bright green (the colour of the vegetation itself), cylindrical and about as long as the little finger. If not soon destroyed by heavy rain, they dry and bleach and finally become almost white and hay-like. In sheltered places they may lie for years. Kean has described the changes in appearance that occur in the open with time and these are often useful for roughly dating events and for tracing birds Because chick droppings are of a different size they may be used for establishing one's presence when no other evidence is available.

In captivity, takahe have been found to be very dependent on water, even in cold weather. Not only do they drink and bathe very frequently, but artificial foods such as lettuce and fruits are carried to water and immersed before eating. This degree of dependence—hitherto unsuspected of being so great—perhaps explains why takahe do not occupy some parts of their main range previously thought to be suitable, for in a dry spell these places become temporarily devoid of water.

Calls. The “contact” call, used by birds of a pair that have become separated or in territorial dispute at a distance, is an almost monosyllabic “klowp” uttered on a rising inflection. It resembles the disyllabic but rather more flute-like call of the weka, common in the same range. The two have been confused in the past because takahe will frequently answer wekas against whom they are very aggressive during the breeding season. Fleming's (1951) description of a pair of takahe calling in a “two-part canon” is an example of this confusion, for though this kind of stimulus and response occurs with takahe, the transliteration of the call as “cooeet” is more accurately descriptive of that of the weka. Occasionally the contact call is heard late at night which implies some nocturnal activity.

The alarm note is percussive, described in Maori tradition as sounding like the knocking together of pieces of greenstone underwater, but nowadays less picturesquely transliterated as “oomp”. It is heard when takahe are aware of the presence of an observer, and also serves as a warning to chicks to remain under cover. Again, the weka has a similar call but more staccato and continuous and given with a greater frequency A quieter version of the alarm note is used between birds of a pair when close together and undisturbed, and this, with a soft rhythmical “kau” superimposed, serves as a recall note for chicks.

A variety of clucking sounds, rather like those of the common owl, are made by undisturbed adults when feeding; and, rarely, if suddenly alarmed at close quarters, takahe utter a loud screech.

Chicks' pipings have little that is characteristic about them, but immatures or yearlings readily call, when alarmed, in a distintive way—rather like the distress call of the pukeko. Even in the absence of any other evidence, this call may be taken as proof that a young bird is present Three chicks taken from Takahe Valley when only a few days old developed the “contact” call at about 4½ months, and the alarm note at about 7 months of age.

Display. Thick cover makes observation difficult in the wild, and little has been recorded. The following brief notes are in part based upon observations of captive birds

(I) Alarm. When a bird is uneasy the tail is continuously and rhythmically flicked upwards to expose the white under-tail coverts. In the presence of other takahe this no doubt acts as a warning signal. This display, characteristic of the Rallidae, is recorded for almost all species. Once, an alarmed adult, being stalked, took to the water, dived, and kept itself totally submerged by grasping underwater vegetation with its feet. It remained thus for five to ten seconds until it was picked up and placed on dry land. (See Witherby et al., 1941, for similar behaviour of the moorhen, Gallinula chloropus).

(II) Extreme Threat. In territorial dispute, either just before attack or interspersed with fighting, the wings are held high and arched, the neck feathers ruffed out and the wing coverts made prominent. If attack ensues blows are struck with beak and neck. An account by Moon (1957) of an intimidatory display by an incubating pukeko resembles this closely. Gullion (1952), under the name “paired display”, describes similar behaviour in the American coot, Fulica americana.

(III) Mutual Display. Falla (1951) describes such between birds of an incubating pan thus: “The sitting bird was called off by its mate. The bird responding darted with a crouching run to the caller and straightened up facing it with the two bill tips almost touching and both necks upstretched. After some seconds one bird, I think the original caller, crouched and moved round the other with a gyratory movement which presented the spread white under-tail coverts to the other's view. Drooped wings and fluffed-out flank feathers gave the general impression of a round white target ringed with blue” A very similar display, with mutual necknibbling, occurs in mated birds that have not yet bred and is probably a preliminary to coitus. Gullions's “arching” display seems to be an equivalent in the American coot.

(IV) “Gaping” Occasionally birds will continually raise their heads and open their bills wide as if calling, but no sound emerges, nor does it appear that nearby birds are responding. This behaviour may go on for some time. It may be elicited after disturbance from sleep, which suggests it a form of threat; on the other hand, it is quite commonly seen when the birds are bathing. I have not seen this behaviour described for other rails.

Pair Bonds. Pairs once formed probably last until the death of one of the partners. Fig. 6 gives details of the duration of pair bonds and of changes that have occurred. In four of the five changes there has been no subsequent record of the superseded bird, though it could have escaped observation or, even less likely (as will be seen presently) have moved out of the study area altogether. In the fifth change the first mate was unbanded and so could not be identified later.

Territorial Behaviour. The obvious year-round occupation by a pair of a fairly well-defined home range or territory^* is an aspect of behaviour that has often been commented on (Falla, Williams, 1952; Williams and Miers, 1958a). Even in the depth of winter when snow lies deep on the valley floor, most of the birds are still in at least part of their summer range though, as they keep mainly to the shelter of the forest at this season, they are not easily seen. However, whenever an extensive thaw occurs they immediately move out into the exposed tussocks. Other rails are known to hold territories throughout the year, for Cramp (1947) has described such behaviour for the European coot, Fulica atra and Gullion (1953) for the American coot.

Various evidence shows that mated pairs or family groups of takahe are attached at least semi-permanently to an area: The characteristic droppings do not occur randomly distributed over the whole of country occupied by the species in the main colonies, but are concentrated (fresh and old alike) in well-marked localities

Fig. No. 7.—Line hatching shows approximate extent and boundaries of the “permanent” takahe territories in Takahe Valley. The cross-hatching on the N. E. shore of L. Orbell indicates an occasional extension of the territory of the pair on the northern bluffs. Stippled areas show the other occupied parts of the main colony which are not well known because of the difficult terrain.

that are usually separated by expanses where fresh droppings are scarce. Feeding signs, equally characteristic, show the same distribution. This is not just a reflexion of the distribution of preferred food plants and water, for these are usually widely and abundantly spread and often occur in places where there is little occupation. Furthermore, calling and sight records are most frequent in the vicinity of those places where they have been reported in other seasons and years. Nests, too, are made in well-used localities season after season, even though sites similar in every way are widespread. All these criteria for establishing territorial behaviour show excellent agreement; but it is the sighting or recapture of banded birds that supplies the most reliable data for fixing the approximate boundaries and period of occupation. Williams and Miers (1958a) have discussed the subject in detail and have included a map, reproduced in Fig. 7, which shows the location and approximate boundaries of a number of permanently-occupied areas. The size of each remains fairly constant—the smallest being about 5 acres, the largest about 20; though these are only provisional estimates.

If it is assumed that a bird has been in continous occupation provided that it has never been seen settled elsewhere and no alien birds have been seen living in its territory, then we have the following occupation records up to the end of the 1958–59 breeding season (see Fig. 8). Seven seasons in a territory—1 bird (No. 16378), six seasons—1 (No. 16377), five seasons—3 (Nos. 16380, 16384, 18488), four seasons—4 (Nos. 16385, 16389, 16390, 16391), three seasons—6 (Nos 16382, 16386, 16392, 16393, 16394, 18492), and two seasons—10 (Nos. 2, 3, 8, 9, 16376, 16379, 16383, 16396, 16397, 16398, 16399)

If pair bonds and territories are semi-permanent young birds, unless they oust one of their parents, must generally move over considerable distances in search of a

Fig. No. 8.—Minimum survival times of all takahe banded up to the end of 1958. No. 16378
was found dead in November, 1958. All were banded as adults or yearlings except the following, which were banded as chicks. Nos. 16376, 16380, 16383, 16387, 16397 and 18489.

place to establish themselves, and there is evidence from banding records that this is so. For example, No. 16380, banded as a chick, was found a year later established in a territory about 2,000 yards away. This is the greatest distance a takahe is known to have moved over. Occasionally there are movements well beyond territorial boundaries that do not seem to be connected with wandering of young or displaced birds but rather to have arisen from disputes, the temporary straying of a chick or from disturbace through capture. Two temporary shifts of over 1,000 yards are known to have been made by adults shortly after being banded, and these

involved the crossing of adjoining occupied areas. The movements of one of these birds, No. 16388, is shown in Fig. 9. An undisturbed pair not seen beyond its territory in five years was found to have left it unoccupied, while others in the main colony were still in theirs and to have moved fully three-quarters of a mile away from the main colony altogether. One of the birds of this pair is No. 16380 in Fig. 9. Movements of two other birds in Takahe Valley are also shown in Fig. 9. They are typical of the wanderings that are the general rule in that they extend little, if at all, beyond recognizable territorial boundaries.

Method of Obtaining Data. As a means of studying behaviour in this species banding has not been entirely satisfactory. The birds spend most of their life in thick cover much taller than themselves and are wary and difficult to shepherd into open ground where the colour combinations may be seen. Loss and fading of the plastic colour bands must also be contended with, though both of these difficulties have been largely overcome since nitrocellulose lacquer has been used to seal the bands and colours subject to fading have been withdrawn from use. All the 20 uncemented bands put on between late 1952 and early 1954 appear to have now been lost, but only six of the 27 cemented bands are known to have suffered the same fate since. There is no evidence that any numbered aluminium bands have been lost.

The nature of the cover and terrain in which the birds live and their behaviour make the use of any sort of trap for their capture impracticable If capture is necessary it must be done by hand. This is difficult, for the birds can run fast and have the advantage of concealment. Three men are needed for efficiency and a minimum of disturbance.

Field work consists of a daily traverse on foot of five to ten miles of forest, bog and mountain meadow, with frequent climbs and descents of 1,000 feet or more. Heavy rain or snowfall limit the amount of observation that can be made on any

Fig. No. 9 — Movements of four selected takahe. Digits indicate first and last observed positions and therefore the total number of observations. Further details in text. The three areas enclosed by heavy lines are those that have been censused.

Twenty-four eggs have been measured. Length, 68–78 mm, mean, 73·5 mm breadth, 47·5–52·5 mm, mean, 49·0 mm. They are pale buff and marked irregularly with blotches of brown and mauve.

Takahe occasionally re-nest: apart from the three-egg nest described above, three other re-nestings have been recorded; two rely on circumstantial evidence—an occupied nest has later been found deserted with a new and occupied one nearby in the same territory. In one of these egg measurements from the two nests were in excellent agreement. For the third record there is unequivocal evidence an indentified banded adult was seen with a newly-hatched chick some ten weeks after having been seen in almost precisely the same place with another. Though the first was not found again its death cannot be assumed, for birds of about three months of age are particularly difficult to see in the tall tussock and low scrub.

Incubation. Though the incubation period is not known exactly, two independent estimates agree in fixing it at close to four weeks: (1) A nest was found in which incubation had been going on for about 10 days, judging from the size of the latrine, which is a reasonable guide. Eighteen days later one of the two eggs was very close to pipping, for the chick was moving about vigorously inside and could be heard “peeping” from a yard off; the other chick was alive but silent and not so far advanced Unfortunately, this was the last visit that could be made.

(2) In 1957 four eggs were taken (two 2-egg clutches) in the first attempt to raise the species in captivity. These eggs were very fresh and, judging from candling and from the size of the nest latiines, could have been incubated for only about two days before they were removed. Under a bantam one reached a stage of development so advanced that the chick could be heard calling from within. This took 26 more days. The chick failed to pip the shell and died, but was found to have been within a few hours of hatching.

As is usual among the Rallidae, incubation is shared by the sexes. Eggs are not covered when the nest is left, and this, too, is usual ralline behaviour. Secker (1952. unpubl.) on one occasion heard what he believed to be egg-turning to continue for 10 minutes after one bird had been relieved by another. On taking its place on the nest a bird will sometimes rearrange the cover within reach to improve concealment, and will often gradually turn its red bill from sight when approached by an observer. Both sexes sometimes sit on empty nests as if incubating eggs, one pair for as long as seven to eight weeks, another for at least a month judging from the size of the latrine and the state of the droppings. Two more pairs have “incubated” on nests that have never held eggs. Observations of two captive pairs suggest that at least one of the birds in each of these four wild pairs may have been a yearling (see above, under “Breeding Age”).

Egg Fertility. Of the 67 eggs recorded, the status or ultimate fate of 48 is known and the data available are summarised in Table 3.

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ones), or were destroyed by the takahe themselves before fertility could be investigated. This gives a likelihood of hatching of 57% and a mean productivity of 1·0 chick per laying pair. Three clutches were totally unsuccessful. A truer estimate will take into account those pairs known not to have laid. However, to include all four would give undue weight to the frequency with which non-laying occurs in the nest records as a whole, which is once in every twelve. By including one of these non-laying pairs in the calculation proportionate representation is restored. Thus, 13 pairs hatch 12 chicks, or 0·9 chick per pair. This should be a maximum estimate of chick production, for a higher proportion (75%) of two-egg clutches is included in the sample than occurs in the nest records as a whole (52%), and all three known sets of hatched twins are included also.

A minimum estimate of chick production may be obtained, and one calculation of this has been given in an earlier publication (Williams, 1957). A new estimate may now be made from all 48 nests 21 recently-hatched chicks have been observed. Forty-two pairs produced these (48, less four known to have re-nested, less another two whose eggs were taken in the 1957–58 season for the first attempt to raise the species in captivity). One pair, then, produces 0·5 chick surviving to leave the nest. This is a minimum figure because, as already pointed out, some chicks have certainly not been accounted for—either because they have not been seen or because of their death soon after hatching.

Chick productivity in this species therefore lies between the limits of 0·5 and 0·9 per pair, or 0·25 and 0·45 per adult.

An approximate check on this estimate may be made by using the provisional figure for hatching success of 57%. Forty-two pairs have laid a minimum of 67 eggs. If 57% of these hatched, 38 chicks would result—a productivity of 0·9 chick per pair. At present it is not possible to decide whether 57% is likely to be above or below the true figure, but the agreement of this estimate of productivity with the upper of the two limits found by other means suggests that the true chick productivity is likely to lie closer to 0·9 than 0·5.

Juvenile and Adult Survival. Little is known of chick survival for few have been banded. Of the seven that have (the last in 1955), survival times for each up to the last sighting have been as follows: 1 week, 2 months, 11 months, 12 months, 13 months, 5 years and 3 months, and one not seen again. The banding and recapture programme begun late in 1952 (Riney and Miers, 1956) has furnished some preliminary data on adult survival. Fig. 8 summarises all returns and recoveries up to early 1959. Only one banded bird has been found dead and, as it was at least one year old when banded, it was at least seven years old at death. Only two other adults have been found dead in the main colony area over the last eleven years, both before banding was begun. The implication from this and the data in Fig. 8 is that takahe, having reached the yearling stage, have a good expectation of further life.

By assuming (1) that over the last 11 years there has been a stable population in the main colony (annual estimates of numbers support this), (II) that the breeding age is one year (there is some evidence for this), and (III) that adult mortality is independent of age (an assumption generally accepted as being true for birds), it is possible to make provisional calculations of expectation of further life for adults.

If all chicks leaving the nest survive to breed the replacement rate or annual adult mortality would lie between the limits of 25% and 45%. But there is sure to be considerable first-year mortality. If this were only 50% replacement rate would lie between 12·5% and 22·5%, which implies an average expectation of further life for adults of between 4 and 7·5 years If 67%, then annual adult mortality would lie between the limits of 8·3% and 15% with an average expectation of further life of between six and 10 years.

With 50% first-year mortality 2·2 to 4 pairs would be needed to produce a chick surviving to breeding age; with 67% mortality the necessary number would lie between 3·3 and 6.

A first-year mortality of 67% is by no means high for birds (Lack, 1954). Alley and Boyd found it to be that for the coot, and Gullion (1954) found mortality to be already 46% for the American coot even at fledging.

If takahe are as long-lived as these preliminary calculations indicate, it will be some years before a satisfactory life table can be constructed.

Population Size and Its Regulation. Various expeditions to the north, west and south of the Takahe Valley-Point Burn area have shown that beyond these two valleys the population density is much less—perhaps one or two birds to the square mile, though this is only an estimate. With a density of this order occurring over the known range the total population of the species can be only a few hundred.

Counting of the pairs in the main areas of the Takahe Valley-Point Burn colonies (shown enclosed by heavy lines in Fig. 9) has indicated that there is little annual variation, the number always lying between ten and twelve (Williams and Miers, 1958a). As territories are held throughout the year and often for a number of years, the observer's familiarity with them makes a direct census more trustworthy than it would be otherwise. However, unmated wandering adults are not satistorily accounted for, though long observation makes it certain that they are not numerous. The total adult population, then, in the main areas probably lies between 25 and 35. Another estimation that should take account of unmated adults may be made by using a Lincoln Index technique: The main areas were repeatedly and thoroughly traversed in the 1954–55 and 1955–56 breeding seasons and, from the ratio between birds seen with aluminium bands and those without, estimations made of the adult populations for each of these seasons gave figures of 29 and 33 respectively. The provisos attendant upon the use of a Lincoln Index to calculate population size are well-known, but, as with the direct census of mated pairs, familiarity with the terrain and the birds' habits removes a number of objections and makes the results of the calculations rather more trustworthy than they would otherwise be. For example, it is unlikely that any birds were counted twice or that banded ones were unavailable for census through being outside the census areas. Conversely, immigration of unmarked birds was unlikely also. As the species is known to be fairly long-lived once about a year old and as marking with metal bands was begun only in early 1954, loss of bands through mortality is likely to have been very small. Therefore, the agreement between the two population estimates is probably not fortuitous.

Two parts of the Takahe Valley-Point Burn area—the great cirques at the head of each valley—are difficult country and not so well known as the rest. However, the number of adults living here (the stippled areas in Fig. 9) is less than the combined total of those in the other parts of the main colonies. To sum up: observations over the last seven years clearly show that there has been little variation in the adult population and estimates would fix it as very close to 50 birds.

Territories or home ranges in the main colonies are almost constant in number and position from year to year and it is only very rarely that takahe succeed in establishing themselves on ground already occupied. Nor do new pairs appear in unoccupied ground and this compels one to believe that where such ground is not obviously unsuitable it is part of a defended area even though behaviour in defence of it has not been seen. Territorial behaviour is, therefore, regulating population density and, through it, population size in these colonies. What, then, determines the extent of the territories? Long familiarity with them and with the takahe's food habits and cover needs convince me that the area each pair occupies is far in excess of what is needed to supply adequate food and shelter throughout.

borne on the anterior part of its segment; variability in number and size of spines present is common; number ranges between 7 and 9, the mean of 10 antennae being 8.2; female never possesses spines.

Legs. Fore and middle legs subequal in length, with hind leg slightly less than twice length of fore or middle legs in female, and slightly more than twice length of fore or middle legs in male. Sexual dimorphism is shown by fore and middle legs of female being 0.6 as long as in the male and hind legs of female 0.5 as long as in the male. Fore and hind femora, fore, middle and hind tibiae and proximal segment of hind tarsi armed with variable number of linear spines (Table I). No spines occur on middle femora and fore or middle tarsi Apical spines constant in number, as in generic description. On hind tibiae prolateral apical dorsal spur longer than retrolateral one. Sexual dimorphism is shown in hind tarsi: those of adult male possessing a small, round, dorsal callosity between apical spines of second proximal tarsal segment, Fig. 2 (CA); this callosity is not present in tarsi of female. Length of proximal segment of hind tarsus subequal with other three together. Ratio of length of legs to length of body: Fore leg, male 2.9:1; female 2:1. Middle leg, male 2.8:1; female 1.9:1. Hind leg, male 5.9:1; female 3.6:1.

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(Figures in parentheses represent number of specimens.)

Genitalia. Female: Suranal plate, Fig. 3 (SAP), concave laterally, truncated distally; distal margin clothed with short setae. Subgenital plate. Fig. 4 (SGP), convex laterally, very deeply notched distally; sparsely clothed with setae. Male: Suranal plate. Fig. 5 (SPL), convex laterally, emarginate distally; plate thickly clothed with short setae. Subgenital plate. Fig. 6 (H), tringulate, subequal in width to length, sides spreading slightly proximally, tapering to concave distally with a rounded apex, glabrous on dorsal side, but apical protuberance on ventral surface thickly clothed with short setae. Two styli Figs 5. 6 (S), thickly clothed with short setae length of styli being 0.25 length of sternite IX (S IX), Subgenital

plate covers genitalia. Parameres, Figs. 6, 7 (P), attenuated, broad at base and tapering to a point, subequal in length to width, prolateral margin thickly clothed with long setae, rest of paramere clothed with short setae. Pseudosternite, Fig. 7 (PD), 1.3 wider than long, tapering to a point distally. Penis, Fig. 7 (PN), two-lobed, each lobe subequal in width to length. Paraprocts, Figs. 6, 7 (PP), elongate, 1.7 longer than broad.

Locality. Limestone caves, Waipu (type locality), coll. A. M. Richards, B. M. May.

Types. Holotype male, allotype female, and paratype male and female so designated in Plant Diseases Division collection.

Pachyrhamma waipuensis is most closely related to P. fusca Richards, but differs from it in:

• 1. Absence or marked reduction in number of dorsal linear spines on middle tibiae.

• 2. Presence on hind tarsi of adult male of a round dorsal callosity between the apical spines of the second proximal tarsal segment.

• 3. Shape of subgenital plate of female.

Genus Pallidoplectron Richards, 1958. Trans. Roy. Soc. N. Z., 85, p. 703.

Pallidoplectron peniculosum n.sp. Text-fig. 2, Figs. 1–5.

Colour. Basic colour light brown, anterior and posterior borders of pronotum and posterior borders of mesonotum, metanotum and abdominal terga light brown; lateral borders of pronotum and mesonotum light brown; nota and abdominal terga irregularly mottled with light brown and ochreous; femora and tibiae banded with light brown and ochreous; tarsi ochreous; antennae light brown; ovipositor deep ochreous, pale reddish-brown at tip and along edges of dorsal and ventral valves.

Body. Length, 15 mm in male, 16 mm in female. Ovipositor 0.7 times as long as body. Antennae in male 7 times as long as body, and in female 6 times as long as body. Fastigium twice as long as high, with base touching scape of antennae. Maxillary palps with third and fourth joints subequal in length. Pronotum and mesonotum distinctly margined laterally and posteriorly.

Antennae. As in generic description (Richards, 1958). Third segment narrower than pedicel, but subequal in length with it. All segments thickly clothed with short golden setae. Sexual dimorphism very poorly developed, antennae of male slightly longer than those of female; no spines present on flagellum of male or female.

Legs. Thickly clothed with short setae. Fore and middle legs subequal in length, with hind leg 1.7 length of fore or middle legs. Sexual dimorphism absent. All femore sulcate ventrally. Fore and middle femora unarmed; hind femora bearing 6 prolateral and 2 retrolateral linear spines beneath in male, and 6 prolateral and either 1 or 2 retrolateral linear spines beneath in female. Fore and middle tibiae bearing 2 prolateral and 2 retrolateral linear spines beneath; hind tibiae with 19 or 21 prolateral and 21 or 24 retrolateral linear spines above in male, and 23 or 24 prolateral and 23 or 27 retrolateral linear spines above in female Fore, middle and hind tarsi without linear spines. Apical spines constant in number, as for generic description. Length of proximal segment of hind tarsus subequal with other three together. Ratio of length of legs to length of body: fore leg, 1.6:1; middle leg, 1.6:1; hind leg 2.8:1.

Genitalia. Female: Suranal plate, Fig. 1 (SAP), slightly convex laterally, with distal margin truncated and bearing two groups of setae. Subgenital plate, Fig. 2 (SGP), straight laterally, distal margin bearing two tubercles; plate glabrous. Male: Suranal plate, Fig. 3 (SPL), concave laterally, deeply notched distally; latero-distal parts of plate thickly clothed with setae, rest of plate sparsely clothed with setae. Subgenital plate, Figs. 3, 4 (H), concave laterally, attenuated and rounded distally; the well-developed median keel thickly clothed with setae at distal end, rest of dorsal surface of plate sparsely clothed with setae; ventral surface with distal lobe thickly clothed with long setae, proximal to the median depression are two lobes fused together medianly and each bearing five small processes distally. Two styli. Figs. 3, 4 (S), thickly clothed with short setae, length of styli being 0.4 length of sternite IX (S IX). Subgenital plate covers genitalia. Parameres, Fig. 5 (P), small, broad at base, tapering to a point 1, 2 broader than long, thickly clothed with setae Pseudosternite, Fig. 5 (PD), compressed dorsoventrally, 1.5 broader than long; lateral margin straight, but notched medianly, distal margin trilobed, the two lateral lobes larger and extending beyond the median one. Penis, Fig. 5 (PN), two-lobed, each lobe nearly twice as broad as long. Paraprocts absent.