Transactions
of the
Royal Society of New Zealand

• Ranunculaceae

• Clematis paniculata J. F. Gmelin(syn. C. indivisa Willd.) — C

• — hexasepala DC. — O

• Ranunculus hirtus Banks & Sol. ex Forst. f. — A

• Monimiaceae

• Hedycarya arborea J. R. & G. Fors.t — A

• Laurelia novaezelandiae A. Cunn. — O

• Lauraceae

• Beilschmiedia tawa (A. Cunn.) Benth. & Hook. f. — O

• Litsaea calicaris Benth. & Hook. f. — A

• Cruciferae

• Cardamine heterophylla (Forst. f.)O. E. Schulz — O

• Droseraceae

• Drosera auriculata Backh. — O

• Saxifragaceae

• Carpodetus serratus J. R. & G. Forst. — O

• Pittosporaceae

• Pittosporum tenuifolium Banks &Sol. ex Gaertn. — R

• — cornifolium A. Cunn. — O

• Cunoniaceae

• Weinmannia racemosa Linn. f. — O

• Rosaceae

• Rubus squarrosus Fritsch. — O

• — schmidelioides A. Cunn. — R

• Acaena anserinifolia (J. R & G. Forst.) Druce (syn. A. sanguisorbae Vahl. — O)

• Leguminosae

• Carmichaelia cunninghami Raoul (syn. C. australis A. Cunn.) — O

• Sophora microphylla Ait. — C

• Oxalidaceae

• Oxalis corniculata Linn. — O

• Rutaceae

• Melicope ternata Forst. — O

• Meliaceae

• Dysoxylum spectabile Hook. f. — A

• Coriariaceae

• Coriaria arborea Lindsay — O

• Corynocarpaceae

• Corynocarpus laevigata Forst. — O

• Sapindaceae

• Alectryon excelsum Gaertn. — C

• Dodonaea viscosa Jacq. — O

• Rhamnaceae

• Pomaderris phylicaefolia Lodd. — C

• Elaeocarpaceae

• Elaeocarpus dentatus Vahl. — O

• Aristotelia serrata (J. R. & G. Forst.) W. R. B. Oliver — O

• Tiliaceae

• Entelea arborescens R. Br. — O

• Malvaceae

• Hoheria populnea A. Cunn. — O

• Violaceae

• Melicytus ramiflorus J. R. & G. Forst. — C

• Passifloraceae

• Tetrapathaea tetrandra (Banks &Sol.) Cheesem. — O

• Myrtaceae

• Leptospermum scoparium J. R. & G. Forst. — A

• — ericoides A. Rich. — A

• Metrosideros scandens (J. R. & G. Forst.) Druce — O

• — carminea W. R. B. Oliver — C

• — diffusa (Forst. f.) W. R. B. Oliver — C

• — robusta A. Cunn. — C

• — perforata (J. R & G. Forst.) Rich. — C

• Myrtus bullata Sol. ex A. Cunn. — O

• Eugenia maire A. Cunn. — O

• Onagraceae

• Epilobium sp. — O

• Fuchsia excorticata Linn. f. — O

• Haloragidaceae

• Haloragis erecta (Murr.) Schindler — C

• — procumbens Cheesem. — A

• Araliaceae

• Nothopanax arboreum (Linn. f.) Seem. — A

• Schefflera digitata J. R. & G. Forst. — O

• Pseudopanax crassifolium (Sol.) C. Koch — A

Approximately 300 bronze and Stewart Island shags roosted on a corner of Tamihau Island in Paterson Inlet, their numbers having steadily increased over the past 40–50 years according to the local wild-life ranger. They were spreading gradually inland from the coastal rocks to the marginal scrub, and most of the older Olearia colensoi trees which dominated the scrub had been killed as a result of the guano at their roots. In a few only the lower branches had been killed and a sparse leafy crown persisted 4–5 m above the ground. (Plate 15, a.)

Saplings of Olearia occurred on the inner margin of the shag roost, many of them dead, others with bare stems 1 m or more high topped by a depauperate cluster of leaves 15–30 cm across. The scarcity of lower branches may have been hastened by the shags breaking some of them off, as tree-nesting shags break off twigs about the nest to obtain a clear view (Oliver, 1930).

Wherever a skin of peaty soil overlay the rocks of the roosting area it was occupied by an 80% cover of Tillaea moschata. This was heavily fouled with guano and much of it uprooted, but shoots were able to remain turgid if only one rootlet remained unsevered. The foliage was deep green, due presumably to the high nitrogen content, as it was a characteristic rusty red elsewhere.

Atriplex hastata was the most abundant subordinate species, the plants of the roost, all of them damaged to some extent, being as follows.

The only important species of coastal rocks apparently unable to withstand the trampling and guano deposition was Selliera radicans

The ground flora of the coastal bush adjacent to the roost appeared to be more resistant to disturbance by shags than was the dominant Olearia colensoi, and there was a rich growth dominated by Hebe elliptica 60 cm high under the half dead trees. Hebe was probably the most guano-tolerant shrub encountered in the S of New Zealand as well as one of the most salt-tolerant. Species of this marginal belt were as follows:—

No nesting colonies of blue shags (Stictocarbo p. steadi, the Stewart Island sub-species of the grey or spotted shag) were visited, but the birds were frequent on coastal rocks of Goat Island, Tamihau Island, Tommy's Island and Ulva Island in Paterson Inlet. They were seldom present in large numbers and appeared to have little effect on the rock flora.

2 Otago

A colony of bronze and Stewart Island shags occurred on the western slopes of Green Island, near Dunedin. The ground was saturated with guano, dry and very spongy, giving way underfoot, and no macroscopic plants survived. The remains of dead Hebe bushes were scattered through the colony, suggesting that the area had previously been part of the Hebe elliptica/H elliptica × salicifolia scrab which occupied the main part of the island. (Plate 15b.)

Plants approaching closest to the shags were seedlings of Atriplex hastata and the fringe of vegetation on the landward side consisted of the two species of Hebe, Lavatera arborea and Solanum laciniatum.

A colony of several hundred bronze and Stewart Island shags occurred on Taiaroa Head at the end of the Otago Peninsula, but it was not possible to make a landing there, so no floristic investigation was carried out. The neighbouring grass, bracken and shrubs had been killed out for 8–10 m back from the normal.

seaward margin of vegetation, and few if any angiosperms persisted in the rookery. (Plate 16, a.)

Grey or spotted shags (Stictocarbo punctatus punctatus. or Phalacrocorax punctatus) also occurred on Taiaroa Head, sometimes nesting on Acaena anserinifolia (A. sanguisorbae) which died within two weeks of the shags' arrival according to the local wild-life ranger.

A small colony of grey shags nested on a low stack off Portobello, in Otago Harbour. Some of the rookery area consisted of bare rock, some was covered by crisply erect, thalloid Prasiola sp. forming an algal mat 1 cm or more thick and some by a mat of Disphyma australe.

Hebe elliptica occurred to either side, completely dead where most exposed to guano, taller and only partially dead where slightly more remote from the birds. The native Poa astonii occurred in the rookery together with three of the annual aliens so common in bird colonies of both Europe and Australasia (Poa annua, Hordeum murinum and Coronopus didymus).

The native Senecio lautus and alien Sonchus oleraceus, two composites of widely dispersed seabird colonies in New Zealand and S. E. Australia, the Sonchus also in Britain, occurred a little further from the nests. Close behind these, and again characteristic of seabird colonies beyond New Zealand were Apium prostratum, Solanum laciniatum and the British Holcus lanatus and Sagina procumbens agg.

3 Cook Strait

No nesting colonies of the king shag (Phalacrocorax carunculatus carunculatus) were visited, but birds were observed on the Disphyma australe, Salicornia australis, Senecio lautus, Coprosma repens communities of the chain of islands between Komakohua Island and Long Island and also on the S. of Little Brother Island.

A group of grey shags were found roosting on a stack off the E. side of Motungarara Island beside Copiosma repens ½ m high. Poa caespitosa was colonising the soil of crevices not too fouled by guano.

4 Hauraki Gulf

Five grey shag areas were visited in the Hauraki Gulf, one pair (a nesting colony and a roost approximately 3½ miles apart) about 26 miles E. of Auckland, another nest colony/roosting area pair in the Noises Group c. 15 miles N. E. of Auckland, and a further colony on cliffs of Bush Island, off the W. coast of the Coromandel Peninsula.

In the first pair of habitats the shags nested on Tarakihi or Shag Island, to the N. E. of Pakatoa Island, and roosted on Ponui Island, to the S. E. of Waiheke Island. A “home roost” occurred on sloping rocks above the nesting colony on Tarakihi and several hundred birds were seen here, many taking off in the direction of the Ponui Island roost as the dinghy approached.

The nests were of sticks, mostly in crevices or under ledges on almost vertical cliffs where there was no vegetation, but patches of very fouled Disphyma australe and Parapholis incurva? (Lepturus or Pholiurus) occurred where groups of unoccupied birds sat. The sloping rocks of the roost above were white with guano and dotted with patches of Disphyma with stunted Coprosma repens and Metrosideros excelsa less common.

About 100 shags were seen on the Ponui Island roost before the general exodus in that direction from Tarakihi. Some of the birds were on the N. E. cliffs of the main island, others on a tidal stack forming part of an un-named reef to the N. of Scully's Reef. All repaired to an unvegetated offshore stack further S. when disturbed.

Disphyma australe was dominant in both the cliff and reef roosts, heavily coated with guano but surviving well. Five other halophytes, also much fouled, occurred

in the cliff roost, 1 halophyte and 7 non-halophytes in an adjacent area of similar size and aspect undisturbed by shags (Aliens in the subsequent lists are marked with an asterisk).

Disphyma formed a 75–80% cover on vegetated parts of the reef roost and with it three other halophytes, the annual Parapholis dead. On the seaward side of an adjacent stack unfrequented by shags the vegetation was more diverse.

The vegetation on the landward side of the shag reef was halophytic and resembled that on the seaward side;, that on the landward side of the unoccupied reef was less halophytic and more varied, including species from the grassland above.

The nests in the Noises Group of islands to the N. W. were in vertical niches of the cliffs of Otata Island and constructed mainly of seaweed. Disphyma was fairly frequent among the nests, Coprosma less so—both badly fouled but apparently thriving.

Sixty to seventy birds were observed in the roost one mile to the E. on the S. and E. sides of the most southerly of the stacks forming David Rocks, and a few on the E side of the N. E. stack.

In the main roost the ground was very foul and Disphyma the only surviving plant. As the boat approached the shags retreated up the cliff into the 1–2 m high Metrosideros excelsa scrub above. The soil there was saturated with guano, the ground vegetation had been eliminated and the Metrosideros was in poor condition with most of the lower branches killed back or broken off.

5 Summary of Vegetation in Colonies of Ground-nesting Shags

Where seabirds are ashore for part of the year only, as in tern, gannet and gull colonies, there is an influx of winter annuals, most of them ahens, as the guano becomes diluted after the close of the nesting season (Gillham, 1959a). Shags are ashore for longer, some species seldom going far from the home area, throughout the year, so their suppressive effects on the vegetation are more continuous and the winter influx of annuals less noticeable.

Table I summarises the vegetation of 10 of the more important shag areas visited and shows the number of native plants to be twice as great as the number

The soil was less fouled than was usual in shag colonies, the guano being insufficient to kill the underlying vegetation, but sufficient to induce a very rank growth of coprophiles. Desiccation of the soft nitrogen-induced foliage was hindered by the close shade of the Hebe canopy and Poa annua, averaging 30 cm high, was the dominant ground species. This limiting of the guano to beneficial rather than toxic concentration was quite probably the result of the eggs or young having been taken from the nests early in the breeding season by fishermen who regarded the species as a pest.

Six indigenous and eight non-indigenous plants occurred in the ground flora, the non-indigenous ones being much more robust and covering more ground.Tillaea moschata was again the most abundant indigenous species, and the Hebe was regenerating, small seedlings being scattered through the mat of introduced species. The high proportion of alien plants may well have been connected with the fact that scattered pairs of black-backed gulls (Larus dominicanus) nested on the edge of the scrub. Prasiola sp. was present, and the vascular plants were as follows, aliens marked with an asterisk.

Three pairs of white-throated shags nested in a Hebe elliptica bush 1 m high on the cliff edge below the colony of royal albatrosses (Diomedea epomophora) on Taiaroa Head in 1956 (Sharpe in lit.).

3. Hauraki Gulf

About 20 pairs of white throated shags and five pairs of little pied shags (Phalacrocorax melanoleucus) were found to be breeding on the innermost of three offshore stacks at the W. end of Motuihe Island with red-billed gulls and white-fronted terns (Sterna striata).

The nests were in dense Coprosma repens scrub 30–100 cm high, and the ground flora consisted of Disphyma australe and the alien annual grass Avena fatua.

4. Summary of Vegetation in Colonies of Scrub-nesting Shags

Most of the colonies of scrub-nesting shags investigated were in the S. of New Zealand and the shrub most favoured as a nesting site Hebe elliptica, its counterpart offering similar conditions in the N being Coprosma repens. Small specimens of Metrosideros lucida, a species also favoured by tree-nesting shags, were sometimes used as nesting sites in the S., the tendency being for these to die out and the shags to move to the Hebe understorey which in its turn died out causing the shags to move to other areas.

The ground vegetation was spared most of the trampling suffered by that in colonies of surface-nesting shags, and some of the guano was prevented from reaching it by the dense leaf canopy of the shrubs and by the nests themselves. The plant habitat was thus much less rigorous and fewer areas were denuded of vegetation altogether.

One of the commoner scrub-nesting species of shag, the white-throated, was characteristically associated with red-billed gulls which nested beneath the scrub to the detriment of the indigenous flora. Viable weed seeds ejected in pelletform by the gulls were able to germinate when the height of their nesting activity was over and the guano from the shags was insufficient to prevent this. There was, in fact, evidence that the shade provided by the nests and the diluted excreta

Pied shags nested in Metrosideros excelsa on a sheltered part of the coast S. of Coromandel.

5. Bay of Islands.

About 20 small black shags (Phalacrocorax sulcirostris) and a few pied shags were seen roosting in Metrosideros excelsa by a sheltered arm of Crowell's Bay in the Bay of Islands. There were no nests and, although many of the leaves of the roosting-trees were invisible beneath a coating of guano there was less killing of branches than in the average nesting colony and little plucking of twigs. The following ground species were recorded, none very badly damaged.

A fairly large nesting colony of pied shags and little pied shags with either whitethroated or small black shags occurred in more severely damaged Metrosideros excelsa trees in Manawaora Bay. When disturbed the birds flew inland to a much frequented hilltop perch on a partially dead Leptospermum scoparium tree. A diverse vegetation survived beneath the nesting trees, the species most severely “scorched” by the guano being Arthropodium, Microsorium, Olearia, Pteridium and Stellaria The following were recorded:-

6. Summary of Vegetation in Colonies of Tree-nesting Shags.

The nesting trees, Metrosideros lucida in the S, M. excelsa in the N. and Pinus radiata where the native bush had been destroyed, possessed xeromorphic leaves

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very resistant to damage by shag guano. Both leaves and twigs were often broken off, braches were killed back and sometimes the entire tree had died.

Trees used only for roosting had usually suffered less damage. The configuration of the branches was not as important as where suitable sites were required for nestbuilding and other species were utilised (e. g., Podocarpus ferrugineus and Leptospermum scoparium)

Coastal forest occurred only where there was a reasonable degree of shelter from wind-borne spray (Gillham, 1959b) and the vegetation as a whole was less halophytic. The nesting trees frequently overhung the water so that much of the excreta was deposited in the sea or on the Hormosira, etc., of the mtertidal zone.

Distribution of Native and Alien Plants in 28 Shag Colonies

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

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

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

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

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

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

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

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

Habitat. As for var. australe.

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Habitat. On trees, rarely mosses.

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

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

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

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

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

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

• Nephroma helveticum Ach. var. helveticum

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

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

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

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

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

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

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

Habitat. Tree trunks, rarely on damp rocks

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

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

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

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

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

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

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

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

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

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

Habitat Trees. damp rocks in shade.

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

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

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

Nephroma lyallii Bab.

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

• Nyl., Synops. Lich I, 322 (1860).

• Hook Handb N. Z. Fl. 566 (1867).

• Nyl., Lich N. Z. 42 (1888)

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

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

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

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

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

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

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

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

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

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

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

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

P. Caninâ

Peltigera nigripunctata Bitt.

Peltigera nigripunctata Bitt. Berichte Deutsch. Bot. Ges., 27, 194 (1909).

Peltigera nigripunctata Bitt. f. farinosa Gyeln., Annal Cryptog. exot. 4, 168 (1932), and Rev. Bryol. et Lichen., 5, 61, 69 (1932).

This is a Javan species which should be easily recognized by the characters given in the key. The form farinosa is based on a specimen collected by Berggren in 1874 in New Zealand (locality not stated) and now in the Uppsala Herbarium as “P. venosa, Berggren 39”. It is not mentioned by Hellborn (1896) who described Berggren's New Zealand lichens. Gyelnik's form “differt thallo superne partim mcuso” seems a trivial modification not worth retaining. P. nigripunctata apparently differs from P. venosa only in having cephalodia on the upper surface as well as the lower, but I have seen neither specimens nor a complete description.

Peltigera scutata (Dicks) Duby.

Peltigera scutata (Dicks) Duby, Bot Gallic II, 599 (1830).

Peltigera polydactyla var. scutata Müll. Arg., J. Linn. Soc. Bot 32, 201 (1896).

Thallus medium sized, curled at edges, grey to brown, slightly scabrid to smooth, commonly prumose at tips of lobes and with masses of grey soredia on the margins. Under surface without veins or with broad brownish veins and yellowish interspaces. Rhizines short and fasciculate. Apothecia up to 3 mm dia, horizontal or erect, hymemnium up to 200μ thick, spores 3–7 septate, 30–70 × 3–5½μ (Description abbreviated from Thomson, 1950).

Habitat. Logs, rocks.

Distribution. Eurasia, North America, Peru, Chile, New Zealand.

Exsiceata Seen. Lich. suecici (Du Rietz), Fl. suecica (Sandberg). The species is reported from Napier (Colenso, 1658) by Müller, but I have seen no New Zealand specimens, nor is it in the Colenso lichen collection in Wellington.

Peltigera virescens (Stnr.) Gyeln.

• Peltigera rufescens var. virescens Steiner apud Zahlbr. et Zederb., Annal. Naturhist. Hofmuseum Wien, 20, 372 (1907).

• Peltigera virescens Gyelnik, Rev. Bryol. et Lich, 5, 73 (1932).

• et apud Zahlbr. Lich. N. Z. 46 (1941).

• Peltigera degeni f. tasmaniae Gyel., Magy Bot. Lapok, 28, 61 (1929).

• ?=Peltigera virescens var. tasmaniae Gyelnik apud Zahlbruckner.

• Lich. N. Z., 46 (1941).

• Peltigera tereziana Gyelnik, Oesterr Bot. Zeitschr, 77, 220 (1928)

• ? Peltigera pellucida f. dilacerata Gyeln. apud Zahlbr. Lich. N. Z., 46 (1941).

Thallus 3–10 cm dia, lobes 10–25 mm long, 3–10 wide, grey-green, smooth, glabrous, veins forming an anastomosing network, raised and narrow, rhizines usually tapering, simple but with some fasciculate and short (less than 3 mm long); interspaces between veins thinly tomentose. Cortex 25μ, algal layer about 40μ, medulla 200–300μ of fairly loosely woven hyphae 7½μ dia. Apothecia vertical on extended lobules, mostly about 3–5 × 2–4 mm; hymenium 100μ thick, including pale brown epithecium, hypothecium brown, 50μ thick; asci 65–95 × 15μ, apparently 6-spored; spores very pale yellow, 3–5 septate, straight or arcuate, 50–95 × 2½–3μ.

Habitat. Logs, stones, moss in damp places.

Distribution. North America, Europe, Australasia. North Island: Tiritea (G2 Zotov, pr. p.) CHR; (Chamberlain, sub P. dolichorhiza) CHR. Marlborough: Onamaluta, Mr. 4214 (pr p.); Waihopai, Mr. 4211 (pr p). Canterbury: Godley

River, 4,000ft, Sc 285. Westland. Franz Josef (C. K. Boey), 4166. Otago: Haast Pass (R. F. Smith), 1182; Trotter's Gorge, T 1400; Flagstaff, 1,000ft, T 732 (and as G21 in CHR sub P. virescens var. tasmaniae); Dunedin, T 1395, T 1071, T 1072; Taieri Mouth, 1423, 1676.

Exsiccata Seen. Fl. Hung. (Filarszky, sub P. pellucida)

A specimen from Jamaica (Plitt) distributed as P. virescens from Gyelink's Herb Lich., has uniformly short fasciculate rhizines, and thus cannot be this species.

This species seems to be commonly known as P. degent Gyelnik, but the name virescens clearly has priority if the species are truly identical. Gyelnik's variety tasmaniae, according to his description and the specimen in CHR (G21) is merely a rather short lobed fruiting form, such as may be found in several species P. pellucida f dilacerata Gyelnik according to the description is identical with virescens, although Thomson (1950) lists it as a synonym of P. polydactyla. P. Lairdii described by Dodge and Rudolph (1955) from Macquarie Id is supposed to resemble P. dilacerata but is tomentose above, and must thus actually be in the P. Canina group I have not seen the original description of P. dilacerata which was founded on a specimen from Auckland P. Tereziana, founded on a specimen from Wellington, is a pusilloid form of virescens according to the mention in Gyelnik (1931, 1932), although it is listed as a synonym of P. canina var. spuria by Thomson (1950), and is said to resemble P. frigida according to Santesson(1944)

It is possible that P. virescens might be best regarded as a variety of dolichorhiza, since the character of the thickness of the veins is the only clear distinction in non-fruiting specimens, and sometimes the veins are in part like those of the latter species In one of my Otago specimens (1423) some of the lobes have traces of white tomentum at the edges, and this plant may belong in the canina group.

Peltigera horizontalis (Huds) Baumg. var. muscorum (Schl.) Schaer.

Peltigera muscorum Schleich (1823) ? in sched.

peltigera horizontalis var. muscorum Schaer, Lich Helvet Spicil, (5) 265 (1833)

Peltigera horizontalis Hellborn, Bihang Kgl. Svensk Vebensk Akad Handl, 21, III (13), 29 (1896)

Thallus flat, to 8 cm dia., yellowish-green or brownish-green, lobes 1½ cm long by ½ cm wide, glabrous, somewhat shining; cortex 40–60μ thick, outer cells 13 × 8μ, algal layer 50–110μ thick, algae mostly 8 × 6μ, medulla 140μ thick, of septate hyphae 4–10½μ dia., lower surface with broad, flat anastomosing veins, rhizines sparse, dark, fasciculate and short. Apothecia on broad horizontal lobules, 2–5 mm dia., distinctly raised above thalline margin; hypothecium reddish-brown, 10–30μ, hymenium 70–95μ thick, paraphyses 2½ thick asci 6–8-spored, spores uniformly 3-septate, hyaline to light brownish, 26–32 (-38) × 5½–8μ, sometimes slightly constricted at middle septum.

Habitat. On soil (in New Zealand)

Distribution. North Temperate Zone, New Zealand Marlborough. Avon Valley, Mr. 4245, Mr. 6918 Canterbury Waipara (Allan), Godley Valley, 2,400ft, Sc 203; 4,300ft, Sc 202; Lake Tekapo, Mason, 43 and 54 Otago Matukituki Valley (R. F. Smith) 0966, Lake Ohau, Mason, 177a.

Zahlbruckner in Cat Lich Univ and Thomson (1950) both reduce the varety to formal rank, the latter stating that it is variable and inconstant and occurs throughout the range of the species European forms of horzontalis which I have seen, however, are different from the New Zealand specimens, being much larger and thicker, with broad lobes (up to 3 cm) and considerably larger apothecia with distinctly longer spores (30–40μ) Consequently I have preferred to retain the variety Hellbom's plant from Porter's Pass evidently also belongs here Peltigera frigida. Santesson (1944) from South America seems very close to the New Zealand specimens of horizontalis var. muscorum, but apparently differs in having radiating veins, thinner cortex and longer, narrower spores.

Peltigera horizontalis var. muscorum f. albido-pruinosa Murray, f. nov.

A varietate differt thallo superne dense albido-pruinoso, planta minima.

Thallus of more or less separate lobes, 3–8 mm long by 2–4 mm wide, mostly densely white pruinose above but with some patches of smooth surface; with broad pale brownish veins below; cortex 75–100μ high, hyaline of plectenchyma with cells up to 25 × 15μ and with thin irregular decomposed outer layer. Apothecia 3–4 mm dia., dark brown, epruinose; hypothecium brown, 25μ thick, hymenium 95μ high with pale, thin epithecium, spores broadly fusiform, 3-septate, 32 × 8μ, hyaline.

Distribution Otago: Matukituki Valley, 4,000ft (D. Scott), 4389 (on soil among scree)

The plant looks very like Santesson's illustration of Peltigera frigida (1944), from which it differs particularly in the pruina and thick cortex. The collection is quite distinct from other specimens of P. horizontalis but may be only an extreme habitat form.

Peltigera polydactyla (Neck) Hoffm var. polydactyla.

• Lichen polydactylon Neck, Method Muscor 85 (1771)

• Peltigera polydactyla Hoffm, Descript et Adumbr Plant Lich, 19 (1790), Bab. in.

• Hook Fl. N. Z. 271 (1855)

• Hellb, Bihang Kgl. Svensk Vetensk Ada Handl 21, III (13) 28 (1896)

• Hook, Handb N. Z. Flora, 566 (1867)

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

• Kirk, Trans. N. Z. Inst 4, 235 (1871). Muller, J. Linn Soc. Bot, 32, 201 (1896)

• Peltigera polydactyla f minor Krmph, Reise Oesterr Fregatt Novara, Bot Vol II, 121 (1870).

? Peltidea polydactyla, Hook Fl Antarctica, l, 197 (1844)

Thallus up to 20 cm dia., commonly of more or less detached lobes 3–8 cm long by about 1½ cm wide, grey, grey-green or brownish, smooth and glabrous above sometimes with somewhat impressed or undulating surface (corresponding to veins beneath) and usually ascending margins Lower surface white to brown with veins ½–1½ mm wide and 02 mm high, carrying mostly short fasciculate rhizines or occasionally some simple ones Cortex 30–35μ of about 4 cell rows, algal layer (10-) 30 (-65)μ, medulla (150-) 250–450μ of loosely woven hyphae 8–15μ dia., with cell walls 2½μ thick Apothecia more or less round, 2–5 mm dia. with thin crenate margin, hymenium 100–150μ, including the pale brownish epithecium, hypothecium brown, 35–65 (-100)μ thick, asci 80–100 × 12–20μ, spores acicular (3-) 5–9 septate, light brownish, straight to slightly arcuate, 50–80 × 3–4μ

Habitat. In damp places on soil, logs, stones, etc.

Distribution. Cosmopolitan; North Island Orongaronga R. (Allan), CHR (pr p), Wellington (KG11) CHR Nelson. Hundalee, Mr. 1306. Marlborough: Pelorus Bridge, Mr. 4168. Canterbury Craigieburn Range, 5,500ft (A. F. Mark), 4163 Westland Greymouth, Mr. 5422, Styx River, 2,100ft, Scott, 141; Fox Glacier (J. M. Anderson), 0744 Otago: Dunedin, Mr. 5426; Mihiwaka, Mr. 706 (pr p), Mt. Cargill, Mr. 5423 (pr p), Mt. Flagstaff, 1,000ft, 1980; Taieri Mouth, 1261, 1464, 1666, 1667, 1672; Southland; Tautuky Bay, 1025 Stewart Island: Port Pegasus, Mr. 5424 (or dolichorhiza var. nana) Chatham Islands: (Colenso, 19) WELT.

Exsiccata Seen. Swed. Lich. (Magnusson, 13788), Boros Lich. Fl. exsicc austrohung No. 41.

P. polydactyla has been frequently reported from New Zealand, and is probably not uncommon, although it is evidently less so than the closely related P. dolichorhiza. Probably many of the early reports actually refer to the latter species. Like most wide-ranging species it is variable, and it has been noted (e.g., by Nylander, 1860) that the New Zealand specimens are distinctly smaller lobed and thinner than the European forms and have a more impressed surface when dry. On this account our plants were distinguished as f minor by Krempelhuber Although this was reduced to synonymy with f. microcarpa (Ach.) Merat by Thomson (1950), the forms are

evidently different. The latter differs from the typical form in the narrower (but not necessarily thinner) lobes and small (2 mm dia.) apothecia, whereas f minor has thinner lobes and small apothecia A few of the specimens listed above have rather thin thalli with impressed upper surfaces, and presumably belong in f minor Kremph. The dimensions of the cortex and algal layer are the same as in more typical specimens Small apothecia which appear on many specimens are immature and are not clearly associated with thinner thalli. I have not thought it worth separating these as a distinct form.

Some New Zealand specimens have lacerate margins in part, and thus correspond to f lophyra (Ach.) Nyl This seems a trivial modification hardly worthy of recognition, the condition is sometimes seen on parts of otherwise normal plants One specimen from Dunedm, Mr. 5426, has abundant 10 mm long black rhizines but is not otherwise different from normal.

There seems no doubt that P. polydactyla is endemic to New Zealand both because of early reports which describe it as common, and because our specimens are usually distinguishable from European or American forms by their generally thinner thalli and narrower lobes.

Peltigera polydactyla var. magyarica (Gyelnik) Murray comb. nov.

Peltzgera magyarzca Gyelnik, Ann. Musei. Nat Hung. Pars Bot., 31, 46 (1937)

This is similar to the species but is usually thicker than the typical form in New Zealand and has smaller, more or less cochleate lobes and few rhizmes below it is often fruiting freely.

Habitat. On clay banks.

Distribution Europe, New Zealand, North America North Island-Mangarakau, T 2630, Huia, Auckland University Botany Department Otago: Matukituki Valley, 1,700ft (D Scott et al) 4388, Dunedm, 1936; Taieii Mouth, 1385, 1502.

Distinguished by the above characters, this has the appearance of a habitat form, but two of the quoted specimens I found growing within a few centimetres of normal plants of P. polydactyla and dolichorhiza. It seems to be of rare occurrence and is perhaps not a good variety. It is not very different from small specimens of the following variety.

• Peltigera polydactyla var. polydactyloides (Nyl) Maas Gest. in sched?

• Peltigera polydactyloides Nyl., Flora, 46, 265 (1863)

• Peltigera crassotdes Gyeln, Magy Bot. Lapok 29, 51 (1930)

• Peltigera polydactyla var. hymrenzna Auct (non Ach.)

• ? Peltigera pusilla Zahlbr. Lich N. Z., 45 (1941)

Plants usually rather small with ascending or incuiled edges, underside with a mo e or less continuous tomentum and the veins absent or very indistinct (except sometimes cntrally) rhizines usually few but sometimes clustered m the centre, cortex 30μ, algal laye 25–50μ medulla 350–450μ of loosely woven hyphae 8–10μ dia. Apothecia sare, hymcnuin ca 100μ thick, hypothecium brown, 50μ thick; spores 7–9 septate, 60μ or more long (ripe spores not seen)

Habitat. On soil or logs, often in dry or alpine situations.

Distribution. Apparently as for the species. Canterbury Upper Godley Valley, 5,200ft, Scott 158. Otago: Merton, 3899; Dunedm, 1934, 3543, Ravensbourne, 1181, 1976, Sandymount, T 1773 (and in CHR as ZA 523 sub P. pusilla), Maungatua, 1,000ft, Mr. 925 (pr. p.), 3,000ft, 0515, Walporn, T 196; Taierl Mouth, 1673, 1675. Southland: Manapour (D Hamilton), 0717., Riverton, T 797.Exstccata SEEN No. 1944–29, ex Herb. R. D. Hoogland, in Auckland University Botany Department.

This does not seem to be a habitat form since specimens are sometimes growing with the typical form of the species and the difference between them is quite marked. It is not a growth form, since specimens 1181 and 1976 are collections from the

same plant taken five years apart. Possibly the variety is produced by an abnormally slow growing plant, since the veined surface in Peltigera is in large part due to separation of the lower surface by the more rapidly growing algal layer and cortex. Specimen 1976 had almost doubled in size in four years, which is about the usual lifetime for Peltigera species. The Sandymount specimen T 1773 certainly has the appearance of P. pusilla (P. canina var. spuria) but lacks any trace of tomentum; it could equally well be a small specimen of var. magyarica. The specimens under this variety name are not very homogenous, the size of the lobes varying from 15 mm in diameter to 3 mm. So far as I can discover Maas Geesteranus' combination has not been published.

Peltigera dolichorhiza Nyl. var. dolichorhiza.

Peltigera polydactyla var. dolichorhiza Nyl., Synops. Lich. Vol I, 327 (1860).

Muller, J. Linn. Soc. Bot. 32, 201 (1896)

Peltigera dolichorhiza Nyl., Lich. N. Z., 43 (1888)

Hellb, Bihang Kgl. Svensk Vetensk. Akad. Handl., 21, III.

(13), 29 (1896)

Zahlbr., Lich. N. Z., 46 (1941)

Peltigera dolichorhiza f javanica Gyelnik, Nyt Mag Naturvidenskap, 68, 269 (1930 et apud Zahlbr. Lich. N. Z. 46 (1941)

Thallus as for P. polydactyla f minor except that the rhizines are up to 10 mm long, simple and tapering, usually dark coloured; cortex 25–30μ thick, algal layer 25–35μ, medulla 180–240μ of rather compact hyphae 8μ dia. Apothecia about 3 mm dia., hymenium 100–130μ high, hypothecium rather dark brown, 35–70μ thick, asci 4–6 spored, spores brownish when mature, (3-) 5–7 (-9) septate, 50–80 × 3–4μ.

Habitat In damp localities on soil, mosses, logs, rarely stones, etc.

Distribution. New Zealand, North and Tropical America, Australia and probably elsewhere in the southern hemisphere. North Island: Whangarei (Given) CHR, (?) Volcanic Plateau (Attwood) CHR; Maungatawhiri (G20 Moore) CHR; Tiritea (G1 Chamberlain, 2 coll. and Allan) CHR; Pirongia (W12 Allan) CHR; Kaingaroa Plain, Allison 246, Wairarapa (Colenso 403) (Colenso 2591) WELT; Kahuraamake (Colenso 2904) WELT Nelson: Korere (Allan) CHR; Hundalee, Mr. 1301 Marlborough: Pelorous Bridge, Mr. 1337. Westland: Greymouth, Mr. 6882 Canterbury. Waipara (G50, Moore sub P. dolichorhiza f. javanica) CHR; Hermitage, 3,000ft, Sc 291; Otago. Kaituna, T 1955, Haast Pass, 0954; Huxley River, 1848, Trotter's Gorge, T 1400 (pr p); Leith Valley, Mr. 749, T 1740 (pr. p) and in CHR (G53), Abbott's Hill, T 936, T 949, Lee Stream, 0695; Flagstaff, 1,500ft, Mr. 1150 (pr p); Taieri Mouth, 1384, 1670, 1674 Southland: Mackinnon Pass, 3,000ft, T 2900 (unusually broad lobes); Doubtful Sound, T 2879, T 2885, 3945, Forest Hill, 0392, Riverton, T 793 Stewart Island Oban, Mr. 78; Wilton's Bush, Mr. 5425.

Exsiccata Seen Gyelnik Herb. Lich. (Jamaica, Plitt) (Thallus thicker than in New Zealand specimens)

Nylander's original spelling of the species name with a single “r” was changed to “rr” in 1888 The difference between polydactyla and dolichorhiza is perhaps hardly enough to justify separation at the species level, but nevertheless is remarkably constant and I have not been in doubt about assigning more than a very few of the specimens to one or the other species. Since dolichorhiza was not segregated till 1860, and it is apparently the commonest member of the genus in New Zealand, it is more than likely that the early records of P. polydactyla refer in large part to dolichorhiza. The polydactyla-dolichorhiza complex is in need of controlled growth studies to ascertain the constancy of the taxa listed as varieties. It is a reasonable suspicion that they are not, in which case the best treatment may be to leave the latter as a variety of polydactyla and reduce the varieties to the rank of forms or to synonymy. It seems not without significance that parallel pusilloid, thinner or

obscurely-veined varieties are found in each of the species virescens, polydactyla, dolichorhiza and canina Gylenik's f javanica is merely a freely fruiting form.

Peltigera dolichorhiza var. nana (? Nyl.) Murray comb. nov.

Peltigera polydactyla var. nana Nyl., in sched sec Knight Peltigera nana Wainio, Philipp J. Sci, 8, 114 (1913) and Gyelnik, Ann Mus Nat Hung, 30, 132 (1936) Zahlbr. Lich. N. Z., 45 (1941)

Peltigera nana f nervosa Gyelink apud Zahlbr., Lich. N. Z. 45 (1941) Thallus thinner than for dolichorhiza, more or less shining, light reddish-brown or green above with reddish veins and rhizines below, cortex 30μ, algal layer 25–38μ thick, medulla 70–100μ thick of rather compact layers of hyphae 5μ in dia. Apothecia 3–5 mm dia., hymenium 90–110μ thick, hypothecium brown, 50μ thick, asci 4–6-spored, spores yellowish, 3–5 (-7) septate, mostly 65 × 4μ

Habitat On mosses and plant debris in damp places.

Distribution Philippines, New Zealand, ? Jamaica North Island Little Barrier Island (G52, Hamilton, sub P. nana f venosa) CHR, Hen Islands (Moore) CHR; Mohikinui (Allan), CHR, Papataki, CHR, Awakino, CHR, Wellington (Allan), CHR, Totara Reserve, CHR; Wai-iti Stream, CHR, Hawke's Bay (Colenso, 1671), WELT, Plimmerton (Knight, sub P. polydactyla var. nana Nyl.), WELT Westland. Greymouth, Mr. 5430, Lake Kaniere, Mr. 1349 Otago: Waitati, T 1893 and in CHR (G55), Mihiwaka, Mr. 921 Southland Freshwater Valley, T 3055, Doubtful Sound, T 2883, Waipai, Mr. 1304 Stewart Island Port William, Mr. 707.

Exsiccata Seen Gyeln. Lichenotheca (Jamaica, Plitt) sub P. nana var. meridiana (doubtful)

I have not been able to find whether Nylander's name has been published, but it appears several times on specimens in the Knight collection. Gyelnik (1936) combined his species P. meridiana from Jamaica with Wainio's from the Philippines, listing the Jamaican plant as a variety and making Wainio's species P. nana var. philippina (an illegitimate new name) The New Zealand plant from Waitati (CHR, G55) he listed merely as the species, it agrees very well with Wainio's and Gyelnik's descriptions, whereas the Jamaican specimen distributed by Gyelnik does not. The differences from var. dolichorhiza are usually fairly clearly marked, although some specimens—e g, T 2883, come close to var. dolichorhiza, so I have reduced the taxon in rank. The veins are usually of a light reddish colour not seen in var. dolichorhiza Gyelnik's nana f venosa according to the isotype specimen is scarcely distinguishable in venation from the more typical forms Some North Island specimens are nearly as thick as var. dolichorhiza but have the other characteristic features of var. nana.

Gyelnik (1931b) describes a Peltigera oceanica apparently from the Pacific Islands which is evidently only a pusilloid variety of dolichorhiza—probably it may be found also in New Zealand, although I have seen no specimens which are clearly referable to this.

Peltigera malacea (Ach.) Funck.

Peltidea malacea Ach, Synops Lich., 240 (1814)

Peltigera malacea Funck, Crypt Gewachse, heft 33, 5 (1827)

Hellborn, Bihang Kgl. Svensk Vetensk Akad Handl 21, III (13) 28 (1896)

Lobes up to 4 cm long by 2 cm wide, but usually considerably smaller, and lobes in alpine plants more or less pusilloid, tomentum sparse, mostly marginal and thallus shining in the centre Tomentum on undersurface brown Thallus rather soft, apothecia marginal, found, 4–8 mm dia., spores 3–5-septate, 58–74 × 5–6μ (Description adapted from Nylander (1860)

Habitat On mosses, usually in alpine situations.

Distribution Europe, Himalayas, North America, Kerguelen, New Zealand.

The species is reported from the Bealey River (Canterbury) and Papakauri (Auckland) by Hellborn (1896). I have not seen New Zealand specimens, but it should not be difficult to recognise. The tomentum on the upper surface may be almost absent, as in a specimen from Europe distributed in Gyelnik's Lichenotheca (Timko).

Peltigera canina (L) Willd var. canina

Lichen caninus L. Sp. Pl. 1149 (1753)

Peltigera canina Willd. Flora Berolinens., Prodom 347 (1787)

Thallus usually large, up to 20 cm across, with lobes (2-) 5 (-10) cm long by (½-)1–2 cm broad, grey-green sometimes turning yellowish or brownish-green in the herbarium. Upper surface dull, thinly tomentose particularly marginally, lower surface with white to brownish tomentum and rather narrow elevated veins bearing mostly simple rhizines, veins smooth to fibrillose. Cortex 25–30μ, algal layer 40–60μ, medulla (100-) 200–250μ of more or less parallel hyphae 5–8μ dia Apothecia on extended lobules, ca 4 mm dia; with thin crenulate margins, hypothecium reddish-brown 50μ thick; hymenium 100μ thick, asci 6–8 spored, more or less cylindrical; spores aciculai, hyaline or pale yellow, 3 (-5) septate, 45–70 × 3–4μ.

Habitat On soil, moss, logs, etc., in damp, shady places.

Distribution. Cosmopolitan. North Island: ? Ruakura, Allan, CHR; New Plymouth, Moore, CHR (thin, apparently f. membranacea); Tahuna (Colenso, 5017) WELT. Otago: Dunedin, T 1740 (pr. p., and in CHR G53), 1187 (pr. p.), 1935, 1973, Maungatua, 1,500ft, 0392a, Mr. 925 (pr. p.); Taieri Mouth, 1668, 1669. Southland: Eglinton, Rawlings CHR (KG10).

Exsiccata SEEN Lich. Suecici (Vrang), Fl. v. Bayern (Royer), Swed. Lich. (Magnusson 7619), Herb Schallert, Pl. reg. mag. (Dusen 260), Fl. Hung (Timkohy).

Peltigera canina has been variously split into forms, varieties and species on the basis of thallus thickness, character of tomentum, colour of veins and rhizines, size of lobes, presence of marginal lacerations, etc., until the list of synonyms includes at least 30 names. No doubt some of the published forms and varieties correspond to constant entities, but many certainly do not Thomson (1950) divides the North American material into var. albescens with white veins and rhizines and var. ulorrhiza with brown veins, but European and New Zealand specimens appear to form a continuous series in this respect according to the few examples I have seen, and I have not used these names.

Thomson also separated North American specimens with thinner thalh and a penicillate appearance of the veins as P. membranacea Nyl, distinctions which also seem difficult to maintain (compare Lindahl, 1953). According to Thomson the thickness of the medulla in P. canina is 300–500μ and in P. membranacea is 70–110μ, but several of the above European specimens of canina come between these ranges and one (Magnusson 7619, Swed. Lich.) has both smooth and penicillate veins in the same specimen. New Zealand specimens of canina are distinguished from European by their generally thinner, narrower lobes, and closely resemble Schallert's and Dusen's American specimens. A penicillate appearance of veins and rhizines is sometimes shown by our specimens of var. canina and var. rufescens, but it is not constant even in the same specimen Variations of this sort are similarly apparent in New Zealand specimens of P. polydactyla; I have seen no specimens in either of these species approaching the large size apparently common in Europe.

It has been noted in other cases that distinctions between certain lichen species or varieties may be clear-cut in one country but not in another, and this phenomenon may explain the different treatments of the P. canina complex in North America and in Europe. The fact that New Zealand specimens of P. canina do not match the European forms exactly favours the assumption that the species is truly indigenous.

A specimen from Doubtful Sound (3946) seems to belong to the P. canina complex, although it differs from all varieties or species of which I have seen descriptions. The lobes resemble those of canina, but have a few sorediate spots, while the undersurface has no distinct rhizines but caninaeform veins with a 2 mm thick mat of loose anastomosing hyphae. It is sterile, so I have preferred not to describe it as new in the meantime.

• Peltigera canina var. rufescens (Weis) Mudd.

• Lichen caninus var. rufescens Weis., Plant Cryptog Flor Goettigens, 79 (1770)

• Peltigera canina var. rufescens Mudd., Manual Brit. Lich. 82 (1861)

• Peltigera rufescens Humb., Fl. Friburg Specim, 2 (1793) Bab in Hook Fl. N. Z.

• Vol. II., 271 (1855)

• Linds, Trans. Linn. Soc. 25, 521 (1866). Nyl., Synops Lich Vol. 1, 325 (1860)

• J. Linn. Soc. Bot. 9, 246 (1865)

• Hellborn, Bihang Kgl. Svensk Vetensk Akad. Handl. 21, III (13) 28 (1896)

Peltigera rufescens var. spuria Hooker, Handb N. Z. Fl 566 (1867)

Thallus 3–12 cm dia., with lobes 5–25 mm long by 5–12 mm wide, brownish or reddishbrown, veins dark-brown to nearly white but lighter near the periphery., other characters as in var. canina, but tips of lobes sometimes more scabrid than tomentose.

Habitat. Mossy banks, rocks and logs, in more open situations than var. canina.

Distribution. Probably cosmopolitan. North Island: Waitakere, 3456, Wellington. Allan (CHR, pr. p.) Marlborough Waihopai, Mr. 4211 (pr. p.), Onamalutu, Mr. 4214 (pr. p.) Otago: Waikouaiti, T 2483, Dunedm and vicinity, T 733, T 2324 (and in CHR), Mr. 5421, 038, 0391, 1213, 1214, 1977., Mihiwaka, Mr. 706 (pr p), Mt. Cargill, 3782, 3783, Mr. 5423; Flagstaff, 1,500ft, Mr. 1137; Taieri Beach; 1424; Akatore, 1556, Campbell Island Oliver (WELT 7) (uncertain)

Although the differences between var. canina and var. rufescens are difficult to express quantitatively, they are usually easily seen in both fresh and herbarium specimens, and most of the specimens in a moderate-sized collection can be placed in one or other variety without hesitation. Besides the generally smaller size of the lobes and the brittleness of var. rufescens, there is a definite habitat difference, although occasionally they can be found growing together. This is the case also in Europe according to Lindahl (1953). This is certainly the commonest form of P. canina in New Zealand, as it is also in North America In alpine situations it has a thicker thallus with ascending or mrolled crisp margins and dark veins (e. g., Mr. 1137, from Flagstaff). In Europe it has evidently been confused with var. canina, and Central European specimens. I have seen distributed as P. rufescens can mostly be referred to P. canina var. canina (e. g., Flor Hungarica Nos. 75 and 227 and specimens from Budapest and Bucharest Museums)

The specimen from Campbell Island (WELT 7) does not exactly match any I have seen from the mainland, most of the lobes are longer than normal with relatively broad and little elevated veins, which have an appearance intermediate between those of rufescens and dolichorhiza Peltigera coloradoensis Gyelnik (1930)is a species like rufescens but with polydactylaeform veins and fasciculate rhizines (although Thomson (1950) treats it as synonymous with var. rufescens) and thus differs from the Campbell Id specimen only in the nature of the rhizines Pending further collections from Campbell Id I have left this specimen under var. rufescens

Specimens of P. canina var. rufescens are sometimes attacked by the fungus Illosporium carneum Fr. which forms small pink powdery spots on the upper surface; old specimens of the fungus can be mistaken for soredia.

Peltigera canina (L) Willd var. spuria (Ach.) Schaer.

• Lichen spurius Ach., Lichenogr Suec Prodom 159 (1798)

• Peltigera canina var. spuria Schaer, Lich Helvet Spicil (6), 265 (1833)

• Peltigera pusilla Muller, J. Linn. Soc. Bot. 32, 201 (1896)

• Peltigera rufescens Hook, Handb. N. Z. Fl. 566 (1867)

• Peltigera canina var. pusilla Bab in Hook., Fl. N. Z., 271 (1855)

• Peltigera rufescens var. spuria, Kirk. Trans. N. Z. Inst., 4, 235 (1871)

Plant varying from single small cochleate lobes 3 mm in dia to thalli 15–20 mm across, lobes ascending with incurled margins and small apothecia on short lobules, tomentum often restricted to margins.

Habitat Clay banks, alpine localities, usually in dry places.

Distribution. Probably cosmopolitan North Island. Napier (Colenso) WELT Canterbury Godley Valley, 6,200ft, Scott 156 Otago Haast Pass (R. F. Smith) 0937, Ravensbourne, 0596.

Exsiccata Seen. Fl. Suecica (Hülphers) (sub P. spuria).

Although commonly considered a separate species, this has every appearance of being a habitat form, although I have not seen specimens intermediate between this and var. rufescens or var. canina. The microscopic characters do not differ from those of var. rufescens.

Peltigera canina (L) Willd var. spuria (Ach.) Schaer f. sorediata

Schaer., Enumer. Critic. Lich. Europ. 20 (1850)

As for var. spuria but with small round spots of coarse greyish-blue or greyish-green soredia, like eroded places.

Habitat and Distribution. As for the variety. Canterbury: Lake Tekapo, Mason, 10 Otago Maitland Valley, 1734, Flagstaff, 1,000ft, 0900; Taieri Mouth, 1671.

Exsiccata Seen Swed. Lich. (Magnusson, 13735) (sub P. erumpens). Although there is a considerable number of synonyms at the species level for this form there now seems little doubt that it is no more than a growth form of the variety (of Thomson, 1950 and Dahl, 1950).

The earliest valid name for this form would seem to be Peltigera canina var. sorediifera Schaer, but I have not been able to see the relevant literature.

Peltigera praetextata (Flk.) Wain.

Peltidea ulorrhiza var. praetextata Flk. apud Sommerf., Suppl. Flor. Lappon, 123 (1826)Peltigera praetextata Wain, Termeszetr Fuzetek, 22, 306 (1899). Peltigera nitens f zeelandica Gyeln, Bot Lapok, 28, 60 (1929)

Thallus about 8 cm dia, with lobes to 25 mm long by about 10 mm wide, brownish, with clusters of subsquamulose isidia along margins and imperfections on the upper surface and white scales near the tips of the lobes which curl downwards, veins elevated (caninaeform), whitish to brown, obscure near the margin and with a fuzzy appearance due to projecting hyphae rhiznes simple to more or less fasciculate. Other and miciroscopic characters identical with those of P. canina.

Habitat. On soil and damp rocks, etc.

Distribution. Eurasia, North America, Japan. Otago: Flagstaff, 1,500ft, Mr. 1150 (pr. p.), Mr. 1152 (pr. p.)

Exsiccata Seen. Fl. suecica (Hulphers), Swed Lich. (Magnusson, 11650). This species has been very variously treated by lichenologists recently. Thomson (1950) reduced it to a form of canina var. rufescens, remarking that it was merely a form of this variety with the edges regenerating after damage from “insect bites”, but Lindahl (1953) has shown from experiments on P. canina and rufescens that this is not so Sometimes, apparently, particular plants do not develop isidia, in which case they are hardly distinguishable from var. rufescens Although the species is usually described as “tomentose above”, Lindahl mentions the presence of scales, and the Otago plants are clearly scaly rather than tomentose under the microscope,

and are thus separable from most states of canina var. rufescens. The scales are formed by the growth of several adjacent cortical hyphae which fuse together to form a white mass 20–50μ square and about 30μ thick. The scales have rather the appearance of a decomposed areolate cortex In specimens of P. praetextata var. subcanina distributed by Gyelnik the tips of the lobes are minutely tomentose, and these may be more nearly related to P. canina. Lindahl (1953) reduces P. nitens (Anders) Gyelnik to synonymy with praetextata, but it is clearly regarded by Gyelnik as smooth above and thus close to P. virescens. The Otago specimens like Gyelnik's P. nitens f zeelandica have crisp margins, but I consider this appearance to be due to the plants growing in rather dry or alpine situations, particularly since similar forms are shown by all the Peltigera species under these conditions P. nitens f zeelandica was founded on one of Berggren's specimens (exact locality not stated)now in Stockholm, it is not impossible that it is a form of praetextata with the scales almost absent, as in some lobes of Mr. 1150.

Solorina Ach.

Thallus small foliose, corticate above, ecoiticate below and with indistinct veins and scattered rhizines below Algae commonly green and blue-green in the same thallus Apothecia immarginate, scattered over the upper surface, spores brown several celled.

The genus is predominantly Northern and African, and until now there has been only one record of it for New Zealand or Australia.

Solorina crocea (L) Ach.

Lichen croceus L., Sp. Pl., 1149 (1753)

Solorina crocea Ach., Lich. Univ. 149 (1810)

Du Rietz, S [ unclear: ] bot Tidskr 20, 300 (1926)

Ibid., Rept. Austral Ass. Ado. Sci. 628 (1928)

Thallus roundish, lobed, 1–10 cm dia, greenish brown above, copper-coloured below with indistinct subreticulate veins and a few rhizines, cortex 300–500μ thick, of vertical thick-walled hyaline hyphae about 4μ dia and containing pyramidal or tooth-like sections containing mostly small algae (said to be Dactylococcus) which reach almost to the surface Medulla and algal layer 300–600μ thick, of parallel, compactly layered hyphae 3–8μ dia coated with orange granules, K + purple Algal layer not clearly delimited, of small green algae mixed with scattered colonies and frequent more or less ovoid cephalodia containing Nostoc Apothecia oblong to round, plane, dark brown, not depressed, to 1 cm dia, hypothecium hyaline, 25μ thick, hymenium 125μ, hyaline with thin brownish epithecium, paraphyses simple, 2μ thick, conglutinate, asci 6–8-spored, spores 1-septate, hyaline becoming brownish, 26–34 × 8–10½μ, oblong-ellipsoid, sometimes slightly constricted at septum.

Habitat On soil in subalpine situations.

Distribution Eurasia, North America, Himalayas, New Zealand Canterbury Craigieburn Range, 5,500ft (A. F. Mark), 4162, Mount Peel, Allan (CHR), Godley River, 6,000ft, Sc 268, (D Scott) 4164 Otago Old Man Range, 5,000ft (D Billings, NZL) 4160.

Exsiccata Seen Schaer exsicc no 24 (WELT), Metzger (WELT), Lich. scand (Zetterstedt & Wickborn) (Otago Museum)

The New Zealand specimens are macroscopically indistinguishable from European S. crocea, although the spores are smaller and paler than reported for European plants (Nylander, 1860, gives the spores as 34–53 × 10–13μ). The occurience of S. crocea in New Zealand is of considerable phytogeographical interest since the nearest known report is from Java. It is curious that it was collected only once before 1959, despite the characteristic and striking appearance of the species. Although apparently not common, it is evidently widely distributed on the mountains of inland Otago and Canterbury, and can hardly have been introduced in recent times. If further collections sustain the spore differences seen (only one fruiting specimen, 4162, has been found), the New Zealand plants may prove to be taxonomically separable, and support a long isolation of the South Island lichen population.

of the Stephen's Island bush. This species, which was so common in Captain Cook's time, was virtually extinct on the mainland by the early twentieth century, but abounded on certain of the small outlying islands (Cockayne, 1907a), where it is still frequent in seabud colonies.

The Hymenanthera obovata was of low creeping form, seldom exceeding a height of 30 cm, the Coprosma repens averaged about 50 cm and the Hebe elliptica not much more than 1 m.

2. Local Distribution of Communities

The general distribution pattern of communities was from Poa caespitosa tussock and low Muehlenbeckia complexa on the eastern summit (Muehlenbeckia on the more sheltered parts) to stunted Coprosma repens/ Hebe elliptica scrub on the sheltered N. E. slopes; a mixed crevice flora on the S.E. and S. and Disphyma australe leading down to the western Salicornietum. Coprosma on the W. side descended to about 120ft (37 m) above sea level, Hebe to about 200ft (60 m).

On the extensive areas of eroding gravel in the N. tentative colonisation by prostrate clumps of Lepidium oleraceum and Rhagodia triandra had occurred with a liberal scattering of autumn-germinated Senecio lautus seedlings. The Senecio descended closest to sea level, below the Salicornia in places, but it was doubtful if seedlings in such situations would reach maturity. Desphyma australe was creeping across the denuded gravel from the margins, and a few tufts of Poa caespitosa and Agropyron scabrum had become established on the upper parts.

The chief subordinate species in the Poa/ Muehlenbeckia community of the island crest were Coprosma repens and Wahlenbergia flexilis with Apium prostiatum at the base of the lighthouse tower. The Coprosma repens and Hebe elliptica possessed a dense, almost unbroken leaf canopy which excluded most other plants but was penetrable by the hundreds of petrels which tunnelled beneath. (Plate 17, b.)

Considerable portions of the Disphyma australe communities were quite pure or with but a few grasses and low shrubs (Plate 18, a). The Salicornia australis community typically included Suaeda maritima, Disphyma australe and Senecio lautus with a little Rhagodia triandra towards the top.

Figure I
Colony of Dove Petrels Diving Petrels & Blue Penguins. Plants in Relation to Exposure I. Little Brother Island, Cook Strait

Much the greatest floristic variety was to be seen on the broken upper slopes of the S, S. E. and S. W. in positions too exposed to permit close growths of shrubs but not sufficiently so to eliminate all but the most salt-resistant species of Salicornietum and Disphymetum.

Figure 1 shows the plant zonation along a 195.5 m long transect across the island from W. to E. The transect ascended from the lower limit of vegetation approximately 5.5m above h.w.m. up the exposed W. slope, across the most sheltered part of the island in the northern lee of the lighthouse buildings and out to more exposed conditions again on the E. where it terminated at the edge of a precipice approximately 46m high.

Salicornia australis dominated the first 56 m of the transect with Suaeda maritima, a form with flattened leaves very different from the terete-leaved British S. maritima of salt marshes, rising to local dominance in the middle and upper part of the belt and Senecio lautus seedlings maintaining a steady 5–10% ground cover throughout.

The soil pockets between a series of vertical steps in which the upper end of the slope terminated were co-dominated by Salicornia and Disphyma, and the latter remaind dominant until about 92 m.

Rhagodia triandra occurred spasmodically in the open and Poa caespitosa, Agropyron scabrum, Apium prostratum and Coprosma repens, in the slight local shelter afforded by rock outcrops. Other less common species recorded in this part of the transect were Asplenium obtusatum, Spergularia media, Tillaea moschata, Crepis novae-zelandiae and Senecio lyallii.

The area sheltered from the S. by the lighthouse buildings was occupied by a dense Hebe elliptica scrub which merged into equally dense but lower Coprosma repens and finally to Disphyma australe and a scatter of mixed cliff plants including Lepidium oleraceum.

The S. to N. transect illustrated in Fig. 2 did not show such a regular zonation, the vegetation on the southern slopes being more diverse. It was 212 m long and passed over the crest of the island where it was displaced 9 m to the E. to avoid the lighthouse tower. Twenty species, a moss and a leafy liverwort occurred in the belt; only 16 angiosperms and no bryophytes in the W. to E. belt. Considerable amounts of filamentous and thalloid green algae occurred in both transects.

The Salicornia zone started approximately 11 m above h.w.m., below which there were no suitable crevices, and was quite narrow. The Disphyma above was confined as dominant to the rockier patches, Poa, Agropyron and Coprosma sharing dominance on the deeper soil with Muehlenbeckia, Wahlenbergia and Rhagodia quite frequent. Poa and Muehlenbeckia occupied the summit, Disphyma the rocky upper slope to the N. and Coprosma the deeper peaty soil below, extending almost to the edge of the northern cliff about 32 m above sea level.

Both Cockayne (1928, p. 100) and Cooke (1911) state that Salicornia australis approaches closer to the sea in exposed conditions than does any other species, but Hamilton (1936) writing of Little Barrier Island, states that Disphyma australe is the only species present where excessive spray falls, and that “with less spray Disphyma is less important and often supplanted by Salicornia”.

Observations made on the Bass Strait Islands of Australia suggest that the above ground parts of Disphyma are quite as resistant to deposition of wind-borne sea spray as are those of Salicornia, but that the species is much less tolerant of salt water at its roots and tends to yield place to Salicornia where the spray is sufficiently heavy to run through the foliage mats to the underlying soil.

3. “Scorching” Effect of Salt Gales.

The unusually severe salt gale left its mark on the vegetation, and in places the Salicornia mat of the W. had been ripped from its anchorage and rolled back like a carpet (Plate 18, b). Considerable clumps of this and other species were blown into the sea or washed off in the heavy rains which followed the storm and leached much of the accumulated salt from the soil.

It was not possible to crawl to the windward side of the island while the gale was in progress to observe deposition of sea water there, but a continuous rain of it poured down on the crest and upper leeward slopes Waves sweeping round both sides of the island met with terrific impact at the northern “Blowhole”, shooting high into the air and deluging most of the rest of the leeward slopes.

Thus, although the more halophytic communities received a greater drenching than the less halophytic, all received more than the average which had allowed that particular type of community to develop, and the component species were damaged accordingly.

The soft, glaucous leaves of Sonclius oleraceus, Rhagodia triandra and Crepis novae-zelandiae and the succulent ones of Lepidium oleraceum had suffered a general blackening and curling of the margins except where sheltered by rocks or adjacent vegetation. (Lepidium, although possessing halophytic characters, was seen to have suffered in this way on islands visited in various parts of New Zealand).

The prostrate rosette habit of Senecio lyallii, Hypochoeris radicata and, to a smaller extent, Wahlenbergia flexilis had enabled many of these plants to escape in the shelter of taller species. Apium prostratum probably owed its high survival rate to the fact that it occurred largely in sheltered crevices. Coprosma repent and Hebe elliptica showed quite severe leaf curl at the extremilies of their range, but had escaped relatively unharmed further from the sea. Of the grasses the softerleaved Agropyron scabrum suffered more harm than did the needle-leaved Poa caespitosa.

The relative damage suffered by some of the more important species is set out in Table I.

This anomaly masks to a large extent the differences between the chloride contents of the two groups—the mineral soils occurring in all the more exposed situations, the peats usually only where sheltered. In spite of this, however, the differences are still apparent and are set out diagrammatically in Fig. 3.

The black vertical columns at the base of the figure represent the percentage of salt present, the name of the species of the immediate sample area is placed at the top of each column and the generally dominant species for the region as a whole indicated by the horizontal bars at the top of the diagram.

Five readings showed more than 2% by weight of chlorides, and three of these were rocky outcrops catching the full force of the gale, whilst the other two were in the lower part of the western Salicornietum.

The trend in the W.-E. transect was for salinity to diminish from W. to E. with an unusually high result on a Disphyma-clad outcrop at 55 metres and an unusually low one in the hollow which this sheltered at 73 metres.

The highest reading in the S.-N. transect was on the island summit, where 3.92% of chlorides by weight of the light, organie-rich crevice soil was recorded. Both the summit and an outcrop at 20 yards with 2.92% of chlorides afforded appreciable shelter to the land to their immediate N. where readings dropped as low as 0.28%.

With the salinity, as with the vegetation, no such marked gradation was apparent in the S.-N. transect as in the W.-E. one, this being largely due to the broken nature of the ground and alternation of hollows and eminences. Where an outcropping rock occurred at the sampling point the sample was taken from as close as possible in a soil-filled hollow, so results may err on the low side of the true figure. The soil giving the high reading near the northern cliff edge was affected by splash from the nearby “blowhole”.

“Salt-scorching” of the more mesophytic species, particularly the English pasture plants, had occurred up to about 400 feet above sea level, these plants having a generally wilted and blackened appearance as though frosted (Table I).

Only eight soil samples were collected, too few to be of great significance but nevertheless showing decreasing salinity and pH gradients from the lower slopes upwards. They fell into the three categories of broken cliff face about 100ft (31 m) above sea level, tussocky pasture from about 100–300ft (31–93 m) and tree-clad areas higher up.

The first sample area showed a chloride content similar to that of the Little Brother Poa and Disphyma communities (1.06% Cl¹) and supported a mixed crevice vegetation with these two species, Coprosma repens, Rhagodia triandra and Senecio lautus. The pH was higher than elsewhere, 7.0 (air dry), 7.8 (moist) m spite of the peaty nature of the soil, again suggesting that sea salt may have a similar effect to calcareous soil washings m producing a neutral “fen peat”.

Chloride content dropped to approximately one-third on the petrel slopes above, the two light, retentive, peaty samples. giving readings twice as high as did the two heavy, non-retentive mineral samples. The average pH was much lower, 4.52 (air dry) and 5.8 (moist).

In a patch of cliff bush dominated by Myoporum laetum the chloride content had dropped to 0.1% and pH to 3.97 (air dry) and 3.6 (moist). In the more extensive inland bush with mixed dominants (Dysoxylum spectabile, Macropiper excelsum, Melicytus ramiflorus, Olearia paniculata, Rhopalostylis sapida, Suttonia australis, etc.) most blown salt was prevented from reaching the soil by the dense leaf canopy and the average chloride content of the peaty soil was only 0.04%, pH 3.78 (air dry) and 36 (moist).

3. Islands Near the Entrance to Queen Charlotte Sound.

Islands visited after leaving Little Brother on May 23 were Motungarara, the Westerly Twin, Komakohua and Long Island. All were afforded fair protection by the rocky headlands at the entrance to Queen Charlotte Sound and woody vegetation descended to within a few metres of the sea on leeward shores.

Seven soil samples were taken from among shearwater burrows on Motungarara Island and the most saline came from open Disphymetum at the exposed seaward end. The chloride content was 0.72%, pH 6.1 (air dry) and 7.4 (moist). The least salme samples were taken from the mixed bush of the island summit (Cl¹ 0.16 and 0.15%, pH 5.2 and 5.7 (air dry), 6.8 and 7.3 (moist)). This area was covered with a dense low canopy of Myoporum laetum, Melicytus ramiflorus, Nothopanax arboreum and Olearia paniculata.

The four samples of mtermediate salmity (0.19–0.42% Cl¹) came from Coprosma repens scrub, sometimes an open community with Disphyma and Rhagodia, sometimes a closed one with Nothopanax.

The Westerly Twin was a small stack on which only 22 plant species were recorded. The steeper, more exposed sides and the crest were burrowed by blue pengums and occupied by tall grass (chiefly Poa caespitosa, Poa anceps and Agropyron scabrum) with Disphyma, Rhagodia, Senecio lautus, etc. The leeward slope was occupied by low Coprosma/Nothopanax scrub with marginal Parsonsia heterophylla and Hymenanthera obovata and Asplenium flaccidum and A. lucidum where sufficient light penetrated Only four soil samples were taken, and these showed no correlation between soil salinity and plant zonation. Soil chlorides ranged from 0.18–0.36%, pH of air dry soil from 4.8–6.7 and of moist soil from 6.7–7.3.

The western slopes of Komakohua Island were dominated below by open, eroding and heavily burrowed Disphymetum, above by Coprosmetum. As on Little Brother Island, Salicornia australis was locally dominant at the lower margin of the Disphyma and Poa caespitos at the upper. The commonest subordinates were.

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All sample areas heavily burrowed by fairy prions or mutton birds except the first, which supported a small colony of black-backed gulls.

Agropyron scabrum, Coprosma repens, Hymenanthera obovata, Rhagodia triandra, Senecio lautus and Sonchus oleraceus. The upper Coprosma zone included species not present on Little Brother, principally conspicuous open patches of Cassinia leptophylla, a common coastal and hill species on the adjacent mainland, and Phormium colensoi.

A landing was made on the N.E. of Long Island, a much larger island with a vegetation resembling that of the mainland coasts rather than of the islands described above. Most of the area was occupied by any open heathy scrub of Cassinia leptophylla, Leptospermum scoparium and Olearia solandri with tall, coarse grass between. Halophytes (Apium prostratum, Crepis novae-zelandiae, Disphyma australe, Lobelia anceps, Salicornia australis, Samolus repens, Selliera radicans, Senecio lautus and Spergularia media formed no coastal belt but occurred as scattered individuals just above h.w.m.

Correlation between increasing soil salinity and ph in 10 cook strait plant communities

Soil samples from the E. coast and N. crest showed 0.12 and 0.18% of soil chlorides and soil reaction by the coast was pH 7.4 (air dry), 8.0 (moist), and beneath Cassinia on the crest pH 6.4 (air dry), 7.1 (moist), neither sample being at all rich in organic matter.

Soil salinity and pH in relation to community dominants on Little Brother Island are compared diagrammatically in Fig. 4 with those on the other islands visited. The diagram shows the parallel trends in the two cases, but these were much clearer on Little Brother, where the environment was more uniform and the number of samples more representative (55 and 21 respectively).

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Salicornia and the other by Samolus. Beyond this the agriculture permitted by the level nature of much of the ground and the deep red volcanic soil at Egmont invalidated any comparison between this receding flat and the Brother's steeply rising cliff.

2. Bay of Islands, N. New Zealand

Bay of Islands lies approximately 400 miles N. of Little Brother on the E. coast of New Zealand, sheltered from the Tasman Sea by the land mass of Northland and from the Pacific Ocean by the land about Russell.

The principal coastal vegetation where the land has not been cleared for agriculture is Metrosideros excelsa forest, usually lacking the coastal fringe of Coprosma repens which occurs in more exposed regions such as the Hauraki Gulf and Bay of Plenty.

Some of the inlets, such as Crowell's Bay, are sufficiently sheltered to support mangrove swamp, some of the stacks such as those off Tapeka Point lighthouse, sufficiently exposed to be characterised by a Disphyma-Coprosma community as on the N. of Little Brother.

Other Little Brother species occurring on low spray-washed cliffs on the Black Rocks off Motoroa Island were Salicornia australis, and Senecio lautus. Three species occurring on more sheltered sandy pockets of these rocks and characteristic of less exposed positions on Little Brother were Apium prostratum, Atriplex hastata and Rhagodia triandra. Species of higher grassy slopes included Muehlenbeckia complexa and the commoner Little Brother aliens, Sonchus oleraceus, Hypochoeris radicata and Anagallis arvensis, together with Cassinia, Leptospermum and Phormium spp. as on the more sheltered Cook Strait islands.

3. Stewart Island, S. New Zealand

Half of the Little Brother species were found fairly commonly on the shores of islands in the Stewart group, 500 miles to the S. These were:

There was a marked difference between the type of protection found most commonly in the leaves of the dominant marginal, woody evergreens in the two areas. Those of the Stewart region possessed a thick woolly tomentum, those of the Cook Strait region a resistant shiny surface—both effective means of minimising wind abrasion and salt entry.

Few or no woody species are found in similarly exposed areas on British islands, whether affected by grazing or not, and this may be due to the higher proportion of soft-leaved deciduous species in the flora as well as to climatic differences.

The commonest tomentose species found at the seaward limit of vegetation in the S. were Senecio puffini (S. rotundifolius), Olearia angustifolia, O. colensoi and the hybrid O. traillii. One of the few seen in the Cook Strait coastal flora and considerably further back from the spray zone was O. paniculata, this leaf type occurring again in one of the commonest coastal trees further N., Metrosideros excelsa.

Hebe elliptica was the shiny-leaved species found approaching the sea most closely on Stewart Island. Typical plants of this group in the spray zone in Cook Strait were this species, Coprosma repens, Hymenanthera obovata, Myoporum laetum, Nothopanax arboreum and Parsonsia heterophylla, softer-leaved species such as Macropiper excelsum occurring further back.

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internal characters of the shell were unknown. From the external appearance Zittel referred the shells to Mytilus. Trechmann (1918, p. 202) suspected that the species might belong to the Myalinidae. Later Wilckens (1927) placed the species in Myalina, and proposed a new subgenus, Maoria, for the problem species, though the internal features were still unknown. However, the name Maoria was later found to be preoccupied, and the species was again referred to Mytilus (Marwick, 1946).

Two recent statements have been made on the generic position of problematicus. Marwick (1953, p. 66) has stated that there is no evidence to indicate that the species is Myalinid. On the other hand, Avias (1953, Pl. 24, figs. 1–3) considers that the species is more closely allied to Myalina than to Mytilus.

None of the writers who suggested that M. problematicus might be Myalinid have stated why they disputed the affinity with Mytilus. The reason is probably because problematicus is inequivalve, a condition rare in the Mytilidae, and common in the Myalinidae.

Mytilus mirabilis Trechmann

The relative inflation of the valves is connected with another problem of M. problematicus, that of its relationship to M. mirabilis Trechmann. Zittel (1864) had based the species Mytilus problematicus on a relatively little inflated right valve. In 1918, Trechmann erected a new species Mytilus mirabilis for high inflated shells (Pl. 19, figs, 1, 2.)^* that occur with M. problematicus. These had previously been confused with Gryphaea (Park, 1903, p. 397; Bell, Clarke and Marshall, 1911, p. 20; Marshall, 1911, p. 17; Ongley, 1940). Trechmann (1918, p. 202) stated that the two species were connected by a series of intermediate forms, and considered that mirabilis “suggests … an enormously overgrown specimen of Mytilus problematicus in which the valves have become strongly arched”.

New Name for M. mirabilis Trechmann. The name Mytilus mirabilis is a still-born homonym of Mytilus mirabilis (Lepsius, 1878, Das weslich Süd-Tirole, p. 365., Pl. 6, figs. 3 a-c). Originally placed in Gervillea? by Lepsius, the species was transferred to Mytilus by G. Boehm (1884). Even though Trechmann's species is considered to be a subjective synonym of problematicus it should be renamed, following the recommendation of the International Commission for Zoological Nomenclature (Hemming, 1950, p. 49). Therefore the new specific name trechmanni is here proposed to replace mirabilis Trechmann non Lepsius. The newly named type was figured by Trechmann (1918, Pl. 20, fig. 20 a, b), and is kept at the British Museum (Natural History).

Growth Stage of M. problematicus. Both Wilckens (1927) and Marwick (1953) treated “Mytilus” trechmanni (= mirabilis), tentatively at least, as a distinct species. Wilckens (1927, p. 14) referred to mirabilis as a “puzzling pelecypod”, and noted that in its early growth stages it closely resembled M. problematicus. Avias (1953) considered that the two were conspecific, stating that mirabilis was the gerontic growth-stage of problematicus.

To test the relationship between M. problematicus and M. trechmanni, all the relatively undistorted specimens of the two forms at the N.Z. Geological Survey have been measured and graphs of dimensions have been plotted. One of these graphs, simplified, is presented as Text-fig. 1. Judging from the graphs, it seems impossible to separate early growth stages of trechmanni, indicated by growth lines, from specimens of problematicus. It appears that the two “species” are merely different growth-stages of one species. Therefore, following Avias (1953), trechmanni (=mirabilis) is put into subjective synonymy of problematicus.

Text-fig. 1.—Graph of right valves of little inflated Manticula problematica Zittel and highly inflated M. trechmanni n. sp, plotted from specimens and their growth-lines, showing individual ontogenies.

The best illustration of the change from a form typical of problematicus to one typical of trechmanni is given by Avias (1953, Pl. 24, figs. 1–3). Growth lines show that the young form of trechmanni is little inflated, like problematicus. As the right valves, or growth lines of other specimens, increase in size the inflation increases rapidly, particularly at the posterior-ventral margin. In the growing animal this resulted in a considerable tilting of the first formed part of the shell. Little further increase occurred in length or height; indeed the margins of some shells became slightly constricted. As a result the valve became highly arched and semi-circular in outline.

Few Mytilid species are inequivalve (Nicol, 1958, p. 58) but there are exceptions. A recent Australian genus, Stavelia Gray (1858), is inequivalve, with a short ligament carried on very stout, almost shelf-like projections (Iredale, 1939, p. 410). The hinge has no teeth. Unlike Manticula, Stavelia is twisted in outlie, and the anterior ventral margin of either the right or left valve bulges into that of the other valve, so that the margin is sinuous.

Inequivalve genera are more common amongst the Myalinidae then in Mytilidae. But none are known to be so grossly gibbose as Manticula problematica, and it is generally the left valve of Myalmids which is inflated, not the right. The ligament area of Myalinid genera is broad as in Manticula, though as a rule it is longer and more externally placed. Usually it lies on a thickened hinge area rather than on a plate. The ligament area of the upper Palaeozoic genera Septimyalina Newell (1942) and Atomodesma Beyrich (1864) shows some similarities to that of Manticula, in that it overhangs an umbonal chamber. But in these genera the left valve is more inflated than the right valve, as in other Myalinids, and an umbonal plate is developed. In addition, teeth occur in Septimyalina.

According to Newell (1942, p. 33), the outer ostracum of both valves in Myalinid genera is typically composed of polygonal prisms of calcite at right angles to the surface. By contrast, the outer ostracum in Mytilids may comprise oblique prisms (Bøggild, 1930, p. 42). The outer ostracum of Manticula shows neither of these characteristic structures: it is very thin, and apparently homogeneous, as may be the case in some Mytilids (Bøggild, 1930), and in some Myalinids. Most of the ostracum is made up of a complex lamellar layer, unlike the shell structure described for any Mytilid or Myalinid.

Manticula problematica (Zittel). Figs. 1–4, 26–30.

• 1864. Mytilus problematicus Zittel: 28, Pl. 8, figs. 1a, b.

• 1910. Mytilus problematicus Park: 72, Pl. 3, figs. 1a, b.

• 1912. Mytilus problematicus Marshall: 183, fig. 102.

• 1918. Mytilus? problematicus Trechmann: 201, Pl. 20, fig. 8.

• 1918. Mytilus? mirabilis Trechmann: 202, Pl. 20, figs. 9a, b.

• 1927. Myalina? (Maoria) problematica Wilckens: 13, Pl. 2, figs. 3–6.

• 1927. Myalina? mirabilis Wilckens: 14, Pl. 3, fig. 1.

• 1939. Maoria problematicus Marwick in Ongley: 37.

• 1946. Mytilus problematicus Marwick: 27.

• 1953. Mytilus problematicus Marwick: 66, Pl. 4, fig. 1.

• 1953. Mytilus mirabilis Marwick: 66, Pl. 5, figs. 1, 2.

• 1953. Mytilus problematicus (= M. mirabilis) Avias, Pl. 24, figs. 1–3.

• 1956. Oretia sp. Marwick in Wood: 60.

• 1959. Mytilus problematicus Waterhouse in Kingma: 23.

• 1960. “Mytilus” problematicus Grindley and Waterhouse: 262.

• not 1878. Gervillea mirabilis Lepsius: 365, Pl. 6, Figs. 3 a-c.

Type Specimen. Specimen figured by Zittel (1864, Pl. 8, figs. 1a, b) and kept at the Vienna Museum. Flügel (1959) reported that the specimen could not be found.

Type Locality. Wairoa Valley, south-east of Spring Grove. The exact position is unknown.

Material. Several hundred specimens are available in the collections at the N.Z. Geological Survey, Lower Hutt.

Description. The most striking feature of the species is the grotesque inflation of large right valves, in which the width of the shell may exceed the length and height. Left valves of equal height are only about a quarter to a third as wide.

The umbones are anterior and prosogyrous. The anterior margin is wide and recessed, with a shallow byssal gape, and a weak beak ridge defines a shallow concave false lunule. In right valves the false lunule is subdivided by a second ridge. Behind the beak the short dorsal hinge is inclined at about 80° from the anterior margin. For about 5 mm the angle between the posterior dorsal margin and the line of maximum inflation (angle α of Newell, 1942, p. 25) is low, measuring 20° to 30°. This angle rapidly increases to 50°, and is fairly constant over the rest of the shell.

Variation in shape is considerable. At one extreme specimens are particularly high, with a mytiliform appearance. These high specimens have a narrow umbonal angle of 70°, and angle α is as much as 60°. The shell is well inflated near the anterior margin, and the posterior wing is well developed.

Other specimens are elongate and oval in outline. In these the beak is blunt, with an umbonal angle measuring 90° or 95°. Angle α is low, measuring 50° to 55°. The anterior margin is little inflated and extends a little in front of the beak. The posterior wing is not so well defined in these elongated shells.

The ornament is variable. Some specimens are almost smooth. Others have fine concentric wrinkles, and others strong concentric wrinkles, which pass into overlapping lamellae near the beak. In addition, the shell surface is covered with very fine radial filae, 3 to 5 occurring in 1 mm. Coarser radial costae are visible on internal casts of a few specimens.

Internal. Hitherto the internal features of problematica have never been described: they were not visible in Geological Survey collections in spite of the abundance of material. Most of the specimens which show internal details have been collected and prepared by the writer. Not all details are yet known. However, it is found that the hinge is edentulous, unlike Mytilus (s.s.), and that the ligament rested chiefly, if not entirely, upon a triangular plate, marked by growth-lines and weak subvertical striae (Fig. 3). On the right valve the area is set a little below the commissure, and is weakly concave. On the left valve the area is more obscure, and seems to be at the level of the commissure in one specimen. Because the specimens are somewhat distorted or broken, it is not clear if the ligament also lay at each side on narrow raised shelves as though modified from an Ostreiid or Pteriid pattern. However, this does not seem likely. A short ridge is developed along the shell margin behind the plate. It somewhat resembles the nymph-ridge in Mytilids, but probably served only to strengthen the shell rather than support a ligament.

Part of the posterior muscle scar is visible on a left valve collected by the writer from GS 7615 at the junction of Manganui and Turipoto streams, near the Awakino River, in N.Z.M.S. 1, Sheet N91 (provisional edition). The scar is relatively large, extending for a quarter of the height of the valve, and is lightly impressed. It is apparently oval in shape, though the posterior margin is not well defined, so that its shape and extent are incompletely known. A small pedal scar lies above the adductor closer to the hinge, and a pallial line passes forward from the base of the scar. Traces of a small anterior scar lie close to the anterior margin, below the beak. Faint concentric growth lines lie over the surface of the scar. A scar has been observed in a similar position on another left valve from the same locality (pers. comm., Mr. J. A. Grant-Mackie, Auckland University).

Shell Structure. In large specimens the shell reaches a thickness of 6 to 7 mm, but is usually 3 to 4 mm thick. Few are sufficiently well preserved to be sectioned. Two right valves from GS 2534 with shell 3 mm thick have been sectioned parallel and at right angles to the hinge, and a left valve from beds north of Wairoa Gorge has also been sectioned along the maximum dimension. These sections show that the shell is now made up entirely of calcite. In structure it is complex and variable, and crossed by faint growth-lines parallel to the growing edge (Figs. 26–30). The outermost zone comprises clear, apparently homogeneous calcite, .1 mm thick, well preserved only in the right valve. Within is a zone 2 mm thick of bifurcating and flexuous lamellae. The lamellae are inclined inwards away from the beak, and lie roughly at right angles to the growth-lines. They seem to consist of fine horizontal or weakly oblique fibres. Extinction is parallel to the length. Between these lamellae the calcite extinguishes very weakly, and is closely wrinkled in relief. Dr. W. A. Watters, N.Z. Geological Survey, suggests (pers. comm.) that the calcite is an aggregate of minute fibres, perhaps inclined in two oblique directions. The inner margin of the complex zone is irregular and merges with a zone of either massively crystalline or apparently homogeneous calcite, up to 1 mm thick. In some sections the calcite extinguishes parallel with the lamellae; in other sections it extinguishes weakly, and is possibly made up of the fibrous aggregate. The innermost narrow zone of shell comprises two or three layers parallel to the inner margin. These are homogeneous in some sections and fibrous in others. The structure of the left valve is apparently much the same as that of the right valve.

Over the outer and inner surface, and locally within the inner recrystallized layer, are thin layers of biotite up to .2 mm thick. This association of biotite with the shell of Mollusca is common in fossils from the New Zealand Triassic (see also Waterhouse, 1960). Apparently the biotite has formed in cracks in the shell, and between the shell and matrix.

Occurrence. Manticula problematica is typical of the Otamitan stage of New Zealand. In the North Island, it occurs at many localities in the Huntly-Kawhia district (Henderson and Grange, 1926), Te Kuiti (Marwick, 1946) and the Mokau district (Henderson and Ongley, 1923). In the South Island the species is found on both limbs of a tight syncline in the Triassic beds near Nelson. It also occurs

Along the hinge of each valve is a longitudinally striated groove for the reception of the ligament, but only the anterior part of the area is visible in the specimens available. The shell is thickened under the beaks, but it is unknown if teeth are present, and the muscle scars are not seen. The shell is thin, and its structure unknown.

Rsemblances. P. spedeni is distinguished by its comparatively well rounded outline. It is less oblique than many species. P. tofanae Bittner (1895, Pl. 8, figs. 9–11; Broili, 1903, Pl. 18, fig. 20) is comparable in shape. This species occurs in the St. Cassian and Pachycardien faunas, and is also reported from the Rhaetian of the Apennines. Specimens from Sicily and Bakony have also been compared with P. tofanae by Scalia (1910) and Bittner (1901), and a left valve from the Tropites Limestone of the Himalayas was compared with P. tofanae by Diener (1906, Pl. 17, fig. 7). The European species is smaller than P. spedeni, and has a higher beak, a longer anterior ear, and longer obtuse-angled posterior wing. The concentric ornament is lower and the ligament area is higher.

Krumbeck (1924, Pl. 7, figs. 13 a, b, 14) considered that a new Ladinian species from Timor was close to P. tofanae, but the Timor shells have a larger anterior ear than either P. tofanae or P. spedeni.

A Carnian form from Bear Island, P. torelli Böhm (1903, Pl. 3, figs. 13, 16, 17, 21, 26), is moderately close in outline and ornament. It is less oblique than P. spedeni and has a much larger anterior ear and a more inflated right valve. P. torelli somewhat resembles P. angusta Saurin (1941, Pl. 1, figs. 7–13) from the upper Triassic of Indo-China. This species is also more inflated and less oblique than the New Zealand shell. Its beak is more prominent, and its anterior ears larger. As noted by Saurin, the specimen figured as P. cf. torelli Boehm by Patte (1926, Pl. 9, fig. 2) from Indo-China is close to angusta.

A closer species is P. caudata Stoppani (1858, Pl. 18, figs. 18–19) from Esino, and also described from the Pachycardientuffe by Broili (1903, Pl. 18, fig. 21), the St. Cassian beds by Bittner (1895, Pl. 8, figs. 17–18), the Grigne Group by Ronchetti (1959, Pl. 15, figs. 6, 7), as well as other Ladinian and Carnian beds of Europe. Diener (1906, Pl. 17, fig. 8) recorded a specimen with affinities to P. caudata from the Tropites Limestone of Byans, and Mansuy (1913, Pl. 3, fig. 16) compared a specimen from the Carnian of Tonkin, Indo-China. The anterior ear of P. caudata is moderately prominent, and the posterior margin meets the hinge at an acute angle. However, the species is more oblique than most of the New Zealand shells.

A. cassiana Bittner (1895, Pl. 8, figs. 6–8) from the St. Cassian beds, and Sicily, is only slightly more oblique than P. spedeni, but is smaller and more inflated. A specimen with some affinities to cassiana was recorded from Indo-China by Mansuy (1908, Pl. 17, fig. 8).

Pteria kitakamiensis Hayami (1958, Pl. 24, figs. 10, 11) from the Liassic of Japan has similar concentric ornament, but is more oblique, with a larger anterior ear.

The Problem of Megalodon globularis Trechmann

The true affinity of the species named Megalodon globularis by Trechmann (1918) has been one of the minor mysteries of the New Zealand Triassic. The lectotype of Megalodon globularis Trechmann (1918) is kept at the British Museum (Natural History), London. It is a damaged internal and external cast of a specimen with valves conjoined. Trechmann (1918) referred the species to Megalodon Sowerby (1827) on account of the thick hinge plate. However, the internal and external details of the hinge have apparently been destroyed (Dr. L. R. Cox, letter, 1958), and the real systematic position of the species is difficult to determine. Marwick, in 1953, reported that no further speciemens had been found, and left the species in the genus Megalodon.

of Wairuna trig. N.Z.M.S. 1, Sheet S170 (Provisional Edition), Fossil Sample S170/628. Collected by B. L. Wood, N.Z. Geological Survey.

Diagnosis. Unusually high in outline, anterior dorsal margin high and steeply inclined. Beak narrow and prominent, hinge short. Byssal sinus in the left valve weak. Radial ornament very fine and discontinuous.

Dimensions (in millimetres).

Description. External: The shells are relatively high, with prominent slightly prosogyrous beaks, of which the umbonal angle measures approximately 60°. The anterior margin extends almost to the lower third of the height of the shell, and the ventral and posterior margins are well rounded, the posterior margin being most extended close to mid-height. Angle α measures 60°. The greatest width lies close to mid-length in the lectotype, and at the lower third in the right valve from GS 5156.

A deeply excavated false lunule lies in front of the beaks. The byssal sinus is well developed in the right valve, and is deeply excavated under the beak (Fig. 15). In the left valve the sinus is weak though long in the specimen from GS 5156, and is apparently weak in the lectotype.

The hinge is almost entirely posterior, and has a small posterior wing. A high triangular ligament area is visible in the specimens from GS 5156. The area is marked by weak growthlines and bears a triangular ligament pit. The anterior margin of the area projects forward as a low ridge.

The ornament comprises sharply defined concentric growth-lines and steps. Very fine radial filae are present between the concentric growth lines. About two or three filae occur in 1 mm at the ventral margin of the shell, measured at abòut 20 mm from the beak.

Internal Muscle scars are faintly defined. Both the posterior and anterior impressions lie at mid-height. The anterior scar is rounded in outline, and placed close to the anterior margin below the byssal sinus. The posterior scar is apparently elongated and crescentshaped. The impressions are linked by a low ridge with a groove on each side.

The posterior part of the ligament area is supported by a short buttress, at least in the left valve from GS 5156.

The shell is more than 2 mm thick at the central margin. Its structure is unknown.

Resemblances. One of the closest species is Mysidioptera a kittlii Bittner (1895, Pl. 21, Fig. 15), from the Ladinian Red Marmor of Han Bulog and Haliluci in Bosnia, and also the Schreyeralm-Marmor of the Northern Alps and the Anisian of the Dinarides M. kittlii closely resembles M. globularis in its well rounded outline, long anterior dorsal margin, and high ligament area. Moreover, the byssal sinus of the left valve is small in both species and the ornament predominantly concentric, with fine radial striae. M. kittlii can be readily distinguished by its more roundly inflated profile, its more incurved beak, longer hinge, and larger posterior wing.

Wilckens (1909) made M. kittlii the type species of subgenus Latemaria. The subgenus is supposedly distinguished from Mysidioptera by the incurved beak and globose shell, but these characters seem to be of minor importance, and it is doubtful if the subgenus is warranted.

The well-inflated species, M. globosa Broili (1930, Pl. 22, Fig. 16) of the Pachycardientuffe is another species referred to Latemaria by Wilckens (1909) Diener (1913, p. 110) compared a shell to the same species from the Carnian Myophoria Limestone of Kashmir M. globosa agrees with the New Zealand species in its prominent beak, long false lunule, and ornament, but the European species is more elongated in outline, and has a more incurved beak than that of M. globularis.

Age. The lectotype of M. globularis and another specimen mentioned by Trechmann come from beds below Manticula problematica in Nelson, and are presumably Oretian, unless they were not in place. The specimens from GS 5156 are from

Otamitan beds. They were mistakenly mapped as Oretian by Wood (1956), but the fauna includes the Otamitan key fossil, Manticula problematica.

Discussion. The specimens from GS 5156 were referred to Oretia by Marwick in Wood (1956).

Mysidioptera riceae n.sp. Figs. 8, 12–14, 16–20.

Named after Miss Mabel Rice, former Editor of the New Zealand Journal of Geology and Geophysics.

1939. ? Gonodon mellingi non Hauer Marwick in Ongley: 40 (list).

1958. ? Atomodesma Grindley: 43.

1960. Mysidioptera n.sp. Grindley and Waterhouse: 262.

Holotype. External cast of the left valve, with the hinge and most of the internal cast, specimen TM 2101. Fig. 20.

Type Locality. GS 1431, Waiwera River, 20 to 30 chains below hairpin bend, N.Z.M.S. 1, Sheet S179 (Provisional Edition), Fossil Sample S179/475. Collected by M. Ongley.

Measured Specimens. TM 2096–7; 2101–2109, 2148, N.Z. Geological Survey.

Material. This species is represented in the collections at the Geological Survey by a few valves from a number of localities, and by nine specimens from GS 928.

Localities See Table I.

Diagnosis. Moderately small shells. Usually slightly higher than long, anterior dorsal margin high and steeply inclined. Beaks anterior, moderately prosogyrous, inconspicuous. Umbonal angle close to 90°. Posterior wing poorly developed. Byssal sinus wide in both valves. Ornament of strong concentric wrinkles and subordinate fine radial filae. Ligament area short and high.