Go to National Library of New Zealand Te Puna Mātauranga o Aotearoa
Volume 56, 1926

of the New Zealand Institute.

New Zealand Angiosperms.

[Read before the Wellington Philosophical Society, 24th September, 1924; received by Editor, 31st December. 1924; issued separaiely, 6th March, 1926.]

Plate 1.

Xeronema Callistemon sp. nov. (Plate 1.)

Folia disticha, ensiformia, equitantes, 60–105 cm. long., 3.5–4.4 cm. lat., acuta, multinervata, margine interiore ad basin concava. Scapus 70–120 cm. long., folius pluribus reductis. Racemus 15–30 cm. long., secundus, densiflorus. Bracteae scariosae, acutae, 1 cm. long. Pedicelli patentes, 2 cm. long. Perianthium segmentis, uninervatis, 10–15 mm. long. Filamenta rubra, 20–25 mm. long. Ovarium trigonum, stipitatum; stylus rubrus, 2 cm. long.; stigma obscure tricuspidato. Capsula membranacea, base expansa, loculicide trivalvis, style, staminis et perianthiis segmentis per-sistentibus. Semina extus papillose, testa nigra.

Xeronema Callistemon forms large tussocks or clusters 1–2 m. in diameter, seated on huge masses consisting of the rhizomes, roots, and dead leaf-bases. The whole plant may thus be nearly 2 m. tall from the rock-surface on which it grows.

Roots filamentous with a tough woody stele and a spongy cortex, 2 mm. diam.

Leaves distichous, equitant, ensiform, acute, multinerved, almost coriaceous in texture. Base of sheath completely overlaps bases of inner leaves; inner edge concave where outer edge of the succeeding inside leaf emerges. Measurements of largest leaf collected: Length, 105 cm.; breadth above sinus, 4.4 cm.; thickness at base, 2.7 cm.

Stem, including raceme, may reach a height of 120 cm. Leaves resemble the radical leaves, but have sheaths very long and apical portion very short. Length of sheath, 25 cm.; of lamina, 9 cm. Of these 3 or 4 are present. From just below raceme to tip the bracts are simple and membranous, without sheaths, the lower of these being about 3 cm. in length. Top of stem, including receme, dark red.

Racemes 15–30 cm. long, secund, 5 cm. diam. Bracts scarious, acute, 1-nerved, 1 cm. long. Pedicels erect, dark red, 2 cm. long. Perianth-leaves, 6, free, alternately wide (3 mm.) and narrow (1.5 mm.), 1-nerved,

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10–15 mm. long, red, paler towards the tip. In the unopened flower perianth-leaves are joined along margins, but as stamens grow they are burst apart and then turn back and hang down the pedicel withered but persistent. (Fig. 1.) Stamens 6, filaments bright red, 20–25 mm. long; anthers versatile, filament attached about centre of connective, introrse, splitting along whole length of lobes, orange with a brown margin. Ovary trigonous, shortly pedicellate, green with angles dark purplish-red, lower lobes projecting beyond upper. As fruit ripens, ovary becomes entirely of a deep purplish-red colour. Ovules many, inserted on an axile pink disc in each cell. Style red, grooved at base, grooves corresponding with the 3 sutures at angles of ovary, upper portion with 3 grooves.

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Fig. 1.—Flower of Xeronema Callistemon Oliver.
Fig. 2.—Fruit of Xeronema Callistemon Oliver.

Capsule splitting loculicidally above and below. Base of style also splits into 3 for a short distance. Style persistent, erect; stamens and perianth-leaves withered, persistent.

Seeds numerous, trigonous, black, inner faces smooth, outer face roughened with minute papillae.

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Plant and flower of Xeronema Callistemon Oliver n. sp.

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The present species is referred to Xeronema, a genus hitherto known by a single species, X. Moorei, from New Caledonia. X. Callistemon is distinguished by its much larger size, by the perianth-leaves being alternately wide and narrow, by the leaves being excavated on the inner edge, and especially by the form of the fruit, which is abruptly produced at the angles just below the centre and dehisces above and below (fig. 2). X. Moorei is known to me only by descriptions and figures, but I have little hesitation in describing the New Zealand plant as a distinct though closely allied species of the same genus.

Distribution: Xeronema Callistemon is known only from the Poor-Knights Islets, off the north-east coast of New Zealand. It is there extremely common on exposed rocky faces on the higher portions of the islands. The huge tussocks can easily be seen from the sea. It was not found in the shade of the forest or near the shore.

Time of flowering: At the time of visit a few flowering racemes only were found, unripe fruit was plentiful, and a few heads of ripe capsules were collected.

Relationships: Xeronema now consists of two species, one occurring in New Caledonia, the other in New Zealand. The discovery of X. Callistemon adds another species to an interesting group which allies the flora and fauna of New Zealand to that of New Caledonia. Other genera belonging to this group are Meryta, Rhabdothamnus, Corynocarpus, Agathis, and Knightia among plants, and the land-snail Placostylus.

According to Engler's arrangement (Die nat. Pflanzenf., Teil 2, Abt. 5, 1888, p. 38), Xeronema is a somewhat isolated member of the Liliaceae, forming with Herpolirion the subsection Xeroneminae of the subfamily Asphodeloideae. Herpolirion is a monotypic genus of small herbs confined to the mountains of New Zealand, Tasmania, and south-east Australia. The characters in which these two genera agree are the free perianth-leaves, numerous seeds, and distichous leaves.

Xeronema Callistemon was collected by the Dominion Museum expedition which visited the Poor Knights Islands in December, 1924. Shortly after the party landed, Mr. H. Hamilton brought a specimen into camp, and the following day I came across a large patch near the summit of the island. Later Mr. W. M. Fraser, Whangarei Harbour Engineer, who joined the expedition during its stay in the group, and who knew of the existence of this unusual species, piloted us to where the plant grew more abundantly, and good specimens and photographs were taken.

For the opportunity of comparing specimens of Xeronema Callistemon with Brongniart and Gris's plate of X. Moorei I have to thank the Director of the Sydney Botanie Gardens.

Drimys Forst.

The usage of Wintera for the New Zealand species hitherto referred to Drimys calls for comment. Hutchinson, in the Kew Bulletin, 1921, p. 190, has restricted Wintera to the New Zealand species, and used Drimys for the remaining species of the genus as generally understood. Drimys was founded by J. R. and G. Forster in 1776 (Characteres Genera Plantarum, p. 84) for the two species D. Winteri and D. axillaris. The former belongs to South America, the latter to New Zealand, and both are described and figured in the work mentioned. In 1781 Linné fils (Suppl. plant Syst. veg.) uses Drimys for D. granatensis and D. axillaris. This is practically the same usage as the Forsters', for D. granatensis is synonymous

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with D. Winteri. In 1784 Murray (Syst., ed. 14, p. 507) founded the genus Wintera for W. aromaticum and W. granatensis. This clearly leaves D. axillaris as the type of Drimys. Also, it may be noted, both W. aromaticum and W. granatensis are South American species, and if the New Zealand species are separated from the others of the genus Drimys the name Wintera cannot be applied to them—it must remain with the South American species; and if the course taken by Hutchinson in selecting D. Winteri as the type of Drimys be upheld, Wintera would fall an absolute synonym to Drimys. Hutchinson's action, therefore, appears to be contrary to the principles of nomenclature. Furthermore, Wintera cannot be quoted as of Forster 1786 (Prodr. Fl. Ins. Austr., p. 42), for in the work referred to Forster did not found a new genus, but merely listed Wintera axillaris among the species collected by the expedition during Cook's second voyage.

Myosurus novae-zelandiae sp. nov.

Myosurus aristatus (not Benth.) Hook. f., Fl. Nov. Zel., vol. 1, 1853, p. 8; Cheeseman, Man. N.Z. Flora, 1906, p. 6.

Sepala 5, calcare brevi. Stamina 5. Spica oblonga, acuta, 6–18 mm. longa. Achenia breviore subaristates.

There are apparently several species of Myosurus found in North and South America, mainly on the Rocky Mountains and Andes. The nomenclature of the American species is somewhat involved. The species with which the New Zealand plant has hitherto been united was first described by Gay as Myosurus apetalus. This name was dismissed by Hooker as unsuitable because his specimens bore petals, though he afterwards referred New Zealand examples without petals to the same species. However, the New Zealand plant differs in other respects from the American species, so we are relieved of the necessity of deciding either the correctness or suitability of the name to be applied to it.

Specimens from Awatere River, New Zealand, were compared in the British Museum with the Chilian examples collected by Bridges and used by Hooker to establish Bentham's name aristatus. The Chilian plant was larger, with longer fruiting receptacles and longer beaks to the achenes. In the Kew Herbarium specimens from Lake Tekapo, New Zealand, were compared with specimens from the Rocky Mountains in Missouri and Oregon, collected by Geyer and determined by Hooker as identical with Gay's M. apetalus (Lond. Jour. Bot., vol. 6, 1847, p. 498). In this case the American specimens differed in having long fruiting receptacles, and especially in possessing long beaks to the achenes. The New Zealand species has no petals, whereas Hooker states that all his American specimens possessed them. Some examples both from Chile and the Rocky Mountains approach New Zealand specimens in possessing short fruiting-heads, but in all cases the achenes have long beaks.

Nothopanax kermadecensis n. sp.

Panax arboreum (not Forst.) Cheeseman, Trans. N.Z. Inst., vol. 20, 1888, p. 168.

Nothopanax arboreum (not Seem.) Oliver, Trans. N.Z. Inst., vol. 42, 1910, p. 169.

Affins N. arboreo Seem., sed foliis minute sinuato-serratis, petiolulis breviore, umbellis minor.

A large forest-tree, 10 m. tall, with a trunk over 1 m. in diameter and a large rounded head of foliage. Leaves with a sheathing base and slender petiole; leaflets normally 7 in number, elliptic to obovate-elliptic, produced into an acute point, base usually cuneate and often unequal-sided,

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margin shallowly sinuate-serrate, lower third entire, slightly wavy. Umbels compact, primary rays 3–5 cm. long, secondary rays 1–2 cm. long.

This species is closely allied to N. arboreum of New Zealand, and has hitherto been included with that species; but Dr. Cockayne has recently drawn my attention to its distinctive characters. It differs from N. arboreum in the nearly entire leaves, which are more membranous in texture, and in the compact habit produced by the shorter rays to the umbels and the shorter petiolules. Unlike the New Zealand species, it grows into a large tree.

Distribution: Sunday Island, Kermadec Group; in the damp forest on the tops of the hills.

Colobanthus mollis n. sp.

Colobanthus quitensis (not Bartl.) Hook. f., Handb. N.Z. Flora, 1864, p. 24; Cheeseman, Man. N.Z. Flora, 1906, p. 66.

Planta caespitosa; folia angusto-lineares, acuta, molles; flores tetrameres; peduncules foliis longiores.

After comparison of Chilian examples of Colobanthus quitensis in the Kew Herbarium with New Zealand specimens, hitherto included with the South American species, I advance the New Zealand plant as a distinct species. It is characterized by its tetramerous flowers, short flaccid acute or mucronate leaves not exceeding 15 mm. in length, usually much shorter, and by the peduncles being very short, not extending beyond the leaves. Colobanthus quitensis, which is found along the Andes from Mexico to Chile, and also occurs in Elizabeth Island, is a much larger plant, the leaves being commonly up to 25 mm. in length, while the peduncles are 50 mm. in length, thus far overtopping the leaves.

Carmichaelia arborea (Forster).

Lotus arboreus Forster, Prodr. Fl. Ins. Aust., 1786, p. 52.

Carmichaelia flagelliformis Hook. f., Fl. Nov. Zel., vol. 1, p. 51, 1853.

Forster's drawing is evidently the plant afterwards described by Hooker as C. flagelliformis, and, further, the locality of Lotus arboreus and the full description of Forster's quoted by Richard (Voy. Astrol., Bot., p. 345, 1832) leave little doubt that Forster's name should be applied to Hooker's species.

Metrosideros perforata (Forst.) A. Rich.

Leptospermum perforatum Forst., Char. Gen., 1776, p. 72.

Melaleuca perforata Forst., Prodr. Fl. Ins. Aust., 1786, p. 37.

Metrosideros perforata (Forst.) A. Rich., Fl. Nouv. Zel., p. 334.

Metrosideros scandens Sol. ex Gaertner, Fruct., vol. 1, 1788, p. 172.

Further references are given in Cheeseman's Man. N.Z. Flora, 1906, p. 167.

The types of Forster's Leptospermum perforatum are preserved in the British Museum. They belong to the species generally known as M. scandens; and as the specific name perforata predates all others proposed for this species, and was taken up correctly by Richard, there seems no good reason for disregarding it.

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Descriptions of New Native Plants.

[Read before the Auckland Institute, 25th November, 1924; received by Editor, 28th November, 1924; issued separately, 6th March, 1926.]

Plates 2,3.

Carex McMahoni sp. nov.

Species C. Buchanani (Berggren) similis; differt foliis laete viridibus, longe acuminatis, pendentibus; culmis trigonis laevibus, folia permultum excedentibus; spiculis 5–7 crassioribus, ad culmorum apices approximatis sessilibus, duabus supremis masculis; utriculis minoribus, tenuiter plano-convexis, ellipticis, subalatis, plurinerviis, a parte dimidia superiore delicate serrulatis; rostro brevi bifido serrulato.

Densely tufted, dark green; culms numerous, slender, smooth, trigonous, erect below widely drooping above, 90–110 cm. (3–4 ft.) high; leaves considerably shorter than the culms, narrow-striate, keeled, more or less com plicate below, delicately scabrid along edges, slightly recurved in upper half, produced into long acuminate tips; spikelets 5–7, sessile, closely placed at tops of culms, linear-oblong, cylindrical, rather stout, 3–3.5 cm. (1 ¼ in.) long, 5 mm. broad, the terminal 1 or 2 male (the lower much the smaller and sometimes partially female), the remainder female, a slender and abortive spikelet on a long filiform peduncle often springs from axil of uppermost leaf; bracts long and leaf-like; glumes shorter than utricles, broadly obovate, thin, pale, 1-nerved, wavy and bifid above, produced into a fairly long thin smooth whitish mucro; utricles rather small, shortly stipitate, elliptic, thinly plano-convex, slightly winged along edges, many-nerved on both surfaces, finely serrate along upper half; beak short, bifid, sparsely serrate.

Hab.—Pelorus Valley, Sounds County: J. H. McMahon!

I have pleasure in naming this fine species after Mr. McMahon, who has rendered me many valuable services.

Carex Coxiana sp. nov.

Culmi 45 cm. (18 in.) alti, rigidi erecti triquetri, vix graciles leves, folia vix aequantes; folia linearia, a medio 0.8 cm. (⅖ in.) lata, plerumque plana subcoriacea, utrinque tenuiter ac conferte striata, marginibus sub-revolutis, in apices longiusculos filiformes producta; inflorescentia spicula infima duplo longior; spiculae 6–7 approximatae, sursum gradatim breviores, sessiles vel subsessiles robustae fulvae; infima 3 cm. (1 ½ in.) longa 5 mm. lata, summa mascula claviformis, 3 cm. (1 ⅕ in.) longa, subrobusta; glumae utriculos superantes, ovato-oblongae, plerumque integrae, in aristam brevem levemque productae, apicibus bifidis vel integris; styli 3; utriculi pallidi ± nitentes leves ovoidei, in rostrum subbrevem bifidum angustati, ± plano-convexi, nervis lateralibus duobus percursi, cetera enervia; nux obovoidea, obtuse triquetra, subacuta.

Culms about 18 in. high, stiffly erect, somewhat shorter than leaves, triquetrous, rather slender, smooth; leaves shortly sheathing, almost

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linear, keeled near base, flat for most of their length, moderately coriaceous, 3.6 mm. wide at middle, harsh at margins, finely and closely striate above and below, edges somewhat revolute, drawn out into long filiform tips, midrib conspicuous below, inflorescence twice as long as the lowermost spikelet; spikelets 6 or 7 approximate near tops of culms, the upper gradually shorter, sessile or nearly so, stout, dark brown, the lowermost 3 cm. (1 ⅕ in.) long; topmost male, club-shaped, robust, 3 cm. (1 ⅕ in.) long by 5 mm. wide, remainder female; glumes longer than utricles, ovate-oblong, usually entire, produced into a rather short smooth awn, thin, dark brown with a pale striated band along middle, bifid at tops or entire; styles 3; utricles pale, more or less polished and shining, smooth, obovoid, shortly stipitate, rather sharply narrowed into a moderately short bifid beak with slightly diverging teeth, plano-convex or thinly biconvex, with two distinct lateral ribs, otherwise nerveless or with very obscure nerves; nut somewhat obovoid, bluntly triquetrous, subacute.

Hab.—Chatham Islands: W. Martin!

I have seen only a single specimen, but it is well grown and in good condition. When better known the above description may need amendment.

Carex Martini sp. nov.

Culmi 9–18 dcm. (3–6 ft.) alti, robusti erecti triquetri leves; folia culmos multum superantia, sublate lincaria, a medio 1–1.4 cm. (½ in.) lata, plerumque plana, striata levia, in apices longos filiformes producta; bracteae foliis similes, multo angustiores, culmi florigeri 3–6 dcm. (14 in.) longi; spiculae 8–12 angustae, longitudine variabiles, 5–8 cm. (2–3 in.) longae, saepe aggregatae, pedicellis axillaribus valde inaequalibus; 4 summae masculae, reliquae femineae supra ± masculae; glumae perangustae, ovato-lanceolatae, utriculos multum excedentes, in aristam glabram productae; styli 3; utriculi ovato-elliptici, duplo longiores quam lati, in rostrum gracile sublongum angustati, tenuiter biconvexi, utrinque ± costati, colore viridi cinerascenti maculosi; nux ovoidea.

Culms 3–6 ft. high, stout, erect, sharply and unequally triquetrous, smooth; leaves far exceeding culms, long-sheathing, linear, 1–1.4 cm. (½ in.) wide at middle, keeled at and near base, flat for most of their length, smooth, coarsely striated above, more finely and closely so beneath, produced into long gently narrowed filiform tips, midrib little conspicuous above, edges smooth, bracts like the leaves but much narrower; flowering culm 3.6 dcm. (14 in.) long; spikelets 8–12 (counting the aggregated ones separately), narrow, 5–8 cm. (2–3 in.) long, variable in length; peduncles nearly as long as spikelets, several just exserted from sheaths, frequently aggregated when pedicels vary greatly in length, the shorter spikelets having shorter pedicels; topmost 4 (rarely compound) male, the others female with a considerable tuft of males at top of each, the lowermost distant; glumes very narrow, ovate-lanceolate, longer than utricles, continued into a smooth pale awn, entire or wavy at edges, midrib inconspicuous; utricles ovate-elliptic, twice as long as wide, shortly stipitate, thinly biconvex, two lateral nerves prominent with several finer ones on both faces, narrowed into a moderately long slender straight beak not or scarcely bifid, mottled with greenish-grey; nut ovoid.

Hab.—Wet and swampy stations at Chatham Islands: W. Martin!

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This is a very fine plant. Its range in height may be even greater than here stated. In any case, 6 ft. is the extreme height observed, and such specimens are probably very uncommon. The tallest plant seen grew near the edge of a lagoon and in water of some depth.

Carex McClurgii sp. nov.

Culmi 6–8 cm. (2 ¼–3 in.) alti, graciles triquetri leves, folia paullo superantes; folia angusta, a medio 5.2 mm. lata, prope basim carinata vel ± complicata, a parte superiore plana, tenuia, levia striata, a marginibus subscaberula, in apices longos filiformes producta; spiculae ad 8; duae ultimae perdistantes, tenuiter pedicellatae, reliquae a culmo summo approximatae, plerumque sessiles; suprema mascula 3–6 cm. longa, reliquae femineae sed ab apicibus ± masculae; glumae utriculos aequantes, tenues acutae pallidae, ovato-lanceolateae, vix aristatae, ab apicibus ± laceratae; styli 3; utriculi anguste elliptici, breviter stipitati, biconvexi vel ± subtriangulares, nervis duobus firmis ac compluribus subtilioribus percursi; rostrum sensim angustatum in apicem acutum breviter vel vix bidentatum.

Culms 6–8 cm. (2 ¼ in.) high, slender, triquetrous, smooth, somewhat exceeding the leaves; leaves narrow, 5.2 mm. wide at middle, below keeled or more or less complicate, for the most part flat, thin, smooth, slightly harsh along margins, striate, drawn out into long filiform tips; midrib prominent below; spikelets 8, 3–6 cm. (1 ¼ in.) long, rather thin, the two lowermost far apart on long delicate largely ensheathed peduncles, the remainder closely placed near top of culm; topmost male, rather long, the rest female with some male flowers at tops, sessile or nearly so; bracts like the leaves and as long; glumes about equalling utricles, very thin, ovate-lanceolate, acute, pale straw-coloured, awnless or nearly so, more or less lacerate at tops; styles 3; utricles narrow elliptic, shortly stipitate, biconvex or subtriangular, with two strong lateral nerves and numerous finer ones; rostrum gently narrowed to the acute shortly or barely bidentate beak; nut oblong, obtusely trigonous.

Hab.—Chatham Islands: W. Martin! January, 1924.

Mr. Martin obtained a great deal of important help and information about the island plants from Mr. McClurg, who knows almost all the native plants, with their Moriori names.

Carex rekohu sp. nov.

Culmi ad 54 cm. (20 in.) alti, obtuse trigoni leves; folia culmos aequantia, a medio c. 10 mm. lata, coriacea vel subcoriacea, pro parte majore complicata et a marginibus recurva, raro apicem versus plana, in apices filiformes angustata; bracteae foliis consimiles ac aequilongae. Spiculae ad 10, 4 supremae masculae sessiles valde approximatae, sub-crassae; reliquae foemineae saepe compositae a parte terminali ± masculae, infima 6.5 cm. (2 ½ in.) longa, longe pedunculata, crassior distans, superiorum pedunculi breviores, 2 vel 3 summae sessiles; glumae utriculos aequantes, alte bifidi, nervo medio pallido percursae, costa media in aristam hispidam pallidam producta; utriculi anguste obcuneati, rostrum subconicum brevissime bifidum, nervi laterales duo validi, caeteri obscuri et saepe paene obsoleti. Nux obtusa trigona anguste oblonga.

Culms up to 54 cm. (20 in.) high, obtusely trigonous, smooth, moderately stout; leaves about as long as culms, c. 10 mm. wide at middle, gradually tapering to filiform tips, complicate and more or less recurved at

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edges, rarely flat in terminal third, smooth deeply striate, midrib prominent below; bracts like the leaves and as long, the lowermost long sheathing. Spikelets usually 10, the four topmost male, sessile, very closely placed, rather stout, the others female with some male flowers at tips, lowermost 6.5 cm. (2 ½ in.) long, long pedunculate, stout, distant, those above with shorter pedicels, the two or three topmost sessile; glumes about equalling utricles, rather narrow-oblong, bifid above, with one strong median nerve a dark brown strip on either side with pale margins, midrib prolonged into a pale hispid awn as long as glumes. Styles 3. Utricles narrow obcuneate, stipitate, sharply contracted at tip into a conical very shortly bifid beak with very short erect teeth, polished and when mature almost black, with two strong lateral nerves and several faint and almost obsolete nervures between. Nut bluntly trigonous, narrow-oblong.

Hab.—Chatham Islands: W. Martin!

The specimens collected January, 1924, were much overripe, and most of the glumes and utricles had fallen off.

Carex rotoensis sp. nov.

Calmi 15 cm. (6 in.) longi, teretes graciliores leves; folia culmos paene aequantia, linearia, infra ± complicata plerumque plana, striata, a medio 3 mm. lata, marginibus apices versus tenuiter scabridis; spiculae 6, distantes sessiles, c. 5 mm. (¾–¼ in.) longae, sursum regulariter abbreviatae, pallidae, ultima breviter pedicellata; summa mascula 11 cm. (1 ¼ in.) longa, reliquae femineae; glumae utriculos superantes, ovato-lanceolatae acutae, plerumque integrae, a medio pallidae a marginibus subfuscae vix mucronatae; styli 3; utriculi plano-convexi vel subtriquetri, haud vel vix stipitati, ovoidei politi, enerves vel nervis perobscuris; rostrum moderate breve sublate bifidum; nux triqueter.

Culms 6 in. long, terete, rather slender, smooth; leaves numerous, about as long as culms, the outermost short and scale-like, acute or acuminate, strongly striate; older ones linear complicate or keeled below, flat for most of their length, 3 mm. broad at middle, finely scabrid along edges of upper part, narrowed into fine points; spikelets 6, distantly placed save at tops of culms, sessile or the lowermost shortly pedicellate, gradually shorter upwards, ¼–¾ in. long; topmost male, 1 ¼ in. long, the others female; glumes longer than utricles, ovate-lanceolate, thin and membranous, usually entire rarely shortly bifid at or near tips, pale along the middle, more or less brown at sides, midrib prominent, scarcely mucronate; styles 3; utricles plano-convex or subtriquetrous, not or hardly stipitate, ovate, polished, nerveless or almost so; beak moderately short rather widely bifid; nut triquetrous.

Hab.—Te Roto, Chatham Islands: W. Martin!

Only a single specimen of this was available for examination, but it was in good condition. Further specimens may show occasion for amendments in the description.

Schoenus caespitans sp. nov.

Species dense caesptans depressa c. 4 cm. (1–1 ½ in.) alta. Folia a bas erecta, filiformia, culmos aequantia vel excedentia, basi ± expansa ac vaginantia; culmi foliis crassiores virides erecti; inflorescentia brevis capituliformis; capitulum parvum sublonge bracteatum, e fasciculis tribus 8–10 florigeris arcte aggregatis compositum; glumae distichae, 3 inferiores

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vacuae, quarta florem hermaphroditum gerens, summa vacua raro staminifera; stamina 3; setae plerumque 3; stylus erectus supra in ramos 3 divaricantes divisus; spiculae culmique maturi haud visi.

A densely-tufted depressed species 4 cm. (1 ½ in.) high, forming low patches of considerable size. Leaves numerous, filiform, glabrous, more or less involute, delicately scabrid at tips, equalling or exceeding culms, expanded below and more or less sheathing; culms twice as stout as leaves, erect, green; inflorescence compact, dark brown, composed of 2–3 subumbellate closely-placed clusters of 8–10 flowers; cluster-bracts subulate, long, green much exceeding flower-clusters; spikelets small, lanceolate, nearly sessile; glumes distichous, ovate, acute, dark brown with whitish edges and a whitish median tract along back, three lowermost empty, the fourth with a perfect flower, the fifth empty rarely staminiferous; bristles usually 3; stamens 3, sagittate; style erect with 3 strongly divaricating branches; mature culms and spikelets not seen.

Sab.—Ure Valley, Marlborough: Arnold Wall!

The culms probably elongate after flowering. One of the most interesting of Professor Wall's recent discoveries.

Muehlenbeckia debilis sp. nov.

Species M. complexae (Meissner) affinis; differt caulibus gracilioribus viridibus vel cinerascenti-viridibus, ubique glaberrimis, ramis ramulisque distantibus subdivaricatis pergracilibus, foliis multo angustioribus plerumque lineari-lanceolatis acuminatisque, floribus in spicis simplicibus dispesitis, periantho in fructu maturo herbaceo haud incrassato.

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A branching perfectly glabrous shrub, forming low widespreading more or less entangled scrambling sheets in moist stations, or broad erect and finally pendulous dense rush-like tufts on gravelly terraces (where they are usually much eaten back by stock). Stems up to 15 dcm. (38 in.) long in moist stations, much shorter on gravelly terraces, slender, tough, terete, striate, green or greyish-green, quite glabrous, giving off (in the scrambling form) numerous distant very slender alternate subdivaricating branches and branchlets, with rather long simple ultimate flower-bearing twigs; leaves chiefly on ultimate twigs, more or less distant, usually linear-lanceolate and acuminate, entire, 10–20 mm. (7 13/16 in.) long (rarely shorter and contracted about the middle with rather wide basal lobes), suddenly narrowed into slender petioles about ⅓ as long as the blades; sheathing stipules conspicuous, thin, yellowish, more or less persistent. Spikes axillary and terminal, simple, narrow, 3–4 cm. (1 ½ in.) long in male plants, shorter in female ones; flowers closely placed, alternate, almost sessile in axils of broad subacute yellowish sheathing scales, small, dioecious; males with 8 stamens and perianth deeply cut into semihyaline narrow-oblong obtuse divisions; females with somewhat broader perianth-lobes, 8 staminodia and sparingly fimbriate stigmas; fruiting perianth not enlarged or succulent; nut black and shining, deeply triquetrous.

Hab.—On low gravelly terraces and the immediate banks and rocky faces of the Awatere and Grey Rivers, close to the Homestead of Upcot Station, Middle Awatere, Marlborough.

Flowers in February. Male plants seem to be much more plentiful than female ones. This may be due to the general absence of flowers on the plants scattered over the gravelly terraces, most of which are eaten back to below the level at which the inflorescence appears.

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Ranunculus Carsei sp. nov.

Speciés parva prostrata sparse pilosa; culmi complures pro plantae mangitudine robustiores; petioli graciles ± 1.7 cm. longi, basi vaginantes, vaginis copiose pilosis; laminae 6 mm. latae 5 mm. altae, laeves, forma variabiles, plerumque late triangulares 3-lobatae, lobis varie divisis; flores in anthesi parvi axillares sessiles, maturorum pedicellis ad 2 cm. productis; sepala 5 laevia 1-nervia, scariosa ovata; petala pauca (2 vel 3 ut videtur), longiuscula anguste obcuneata; stamina pauca; achenia parva late obovata, biconvexa, ± atro-fusca; rostrum perbreve erectum.

A small prostrate sparsely-pilose plant growing on dripping rocks or similar wet situations. Roots numerous, slender; stems several, short, c. 5 cm. (2 in.) long, rather stout for size of plant; leaves on slender petioles 1.7 cm. (⅝ in.) long, petioles expanded and sheathing at base, sheaths copiously pilose; blades 6 mm. broad, 5 mm. long, thin, variable in cutting, general outline broadly triangular usually deeply 3-lobed, lobes more or less subdivided, terminal lobe sometimes cut like the main lamina; flowers small, axillary, at anthesis sessile or nearly so, in fruit with peduncles longer than the petioles; sepals 5, thin, glabrous, scarious, 1-nerved, ovate, somewhat boat-shaped; petals few (apparently 2–3), rather long narrow-obcuneate, upper part bright yellow; stamens few, anthers small and terminating filaments; achenes small, broadly ovate biconvex, upper parts more or less mottled with blackish brown; beak straight, inconspicuous.

Hab.—Mounts Tongariro, near the base: H. Carse!

Flowers in January. A very peculiar and distinct species. It has no close ally in the New Zealand species of the genus known to me.

Notothlaspi rosulatum (Hk. f.) var. Hursthousei var. nov.

Planta c. 22 cm. (8–9 in.) alta, purpurata, quam forma typica altior ac robustior, a basi inflorescentiae emittens supra folia c. 12 graciles breviores paucifloros ramos; flores subflavidi, petalis purpureo ± tinctis.

Plant 8–9 in. high, everywhere purplish, longer and stouter than the typical form; radical leaves very numerous, long-petiolate; from base of the ordinary terminal inflorescence about 12 slender rather few-flowered branches spread out just above leaves; main axis of inflorescence 6.7 in. long, erect, flowers and capsules closely crowded throughout its length; flowers cream-coloured, tips of petals tinged with purple; capsules larger than in type.

Hab.—Shingly Range near Molesworth Station, Upper Awatere, Marlborough: F. H. Hursthouse!

Only a single specimen of this remarkable form has been seen by me, and when better known it may prove worthy of specific rank. Mr. Hursthouse, who is an excellent observer, writes: “It grows at a height of about 4,000 ft. on shingly faces that are visible from Molesworth Homestead. There were many fine specimens, like the one I have sent, which stood up like little manikins on the bare shingle.” Further specimens will be awaited with interest.

Pimelea laevigata Gaert. var. monticola var. nov.

Rami numerosi a radicibus quoquoversus sublate patentes, graciles, decumbentes, haud vel vix radicantes; ramulis multum subdivisis, pilis

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sericeis brevibus subdense vestitis; folia ± patentia, oblongo-elliptica, sub-acuta, paene sessilia, glabra, latitudine quam longitudine dimidio minore, costa media infra conspicua; flores colore subpuniceo ± tincti.

Branches numerous, spreading widely all round from top of roots, slender decumbent or almost prostrate, glabrous in lower parts; branchlets much subdivided and clothed rather closely with short silky hairs; leaves smooth, patent, oblong-elliptic subacute, almost sessile, glabrous, about ½ as wide as long; midrib conspicuous on under side; flowers stained with pink, floral leaves slightly larger than cauline.

Hab.—Hauhungatahi, Waimarino Plateau, c. 3,800 ft.: H. Carse and H. B. Matthews! Mount Tongariro: B. C. Aston!

Originally sent me by Mr. B. C. Aston, who considered it a new species. It is, however, very close to Gaertner's species, and is for the present ranked only as a variety. It is always a plant of more open growth with leaves much less glossy than in the type.

Nothopanax McIntyrei sp. nov.

Arbor subhumilis ± glaber paene a basi fastigiate ramosus. Folia alterna, coriacea, satis distantia, trifoliolata; petioli ad 3.5 cm. (1 ½ in.) longi; foliola petiolos breviter superantia, sessilia, elliptica, laete viridia supra, infra pallidiora, ad basim gradatim attenuata, acute ac subconferte serrata, subacuta, costa media evidens supra infraque. Inflorescentia umbellata, axillaris (rarius terminalis); rami principales plerumque solitarii erecti graciles, in ramos secundarios complures divaricantes 2 cm. (¾ in.) longos divisi, denuo subdivisi in umbellulas minores 18 mm. (¾ in.) latas, flores parvos breviter pedicellatos 6–8 gerentes. Styli 2; fructus maturus suborbicularis, ± complanatus, c. 5 mm. diam.

A rather glabrous small tree, fastigiately branched from near the ground. Bark blackish-brown; leaves alternate, rather distant, trifoliolate, dark green above paler below; petioles about 3.5 cm. (1 ½ in.) long (sheaths short stem-clasping); leaflets coriaceous, slightly longer than petioles, sessile or nearly so, elliptic, gradually narrowed to base, sharply and coarsely serrate for ⅔ their length, subacute, midrib evident above and below, nerves obscure. Inflorescence umbellate in axils of upper leaves, more rarely terminal, dioecious; primary branch usually solitary and erect, divided into several (6 or fewer) strongly divaricating secondary branches, which are again subdivided into smaller umbellules bearing 6–8 shortly pedicellate flowers. Styles 2, short, slender; male flowers not seen; mature fruits suborbicular flattened, 5 mm. in diameter.

Hab.—Grown in Dr. Hunter's garden at Mornington, Dunedin, and said to have come from Westland: W. A. Thomson.

Mr. Thomson has transmitted specimens of this to me. The species is named in honour of Mr. J. W. McIntyre, who has had charge of the above garden for many years, and has had great success in growing alpine and other rare plants sent on from time to time by Mr. H. J. Matthews, by whom this fine garden of native plants was established. The pieces seen all belong to the female plant. It has been in the garden nineteen years, and is now a small tree about 8 ft. high. The trunk just above the ground is 15 in. in circumference. Several young plants have grown up near its base.

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Aciphylla Crosby-Smithii Petrie

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Aciphylla Crosby-Smithii Petrie

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Aciphylla Crosby-Smithii Petrie. (Plates 2,3.)

A preliminary notice of this species was published in Trans. N.Z. Inst., vol. 47, p. 48. Thanks to the kindness of Mr. James Speden, of Gore, and Mr. H. L. Darton, of Lawrence, a full-grown plant (not in flower) has been obtained, which is shown herewith in two photographs of different scales. It is thus possible to add some important particulars to the original notice. The natural size of the whole plant is 14 in. by 11 in., and each of the six rosettes is 5 in. in diameter.

Stem erect from root, nearly as thick as a man's thumb, short (c. 8 in.); main branches 3 approximate, the lowermost about 2 in. up stem, subdivided above and nearly as stout as stem, closely covered with more or less withered leaves and ending in large rather flattened or rounded rosettes or heads each 4 in. or 5 in. across, and covered with very numerous most densely compacted leaves. Leaves spreading all round, about 3 in. long, the lower half broadly sheathing, sheaths very thin and glabrous, basal spines like the leaflets; leaflets in 3 to 5 pairs, linear-oblong sub-acute with a short abrupt pungent mucro, channelled above, 1-nerved, thickened at the margins. Flowering-stem rather thin (c. 5 mm. in diameter), about 8 in. high, brownish-red, deeply striate; involucral scales thin and flaccid, sheaths longer than leaf-like part; primary umbels numerous, slender, about 1 in. long; secondary small with many almost filiform pedicels; fruits all shed and no flowers seen.

Hab.—Princess Range, Fiord County, c. 4,600 ft.: James Speden! H. L. Darton!

A most remarkable plant, as the photographs sufficiently show. It probably flowers but rarely. No known native Aciphylla resembles it in habit of growth. The old flowering-stems came from other plants than those shown in the illustrations.

Schizeilema Allanii sp. nov.

Species S. pallidae (Domin) consimilis; differt rhizomate longiore, foliis multo majoribus semiorbicularibus infra pallidis; petioles multo longioribus a marginibus subcartilagineis; umbellis plerumque duabus raro singulis vel tribus; fructibus majoribus, foliis involucratibus apicem versus ± dilatatis.

A matted pale-green perfectly glabrous plant very similar to S. pallida (Domin); rhizome up to 5.3 cm. (12 in.) long, slender, flaccid, with 1 to 3 distant rooting-nodes generally more or less leafy; radical leaves not numerous, petioles rather slender flaccid brownish, up to 6 in. long; blades semiorbicular in outline, 3.8 cm. (1 ¼ in.) wide, ¾ in. long, more or less polished, rather pale green above much lighter below, little coriaceous, variable in cutting, usually trifoliolate but sometimes only more or less deeply trifid or tripartite at tips, segments obcuneate usually with 3 to 4 shallow rather broad subcrenate indentations at tips, margins thinly cartilaginous, slightly incurved when dried, veins obscure above evident below; peduncles rather distant, almost filiform, shorter than petioles, the primary at first terminal, later overtooped by a decumbent branch from the axil of a small entire or trifid bract, the same formation being repeated a second or even a third time, secondary umbellules occasionally present; peduncles short very slender, pedicels nearly as long capillary; involucral leaves linear somewhat expanded near tips; flowers 8–12 in

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each umbel; fruits (still rather immature) shortly and broadly oblong, obtusely 4-angled, deeply furrowed at commisures.

Hab.—Maharahara Mountain, near south end of Ruahine Range: H. H. Allan!

I have to thank Dr. Allan for a good series of specimens.

Gentiana Spedeni sp. nov.

Planta perennis erecta glaberrima, culmos solitarios longiusculos gerens; folia radicalia numerosa, conferta, arcte imbricantia, subrosulata spathulata integra, 1 cm. (¼–⅝ in.) longa; pars petiolaris quam lamina bis terve longior, lamina ovata subacuta, parum coriacea, integra, nervo medio evidente supra infraque, cetera enervia; folia caulina 8–10 mm. longa in paribus duobus distantibus disposita, pari superiore ⅓ culmi longitudinem aequante; culmus simplex teres pergracilis, 14 cm. (5 ¼ in.) longus; bracteae florales 4, sessiles ovatae congestae; flores 4 pedunculati, pedunculi 7 mm. (c. 1 ½ in.) longi, ex bractearum axillis editi. Calyx alte in lobos 5 angustos acutos sectus; petala 5 calyce ⅓ longiora, late obovata obtusa, venis subparallelis purpureis percursa; pistillum apice 2-lobatum. Capsula haud visa.

A slender glabrous plant with solitary culms, numerous subrosulate spathulate radical leaves, and terminal umbel-like clusters of 3 or 4 flowers. Base of stem nearly horizontal, slender, naked, closely scarred, dark brown; radical leaves crowded, overlapping, entire, 1 cm. (¼–⅝ in.) long; petiolar part 2–3 times as long as the laminar, flattened; lamina ovate, subacute, little coriaceous, entire, nerveless but for the evident midrib; cauline leaves in 2 pairs, the lower a little above the radical, the second ⅓ way up the stem, small, subsessile; stem simple, terete, very slender, 14 cm. (5 ¼ in.) high; bracts 4, crowded, obovate, sessile, each subtending a flower; flowers 4, forming a terminal cluster 1.5 cm. (c. ½ in.) long; peduncles almost capillary, about equalling the flowers; calyx cut almost to the base into 5 broadly-linear acute segments; petals ⅓ longer than calyx, broadly obovate, traversed by numerous subparallel “purplish or violet” (J. Speden) veins; stamens ¼ length of pistil, filaments about equalling anthers; pistil shortly two-lobed at top.

Hab.—Princess Range, Fiord County, c. 4,000 ft.: James Speden!

Early January, 1924; rare. Only a single plant of this very beautiful and distinct species has been seen. Mr. Speden reports that only a few plants were observed, and that flowering ones were extremely rare. It would probably make a fine garden-plant.

Gentiana amabilis sp. nov.

Herba nana, ad 4–5 cm. (2 ¾ in.) alta, perennis (?), in locis palustribus crescens. Caules solitarii, a radice erecti vel ± flexuosi, flores singulos terminales gerentes; folia radicalia subspathulata; laminae 1.5 cm. (⅝ in.) longae 6 mm. latae, anguste ellipticae, tenues subacutae, in petiolos ± aequilongos ± complanatos angustatae; caulina sessilia, in paribus duobus subdistantibus disposita; flores magni albi, 1.8 cm. (¾ in.) longitudine ac latitudine; calyx corolla dimidio brevior; corolla alte secta, lobis late obovatis; stamina corolla dimidio breviora, antheris ± complanatis ac basim versus dilatatis; pistillum in anthesi staminis aequilongum; stigma breviter bifidum.

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A dwarf herb 4.5 cm. high, perennial (?), growing in boggy ground. Stems solitary, slender, dark brown, erect or more or less flexuous, often laterally placed through rapid growth of a side shoot (rarely once divided and bearing 2 flowers); radical leaves subspathulate, with narrow more or less elliptic thin acute entire blades, 1.5 cm. (⅝ in.) long and 6 mm. wide, and somewhat flattened slightly expanded at base; petioles equalling blades or shorter; cauline in 2 rather distant pairs, narrower and smaller, sessile or nearly so; flowers solitary, terminal, large for so small a plant, white, 1.8 cm. (¾ in.) long and as wide; calyx scarcely half as long as corolla, cut half-way down into broad-based acute lobes; corolla deeply divided, segments broadly obovate; stamens half as long as corolla; anthers flattened and expanding below; pistil about equalling stamens; stigma shortly 2-lobed.

Hab.—Bogs at the top of Mount Tennyson, Garvie Range, Southland, 4,800 ft.: W. A. Thomson!

Mr. Thomson has supplied a good series of specimens, which are most uniform in their characters.

× Veronica Bishopiana species hybrida nova. (V. salicifolia Forst. × V. obtusata Cheeseman.)

Frutex humilis habitus aperti; caules c. 60 cm. alti, infra secundum solum patentes ac saepe a nodis radicantes, deinde ascendentes vel ± erecti, graciles apices versus purpurei ± ramosi; folia decussata anguste lanceolato-elliptica, 5–8 cm. (3 in.) longa c. 2 cm. (⅘ in.) lata, glabra sessilia acuta; racemi e foliorum superiorum axillis, infra nudi, graciles glabri, 8–11 cm. (4 in.) longi; flores dense congesti, breviter pedicellati; stamina stylique longe exserti corollam duplo superantes; capsulae subparvae deflexae ovoideo-ellipticae.

A straggling low shrub of open habit growing on bare or mossy rocks. Stems about 2 ft. high, several spreading more or less closely to the ground and often rooting at nodes, then ascending or nearly erect, slender, glabrous, dark purple, considerably branched near extremities; branches spreading at a wide angle, not further subdivided; leaves decussate, moderately distant, narrow lanceolate-elliptic, 2–3 in. long, ½–¾ in. broad, glabrous, sessile, sharply acute; racemes in the axils of the upper leaves, naked below slender glabrous, 3–4 ½ in. long, about ½ in. broad; flowers closely crowded, shortly pedicellate, rather small; stamens and styles strongly exserted, twice as long as corolla; capsules rather small, deflexed, ovoid-elliptic.

Hab.—Rocky knobs between Huia Hill and Little Huia, near Manukau North Heads: J. J. Bishop! H. Carse! E. Jenkins! April, 1924.

Mr. Bishop has had this plant in cultivation for several years, having transferred young wild plants to his garden, where its position and relations have hitherto puzzled observers. Both the parent species grow on the coast west of the Waitakerei Range as far as Manukau Heads, V. salicifolia being much the more plentiful. V. obtusata, in this district, grows on coastal cliffs and on rocky hummocks in the forest often some way back from the sea.

Euphrasia Hectori sp. nov.

Rami complures a radice patentes ac radicantes, 8 cm. (3 in.) longi vel ultra, ramulos complures laterales breves e foliorum axillis edentes;

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folia in paribus oppositis disposita, ± approximata punctata, anguste obovata, subspathulata, connata, culmos ± vaginantia, sub apice obtuse breviterque 3-dentata, obtusa; culmus florigerus ascendens, deinde erectus; bracteae sessiles, in paribus duobus distantibus dispositae, subtriangulares apicibus breviter 3-lobae; flores majusculi, solitarii, ex superiorum bractearum axillis orientes; pedicellis capillaribus simplicibus, ± pendulis; calyx corollae tubo lato ¼ brevior, ad medium in segmenta obtusa 4 secta; corollae labium superius alte 3-loba, inferius fere ad basim 2-laba, lobis obtusis; capsula anguste ovata, acuta; pistillum capsulam subaequans.

Branches several from the root, spreading and rooting, 8 cm. (3 in.) long or more, light brown, terete, glabrous, giving off short lateral branches from axils of leaves and terminating in a slender ascending or erect inflorescence; leaves in opposite pairs 1.5 cm. (½ in.) apart or less, narrow obovate-spathulate, connate, and stem-clasping at base; petiolar part thin, rather broad, flat or slightly incurved at scantily ciliate edges, about ½ as long as entire leaf; laminar part thicker, narrow obovate with 2 shallow rounded teeth near obtuse tip, punctate on both surfaces, glabrous, vivid green above, paler below, the edges slightly recurved; midrib evident above and below; flowering-stems erect or ascending from the ends of branches, very slender, dark brown; cauline leaves in 3–4 distant pairs, sessile or the lowermost nearly so, coriaceous, subtriangular shortly 3-lobed at tips; flowers large, springing singly from axils of uppermost bracts, pedicels capillary about 2.5 cm. (1 in.) long, dark, simple, drooping more or less; calyx about ¾ as long as the broad corolla-tube, cut halfway down into 4 oblong obtuse segments; corolla white, large, 2 cm. (¾ in.) long and nearly as broad, upper lip deeply 3-lobed, lobes obtuse, lower 2-lobed to base, lobes obtuse; capsules (rather immature) as long as sepals, narrow ovate acute; pistil about equalling capsule; seeds not seen.

Hab.—Haast Valley, South Westland: R. A. Wilson!

The specimen described is incomplete, but the characters above set forth show it to be very distinct. The specimen was forwarded to me by Mr. B. C. Aston, F.N.Z.Inst.

Petalochilus: a New Genus of New Zealand Orchids.

[Reprinted from the Journal of Botany, March, 1924, on the suggestion of D. Petrie.]

Plate 4.

Petalochilus nov. gen.

Herbae terrestres, hirsutae, tuberibus globulosis parvis, caule simplici 1-foliato. Folium ad basin, lineare. Flores pedicellati, 1 vel 2. Segmenta perianthii subaequilonga, fere similia, angustiuscula; posticum erectum vel leviter incurvum, cetera plana et patentia. Labellum ceteris segmentis simile sed breviusculum. Columna elongata, leviter incurva; in parte superiore late alata; inferiore anguste alata. Anthera suberecta,

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Figs. 1–3.—P. calyciformis. Figs. 4–7.—P. saccatus.
1, Column from the side, showing appendage; 2, front view of the flower (the artist has represented the labellar segment too long); 3, column from the front, showing appendage; 4, column from side, showing anterior union of the columnar wings; 5, front view of flower (the artist has shown the labellar segment too long); 6, column from the front, showing the union of the wings to form a pouch; 7, column from the back. All details much enlarged.

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bilocularis, mucronata; pollinia pulverea; caudiculae nullae. Stigma sub antheram, concavum.

Species nobis notae 2, Novae Zelandiae incolae.

Leaf basal, linear. Flowers pedicellated, 1 or 2. Segments of perianth subequal, similar, rather narrow; the dorsal one erect or slightly incurved over the column, the others flat and spreading; labellum similar to the other segments but slightly shorter. Column elongated, slightly incurved, widely winged above, narrowly below. Anther suberect, 2-celled, mucronate; pollinia powdery; caudicles absent. Stigma concave, just below the anther.

Terrestrial hairy herbs, with small globular tubers.

The outstanding feature of the new genus is, of course, the ancestral form of the labellum.

The excellent vernacular names, cup and pouch orchid, originally applied by the discoverer, Mr. H. B. Matthews, have been latinized and retained as specific designations for its two representatives.

The plants are endemic to New Zealand, and, so far as is known, are restricted to the neighbourhood of the little town of Kaitaia, situated in the extreme north-east of the Dominion.

The peculiar appendage in the first species is probably staminodial in origin, and occupies the position of stamen a3 of the inner whorl. In P. saccatus the size and shape of the capacious pouch suggest that the wings of the column are not its sole constituents, but that the staminode is also a component part.

Superficially both species bear a striking resemblance to the diminutive orchid Caladenia minor Hook. f., which is likewise endemic to the same islands.

The affinities of the new genus are certainly with Caladenia rather than with Thelymitra, with which it has few features in common beyond the hitherto unique distinction of an undifferentiated labellum. The Apostasieae, in which, of course, the labellum is also undifferentiated, must be regarded only as doubtful members of the Orchidaceae. It also approaches very closely to Glossodia, especially if it be admitted that the pouch in P. saccatus represents a fusion between the wings of the column and the staminode. In the living state, however, there is little that is reminiscent of that genus in which the labellum, although almost quite plain, is nevertheless very dissimilar in appearance to the petals, and the basal appendage is either bifid or distinctly dual in character.

Mr. Matthews has had these orchids under observation since the year 1912. He says that both species were plentiful, that they seeded freely, and showed practically no variations. On one occasion he collected about a hundred specimens of P. calyciformis and seventy of P. saccatus. In January, 1919, he wrote stating that he had recently visited Kaitaia, and found that in all three places where previously he had been accustomed to collect the cup-orchid, the tea-tree (Leptospermum scoparium), among which it grew, had been cleared for agricultural purposes. Thus, unless a fresh locality should be discovered, this interesting plant will be no longer available to students of botany.

Column-wings open anteriorly, not connate; a linear sigmoid appendage, with a cup-shaped summit, erect against the column 1. P. calyciformis.
Column-wings connate anteriorly throughout their entire length, forming a pouch 2. P. saccatus.
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1. P. calyciformis n. sp.

Gracillima, circiter 7–22 cm. alta. Folium anguste lineare, fere glabrum, circiter 4–15 cm. longum. Caulis gracillimus, hirsutus, infra vel ad medium bractea lineari—lanceolata instructus. Flores 1 vel 2, subvirides, circiter 12–13 mm. in diametro. Ovarium elongatum subvillosum. Segmenta perianthii subacuta, 5–nervosa, circiter 6 mm. longa; labellum ceteris segmentis breviusculum et aliquando latiusculum. Columna circiter 5 mm. longa, in parte superiore late alata. Appendix longiuscula linearis sigmoidea, apice calyculo instructa, ante columnam erecta.

N.Z. : Kaitaia, County Mongonui, H. B. Matthews, 27 Oct.–15 Nov., 1916.

A very slender plant, about 7–22 cm. high. Leaf very narrow linear, nearly glabrous, basal, from 4–15 cm. long. Stem very slender, hairy, a loose linear-lanceolate bract at, or a little below, the middle. Flowers greenish, about 12–13 mm. in diameter, usually single, but occasionally 2, the very slender pedicel subtended by a narrow acute bract. Ovary elongated, rather hairy. Segments of the perianth not very acute, pubescent-glandular on the outside, 5-nerved; the dorsal one erect or slightly incurved, the others spreading; about 6 mm. long; the labellar segment a little shorter than the rest and sometimes a little wider. Column about 5 mm. high; winged throughout, rather widely in the upper half, narrowly below. A linear appendage with sigmoid flexure, furnished with a little cup at the apex, erect in front of the column.

2. P. saccatus n. sp.

Gracillima, circiter 7–14 cm. alta. Folium anguste lineare, fere glabrum, cauli subaequilongum. Caulis hirsutus, supra medium bractea acuta instructus. Flos solitarius, carneus, circiter 2 cm. in diametro. Ovarium elongatum, villosum. Segmenta perianthii subacuta, 5-nervosa, circiter 10 mm. longa; labellum breviusculum et aliquando latiusculum. Columna circiter 4.75 mm. longa; alae antice connatae, saccum membranaceum formantes.

A very slender species, about 7–14 cm. high. Leaf almost glabrous, narrowly linear, usually about as long as the stem. Stem very slender, hairy, with an acute bract above the middle. Flower pink, solitary, about 2 cm. in diameter, its pedicel subtended by a narrow acute bract. Ovary elongated, rather hairy. Segments of perianth not very acute, the sepals glandular-pubescent on the outside, 5-nerved, a pink stripe down the middle; the dorsal one erect or slightly incurved, the others spreading; about 10 mm. long; the labellar segment a little shorter than the rest and sometimes a little wider. Column about 4.75 mm. high with transverse pink bars; the wings uniting behind the anther, coalescing throughout anteriorly, so as to form a well-marked membranous pouch below the stigma.

N.Z.: Kaitaia, Mr. H. B. Matthews, 10–31 Oct., 1917.

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A Proposed New Botanical District for the New Zealand Region.

[Read before the Philosophical Institute of Canterbury, 12th November, 1924; received by Editor, 31st December, 1924; issued separately, 6th March, 1926.]

In 1921 (The Vegetation of New Zealand, p. 303) L. Cockayne extended the boundaries of his Marlborough-Sounds Subdistrict southwards and westwards, and changed the name to “Sounds-Wairau.” As defined by that author, the area was bounded on the south by the River Wairau, and on the west by the average limit of the westerly rainfall. Although this area in that portion of its coastal-lowland-montane belt exposed to a forest-climate may rightly be united to the North Island part of the Ruahine-Cook Botanical District, yet its mountainous portion above the forest-line is markedly different, containing as it does so many species absent on the Ruahine-Tararua Range. It seems to us better, then, to raise it to the rank of a botanical district, under the name “Sounds-Nelson.” Even when this is done there are three distinct sections–(1) the former Marlborough Sounds Subdistrict; (2) the area lying westwards and southwards of Nelson City between the mountains (Ben Nevis, Gordon's Nob, &c.) and Tasman Bay, extending to the western boundary; and (3) the high mountains as a whole.

This separation of the South Island portion of the Ruahine-Cook Botanical District raises the North Island part to the rank of a district, to which we apply the name “Ruahine-Cook.” Thus the North and South Islands are kept distinct floristically and ecologically.

The flora of the Sounds-Nelson Botanical District is marked by a fairly high degree of local endemism, as shown by the following list:—

Gramineae.—Poa acicularifolia–a variety probably confined to the Mineral Belt; Festuca–a species possibly confined to the Mineral Belt.

Ranunculaceae.—Clematis parviflora var. depauperata—given on the authority of Man. N.Z. Flora, but we know. nothing about it.

Cruciferae.—Notothlaspi australe var. stellatum.

Leguminosae.—Probably an undescribed species of Carmichaelia—grows in the Rai and Pelorus Valleys, the stems extremely slender. the flowers small.

Euphorbiaceae.—Poranthera microphylla.

Thymelaeaceae.—Pimelea Suteri.

Boraginaceae.—Myosotis Monroi.

Labiatae.—Scutellaria novae-zelandiae.

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Scrophulariaceae.—× Hebe Simmonsii Ckn. et Allan comb. nov. = × Veronica Simmonsii Ckn. in Trans. N.Z. Inst., vol. 48, 1916, p. 202; H. rigidula Ckn. et Allan comb.' nov. = Veronica rigidula Cheesem. in Man. N.Z. Flora, 1906, p. 514; H. divaricata Ckn. et Allan sp. nov. = Veronica Menziesii Benth. var. divaricata Cheesem. in Man. N.Z. Flora, 1906, p. 512; H. Gibbsii Ckn. et Allan comb. nov. = Veronica Gibbsii T. Kirk in Trans. N.Z. Inst., vol. 28, 1896, p. 524.

Compositae.—Olearia serpentina Ckn. et Allan sp. nov. ined.—Common on the Mineral Belt at all altitudes; Celmisia Rutlandii C. cordatifolia, C. Macmahoni; Raoulia Gibbsii; Cassinia Vauvilliersii (Homb. et Jacq.) Hook. f. var. serpentina Ckn. et Allan.

With regard to the vegetation, that of the northern and eastern portions of the district in the coastal-lowland-montane belt has a plant-covering similar to that of the southern part of the Ruahine-Cook Botanical District. Thus the lowland forest is taxad or Beilschmiedia tawa rain-forest on the better ground, and Nothofagus truncata-N. Solandri forest in exposed positions or where the soil is poorest, while near the sea there is on the most fertile soil Dysoxylum spectabile coastal-forest.

On coastal rocks Arthropodium cirratum (absent in Ruahine-Cook), Phormium Colensoi, Astelia Solandri, and Griselinia lucida are characteristic. Digitalis purpurea, Rubus fruticosus (agg.), and the indigenous Cassinia leptophylla are common weeds which form pure associations. Dairy-farming is carried on in the fertile valleys on artificial meadows which replace rain-forest proper.

In the lowland-montane area southwards and westwards from Nelson City, which is the driest part of the district, there are wide stretches of Leptospermum scoparium, mostly induced. In places this has been replaced by orchards. The hills are occupied by Leptospermum scoparium, Pteridium esculentum, or Danthonia pilosa grassland. As the western boundary of the district is approached Nothofagus forest again puts in an appearance with the addition of N. fusca and N. Menziesii. Where forest extends, or did originally extend, continuously from the North-western District to the Sounds-Nelson, it is not easy to decide to which of the two it belongs, but certain forest species are peculiarly North-western—e.g., Podocarpus acutifolius, Dracophyllum latifolum, and Senecio Hectori.

The Mineral Belt is a striking feature of the district. It extends south-west from D'Urville Island for a distance of about sixty miles, and is clearly marked out from the forest by its stunted vegetation, due probably to excess of magnesia in the soil. It is usually quite narrow, being in one part barely 100 yards wide, but its widest part, on the flanks of the Dun Mountain, is more than three miles across. It occurs from sea-level to the subalpine belt, its plant-covering changing considerably with increase of altitude.

The subalpine flora of the district is rich in species, since near its western boundary it has, in addition to its flora proper, a good many species in common with the adjacent North-western District. Much of the high-mountain area is insufficiently explored botanically, so we can give no details of real value.

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Notes on New Zealand Floristic Botany, including Descriptions of New Species, &c. (No. 4).

[Read before the Wellington Philosophical Society, 27th August, 1924; received by Editor, 31st December, 1924; issued separately, 6th March, 1926.]

44. * Acaena microphylla × Sanguisorbae.

A few seeds were sown in 1923 by L. C. of a supposed hybrid as above collected by him in the Dart Valley (Fiord Botanical District). The seed germinated well, and the seedlings betray their hybrid origin in the colour of the leaves (brown, pale green, darker green, slate-colour, &c.) and the size and shape of the leaflets. So far none have flowered. It is possible that one of the parents may be Acaena inermis, while it is not possible to say what particular variety of A. Sanguisorbae was the other parent.

45. Acaena Sanguisorbae Vahl. var. sericei-nitens Bitter.

This is the common upland Acaena of the South Otago Botanical District, and it extends into the adjacent part of the Fiord Botanical District. Seed collected by L.C. in the neighbourhood of Lake Wakatipu in 1923 has produced a number of seedlings most closely resembling one another, showing that the variety is a true microspecies (Jordanon).

46. Astelia nervosa Banks et Sol. ex Hook. f.

This aggregate species includes several distinct varieties. We here segregate the broad-leaved lowland swamp-plant with stout male inflorescence from the common narrow-leaved forest-plant with slender male inflorescence. The former has been given specific rank as A. grandis by Kirk (Trans. N.Z. Inst., vol. 4, p. 245, 1872), but it seems better to retain it in the aggregate. We thus have—


Astelia nervosa Banks et Sol. ex Hook. f. var. grandis (Hook. f. ex T. Kirk) Ckn. et Allan comb. nov.


Astelia nervosa Banks et Sol. ex Hook. f. var. sylvestris Ckn. et Allan var. nov.

Varietas distincta foliis plerumque ± 1.6 m. longis, interdum valde longioribus, ± 5 cm. latis, anguste linearo-lanceolatis attenuatis; paniculis masculis gracilibus ± 40 cm. longis.

47. Carmichaelia australis R. Br. var. egmontiana Ckn. et Allan var. nov.

Ramulis angustis applanatis ± 3 mm. latis; racemis circiter 5-floris.

North Island: Egmont-Wanganui Botanical District—Common as a member of the subalpine scrub on Mount Egmont.

[Footnote] * The numbers follow on consecutively inl this series of papers.

[Footnote] † The abbreviation we are using for “L. Cockayne.”

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This variety is at once distinguished by its narrow branches and branchlets from the more widely-spread forms with comparatively broad branchlets. An examination of the type specimens of Kirk's var. strictissima showed that our plant is quite distinct in its much narrower branchlets, its spreading few-flowered racemes. The flowers of var. egmontiana are small, the standard being dark purple flanked by purple lines, and shading to a white margin. A seedling plant of var. egmontiana examined had the petioles and leaves on both surfaces sparsely clothed with short hairs. The leaves were pinnately 3–5-foliate, with leaflets ± 8 mm. long, broadly obovate, sessile.

48. Cassinia Vauvilliersii (Homb. et Jacq.) Hook. f. var. serpentina Ckn. et Allan var. nov.

Folia minora, laminis circa 4 mm. longis infra pallide brunneis haud. fulvis, costis minus carinatis; inflorescentia valde parviora ± 1.5 cm. diam., capitulis ± 20, circa 5 mm. longis.

South Island: Sounds-Nelson Botanical District—On Mineral Belt of the Dun Mountain: L. C.

Distinguished from C. albida (T. Kirk) Ckn. by the colour of the tomentum and the much smaller leaves. It might quite well be thought an epharmonic form of C. Vauvilliersii, due to the magnesian soil, but, if so, it is apparently “fixed,” as it has kept unchanged to any extent for five years in cultivation.

49. Coprosma Petriei Cheesem.

There are two varieties * of this species, distinguished by the colour of the drupes.


Coprosma Petriei Cheesem. var. atropurpurea Ckn. et Allan var. nov.

This is the type, described by Cheeseman (Trans. N.Z. Inst., vol. 18, p. 316, 1886). He states that the drupes are “bluish,” but perhaps it is more accurate to describe them as translucent, faintly stained very pale blue.


Coprosma Petriei Cheesem. var. astropurpurea Ckn. et Allan var. nov.

A typo drupis atropurpureis differt.

In this variety the drupes are much the colour of port-wine. The two varieties often grow side by side, but usually in pure patches of considerable size. They do not seem to hybridize. We have noted both varieties in various parts of the North-eastern and the Eastern Botanical Districts. Evidently each variety comes true from seed.

50. Epilobium melanocaulon Hook.

Two well-marked varieties constantly occur side by side on stony river-beds throughout the range of the species. Possibly the two varieties cross, but we have no notes on the matter.


Epilobium melanocaulon Hook. var. viride Ckn. et Allan var. nov. Caulibus foliisque griseo-viridibus.


Epilobium melanocaulon Hook. var. typica Ckn. et Allan var. nov. Caulibus intense atropurpureis paene nigris foliis atrorubris.

This variety is by far the more common.

[Footnote] * Since the descriptions were drawn up L. C. has seen a variety near Cass with white flowers.

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51. Epilobium pernitens Ckn. et Allan sp. nov.

Herba parva, prostrata, affinis E. peduncularis A. Cunn. sed foliis floribusque valde distincta. Caules procumbentes radicantes saepe conferti. Folia maxime nitentia, viridia, interdum purpurascentes, convexo-concava, satis crassa, rotunda ± 6 mm. diam. vel subrotunda, marginibus paulo recurvatis. Petioli brevissimi sed distincti. Flores non multi, ut apparet, maiusculi, usque ad 16 mm. diam., axillares. Pedunculi ± 5 cm. longi, graciles, arrecti, rose0-brunnei. Calycis-lobi oblongo-lanceolati, acuti, ± 4 mm. longi, pallide virides vel rubentes. Petala alba, late obovata, ± 8 mm. longa, apice valde emarginata, ± 10 atris lineis ornata. Capsuli ± 3.1 cm. longi, glabri, pedunculis in fructu ± 5.3 cm. longis.

North Island: Ruahine-Cook Botanical District—Ruahine Mountains, Tararua Mountains, in subalpine herb-field on margins of bare places: H. H. A. South Island: North-western Botanical District—Paparoa Mountains: L. C.

This beautiful little species is distinguished at once from its nearest relatives—E. nummularifolium, E. pedunculare, and E. nerterioides—by its extremely glossy, convex, very small, rotund or subrotund leaves and its comparatively large flowers. It is to be recommended for the alpine garden, as its small patches of glistening foliage and large white flowers remind one somewhat of Linnaea borealis. It sows itself abundantly, but is not likely to become a garden-weed.

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Fig. 1.–Leaves of Gaya : A, G. ribifolia Mount Fyffe; B, G. Lyallii; C, G.ribifolia, Mount Torlesse; D,G. Allanii, juvenile; E, F, G. Allanii, adult. × ½.

52. Gaya Allanii Ckn. sp. nov.

Arbuscula circa 4.5 m. alta; ramuli iuventute striati, plerumque stellatopubescentes, demum glabrati, cortice griseo-brunneo obtecti. Folia juvenilia ± 1.5 cm. longa, ± 8 mm. lata, petiolis ± 5 mm. longis, ovata, crenato-serrata, acuta, sparsis stellatis pilis induta. Folia

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matura ± 5 cm. longa, ± 2 cm. lata, ovato-lanceolata, basi cuneata, submembranacea, grosse et incise bi-serrata vel crenato-serrata, pallide viridia, subtus pallidiora, acuta, supra et infra sparsis pilis stellatis albis brevibus plus minusve deciduis induta, nervis subconspicuis. Petioli graciles ± 1.5 cm. longi, stellatis pilis vestiti, canaliculati. Flores 3–5 fasciculati, brevium ramulorum apicibus versus vel axillares. Pedunculi graciles, ± 1.5 cm. longi, stellato-pubescentes. Calyx late campanulatus, 5-lobatus, dense stellato-pubescens, lobis acute triangulariis, ± 4 mm. longis. Petala ± 1.5 cm. longa, ± 7 mm. lata, oblique obovata, obtusa, retusa, unguiculata, infra satis dense stellato-pubescentia supra glabrata. Columna staminea brevis, basi distensa; filamenta gracilia ± 6 mm. longa, glabra; antherae ± 30. Ovarium ovoideum, ± 4 mm. longum, ± 2 mm. diam., plerumque 7-, interdum 6 vel 8-loculatum, loculis compressis, stellatis pilis vestitis.

South Island: Eastern Botanical District—Peel Forest, on margin of taxad forest: H. H. A. Flowers in January.

This very distinct species differs markedly in its smaller, differently shaped leaves, and rather smaller flowers with fewer carpels, from either of the other two species of the genus. Superficially it bears a much closer resemblance to Hoheria sexstylosa. It will some day become a famous garden-plant and take its place alongside the justly famed mountain-ribbonwood, G. Lyallii. It will be easily raised from seed, and probably from cuttings. Fig. 1 shows outlines of juvenile (D) and adult (E, F) leaves of Gaya Allanii, and of leaves from flowering specimens of G. Lyallii (B, Kelly's Hill, Westland) and G. ribifolia (A, Mount Fyffe, Seaward Kaikouras; C, Mount Torlesse, Canterbury).

53. Hebe as a Genus to replace a certain Section of Veronica.

F. W. Pennell (Rhodora, vol. 23, pp. 1–22 and 29–41, 1921; reprinted as Contrib. N.Y. Bot. Garden, No. 230, 1921) has restored Commerson's genus Hebe (Juss., Gen. Pl., 105, 1789). Hebe, as defined by Pennell, includes all those shrubs and trees hitherto referred to Veronica which occur in New Zealand, subantarctic South America, the Falkland Islands, Tasmania, and south-east Australia. The distinctions between Hebe and Veronica are as follows: the capsule dehisces septicidally (not loculicidally). the thick (not thin) septum splitting and each carpel opening distally by a median slit through the septal wall; the leaves are always opposite and after falling leave more or less conspicuous scars; the flowers are usually in axillary racemes, but these may be greatly reduced, spicate or corymbose; and all are evergreen shrubs, or in a few cases trees. Pennell remarks, “The austral distribution, with its suggestion of genetic remoteness, emphasizes Hebe's claim to recognition as a genus.” It is, indeed, the restriction of the group to those portions of the temperate Southern Hemisphere already noted, together with the evergreen, shrubby habit of its members, which so markedly separates Hebe from Veronica proper, that has induced us to support Pennell's action. Hebe thus becomes of great phytogeographical significance. Pennell also points out that “Hebe has an exceedingly baffling tendency to form local races, a habit at contrast with that of the other ‘Veronicas.”’

Now, L. C., at any rate, has long been of opinion that the section Hebe of Veronica should be raised to generic rank. Therefore, as soon as Pennell's paper came into his hands, he sent it to the late Mr. T. F. Cheeseman, and wrote on the matter to Professor Dr. L. Diels and

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Dr. C. Skottsberg, whose opinion on any subject concerning subantarctic botany must receive the most careful consideration. Cheeseman replied that Pennell's action was probably justified, but that, as in the new edition of his Flora the genus Veronica was completed, he did not intend to make any alteration, and that in the absence of any other systematist of note having dealt with the question he had in the first edition followed Wettstein's treatment in Die Pflanzenfamilien. That a botanist so conservative as Cheeseman should consider the change to Hebe as probably correct speaks strongly in favour of the alteration.

Diels considered the change might quite well be made, and pointed out how certain genera had been removed from the wide conception of Veronica, which were closer to that genus than was Hebe. Skottsberg, though in general considering that there is in certain quarters a tendency to greatly overdo the splitting-up of genera, thought that the case of Hebe was in another category owing to its strictly limited distribution. In New Zealand, in addition to ourselves, Mr. W. R. B. Oliver has decided Hebe must stand, and has adopted the genus in a recently written paper.

The two objections to the change are that it will be very slowly adopted in horticulture and with more or less inconvenience, and that over one hundred species will have to be transferred. Fortunately, this will very rarely lead to changes in specific names. The following changes have already been published or are in course of publication: H. blanda (Cheesem.) Pennell, H. buxifolia (Benth.) Ckn., H. elliptica (Forst. f.) Pennell, H. salicifolia (Forst. f.) Pennell, H. subàlpina Ckn. H. vernicosa (Hook. f.) Ckn.

54. Hebe Allanii Ckn. sp. nov.

Frutex parvus, circa 30 cm. altus, ramulis patulis, profunde cicatricibus fohorum delapsorum notatis; internodia ± 8 mm. longa, dense albis pilis pubescentia. Folia sessilia, patentia vel retroflexa, paene subcordata, ± 1.8 cm. longa, ± 8 mm. lata, oblonga, subacuta, dense supra et infra pubescentes, marginibus ciliatis interdum rubro-tinctis, coriacea, glauca, paulo incurvata; costa inconspicua. Folia subfloralia ± 1.1-cm. lata, ovata vel obovato-oblonga. Spicae ± 25 florae, breves, paucae, axillares apicem ramulorum versus; pedunculi usque ad 2.5 cm. longi, dense mollibus patulis pilis albis induti. Flores sessiles, conferti. Bracteolae ± 5 mm. longae leviter carinatae, ovatae, subacutae vel obtusae, pubescentes, calyce paululum breviorae. Calyx pubescens ± 5 mm. longus; lobi profunde incisi, ovati, obtusi vel subacuti, pallide virides, marginibus scariosis. Corolla alba; tubus ± 6 mm. longus, calycis lobos excedens, extra sparsis pilis ornatus infra glaber; lobi patuli, ± 6 mm. longi, basim versus pilis sparsis infra ornati, ovati vel ovato-oblongi, obtusi. Stamina exserta, antheris atropurpureis. Ovarium pubescens. Capsula adhuc non visa.

South Island: Eastern Botanical District—On rocky outcrops in tall tussock-grassland at about 360 m., Mount Peel: H. H. A.

This well-marked species comes nearest to Hebe amplexicaulis, but is separated at once by its shorter branches, its leaves closely covered with hairs on both surfaces and with ciliate margins, its smaller, denser inflorescences, and smaller flowers, which are hairy in all parts except the upper portions of the corolla-lobes. It may easily be mistaken for H. Gibbsii on account of the presence of hairs on the leaves, calyx, &c., but in the latter species the leaves are ciliated only, and the calyx-lobes

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are much narrower and sharply acute. Although Hebe is so richly represented in gardens, a species so distinct and striking as H. Allanii is certainly a great acquisition. It is easily raised from cuttings and from seed, and will probably grow in poor soil in almost any situation.

55. Hebe amplexicaulis (J. B. Armstg.) Ckn. et Allan comb. nov.

This species was first described by Armstrong (N.Z. Country Jour., vol. 3, p. 56, 1879) from a plant collected in the upper Rangitata by J. F. Armstrong. This (given in the description as “decumbent”) is the well-known almost prostrate garden-plant, which is to be taken as the type of the species, and which is here given the varietal name vera. The species was again described by Armstrong in 1881 (Trans. N.Z. Inst., vol. 13, p. 352), a plant collected in the upper Waiau, Nelson, being included in the conception of the species. The words “or suberect” are added to the habit description. Cheeseman's description (Manual N.Z. Flora, p. 525, 1906) differs from the original in stating the corolla-tube to be “about the length of the calyx,” and he remarks, “Mr. Armstrong describes the corolla-tube as long, but it barely equals the calyx in all the specimens I have seen, including an authentic one from him.”

Our observations in the field and in the garden show that H. amplexicaulis is an aggregate allied to the group that includes H. albicans (Petrie), but differing in the more obtuse semi-amplexical leaves, and the spicate not racemose inflorescences, with larger obtuse bracteoles. We find the corolla-tube of H. amplexicaulis distinctly longer than the calyx, but this is not shown till the flower has fully developed, so that young inflorescences may have the flowers as described by Cheeseman. Although the floral characters are very similar in all, we consider the habit differences sufficient to mark off the following varieties:—


Hebe amplexicaulis (J. B. Armstg.) Ckn. et Allan var. vera Ckn. et Allan var. nov.

This is the type, and needs no further description.


Hebe amplexicaulis (J. B. Armstg.) Ckn. et Allan var. suberecta Ckn. et Allan var. nov.

Caules non multi, pauciramosi; rami primo decumbentes deinde ascendentes, patuli, nudi; ramuli ultimi suberecti deorsum nudi, sursum foliis confertis obtecti; spicae breves floribus confertis.

South Island: Eastern Botanical District—A rock-plant from about 340 m. to 1,200 m., Mount Peel, abundant: H. H. A.

Distinguished from var. vera by its open straggling habit, with long naked branches ending in suberect branchlets with crowded leaves. In cultivation it becomes of denser, less rambling habit.


Hebe amplexicaulis (J. B. Armstg.) Ckn. et Allan var. erecta Ckn. et Allan var. nov.

Caules numerosi, erecti, ramosi; ramuli ultimi stricti, deorsum nudi, sursum foliis confertis obtecti; spicae longiores floribus numerosis.

South Island: Eastern Botanical District—On rocky outcrops at about 350 m. altitude, upper Rangitata River: H. H. A.

Only a few plants of this very distinct erect bushy variety were seen. The spikes are longer with less crowded flowers than in var. suberecta, and are more abundantly produced, so that the shrub presents a handsome appearance when in flower.

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56. Hebe buxifolia (Benth.) Ckn. var. pauciramosa Ckn. et Allan var. nov.

Rami ultimi pauci, arrecti, foliis brevibus patentibus dense obtecti.

South Island: North-western, Western, Fiord, and South Otago Botanical Districts—In wet ground, mainly subalpine: L. C.

Hebe buxifolia is an aggregate species embracing many forms, but all distinguished by the smooth, glossy, green, keeled leaves, with short but distinct petioles (seedling and reversion leaves are deeply toothed or subpinnate), the short spikes of flowers more or less aggregated into heads, and the large bracteoles much resembling ordinary leaves. There are many intergrading forms extremely difficult to classify, but a knowledge of the species throughout its range shows that there are several well-marked varieties, which almost certainly hybridize, so producing the remarkable polymorphy. The varieties so far described are—(a) var. prostrata, which more or less closely hugs the ground; (b) var. odora, much-branched, eventually forming a globose bush; and (c) var. pauciramosa, here discussed. This last, possibly itself an aggregate, is distinguished by its erect final branches (it may be more or less prostrate at the base), straight, erect stems, leafy only in their upper portions, which branch but little and form a shrub of open habit. We have under observation other apparently distinct varieties, but they require further investigation.

57. Hebe Treadwellii Ckn. et Allan sp. nov.

Frutex humulis, ramulis elongatis, depressis, dispersis, cortice pallide brunneo obtectis, inferne cicatricibus foliorum delapsorum notatis. Folia arte imbricata, ± 2.5 cm. longa, ± 1.2 cm. lata, obovato-oblonga, subacuta, fere sessilia, supra valde concava, infra convexa, glabra, crassa, coriacea, satis nitentia, pallide viridia, marginibus pallidioribus; costa depressa. Petioli brevissimi applanati. Racemi duplices in foliorum superiorum axillis positi, aliquanto foliis obscurati, ± 1.2 cm. longi, pauci-flori; pedunculi, rhaches et bracteae (marginem versus) minute pubescentes. Bracteolae subulatae, calyce breviores. Flores albi, parvi, superiori sessiles, inferiori breviter pedicellati. Calyx 4-segmentatus; segmenta ovata, acuta, tubo corollae breviora, pallide viridia, marginibus ciliatis. Corollae tubus brevissimus, latus, calycem aequans: limbus 4-lobatus, haud patulus, lobis obtusis. Ovarium et stigma glabrum. Capsula non visa.

South Island: Western Botanical District—On stony ground at about 1,200 m. on Mount Ollivier (Sealy Range), and near the Mueller Glacier at a similar altitude: L. C.

This species, probably a common plant within its range, comes into the group which contains Hebe pinguifolia and its allies, but it is distinguished by its green leaves, its fewer few-flowered racemes almost hidden by the leaves, its subulate bracteoles, its smaller flowers with acute calyx-segments, and its glabrous ovaries and stigmas.

58. Metrosideros perforata (Forst.) A. Rich. = M. scandens Sol. ex Gaertn.

The specific name given by the Forsters (Characteres Generum, p. 72, 1776) was the first valid one, and Richard's adoption of it (Essai d'une Flore de la Nouvelle-Zélande, p. 334, 1832) must be followed.

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59. Myosotis Colensoi (T. Kirk) Ckn. et Allan comb. nov. = M. decora T. Kirk ex Cheesem.

Exarrhena saxosa Hook. f. was described by Hooker (Fl. Nov.-Zel., i, 202) from a plant collected by Colenso in the North Island at Titiokura. In the Handbook (p. 196) he united with it a plant from the Dun Mountain, Nelson, collected by Monro and Travers. Kirk (Trans. N.Z. Inst., vol. 27, p. 351, 1895) described a plant from limestone rocks at Broken River, Canterbury, collected by Enys and himself, as E. Colensoi, and united with it the Titiokura plant, saying, “A remarkable plant which has hitherto been confused with the Nelson E. saxosa Hook. f.” But, as E. saxosa is based upon the Titiokura plant and not the Nelson one, Cheeseman appears to have adopted a suggestion of Kirk to rename the Broken Hill plant as Myosotis decora (Manual, p. 462, 1906), the Nelson plant receiving the name M. Monroi (ibid., p. 469). But, since the Broken Hill plant provided the material for Kirk's original description of his E. Colensoi, that specific name should be adopted.

60. Nothopanax arboreum × Pseudopanax crassifolium var. unifoliolatum.

Mr. E. Phillips Turner recently discovered on Kapiti Island, growing amongst Leptospermum ericoides, what he thought to be the above hybrid. In this conclusion we are strongly inclined to concur. The plants, three in number, were growing side by side close to a young plant of P. crassifolium var. unifoliolatum. From the largest plant, some 40 cm. tall, Mr. Turner collected four leaves. Fig. 2 should, give a sufficient idea of the leaf-form and size. Except B, the leaves differ greatly from those of any stage of the polymorphic P. crassifolium. The excessive increase in breadth is what might be expected from a cross with N. arboreum. The texture is similar to that of N. arboreum, and the venation with the secondary veins diverging at angle of about 50° is that of N. arboreum, the angle in the case of P. crassifolium being about 20°. The midribs are prominent above and below, as in juvenile P. crassifolium, and the marginal teething is nearer that of P. crassifolium than of N. arboreum. The base of the lamina is intermediate in form.

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Fig. 2.—Leaves of Nothopanax arboreum × Pseudopanax crassifolium var. unifoliolatum. × ½.

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Fig. 3.—Leaves of Nothopanax simplex, N. anomalum, and N. parvum. × ½.

61. Nothopanax parvum (T. Kirk) Ckn.

L. Cockayne * recently expressed the opinion that Nothopanax parvum = N. anomalum × simplex. This view is supported by a number of specimens of what was apparently N. parvum studied by him in the Spey Valley, west of Lake Manapouri. Unfortunately, too few specimens were collected, but they were present in great numbers and made up a most polymorphic group. These juvenile plants differ greatly from those of the supposed parents, but exhibit features which certainly suggest a mixture of parental characters. N. simplex has juvenile leaves ovate-serrate or 3—5-foliolate with the leaflets deeply cut (fig. 3, A—a small

[Footnote] * See L. Cockayne, Hybridism in the New Zealand Flora, The New Phytol., vol. 22, No. 3, p. 126, 1923.

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example). These are succeeded by 3-foliolate leaves with the leaflets lanceolate-serrate (fig. 3, B, C). The adult has simple lanceolate-serrate leaves (fig. 3, D). It is almost certain that there are distinct races differing in their juvenile stages. N. anomalum has small 3-foliolate juvenile leaves with leaflets elliptic-ovate or rounded-ovate with crenate margins (fig. 3, E), succeeded by simple adult leaves oblong-orbicular, crenate-toothed (fig. 3, G, H). The leaves of the Spey Valley juveniles show intermediate characters, as can be seen from the drawings of three specimens (fig. 3, K1, K2, K3; L1, L2, L3; M1, M2, M3, M4). Two juveniles from Stewart Island are figured (fig. 3, N1, N2, N3; O1, O2, O3) which also show an intermingling of parental characters. Adult flowering specimens are figured from Inchbonnie, Westland (fig. 3, P1, P2, P3), and Stewart Island (fig. 3, R1, R2, R3; S1, S2, S3; T1, T2, T3). An examination of the drawings, traced from nature-prints, will be sufficient to show the intricate way in which the various parental characters are intermingled, without the necessity of drawing up an elaborate table. That flowering should occur on forms so different as P, R, S, T is important evidence as to their hybrid origin. This is borne out by the intermediate character of the inflorescences, bridging the gap between the few-flowered simple umbels of N. anomalum and the comparatively large compound umbels of N. simplex.

62. Ourisia Macphersonii Ckn. et Allan sp. nov.

Herba perennis, rhizomatis longis ± 8 mm. diam. Folia radicalia subcoriacea. Laminae ± 4.5 cm. longae, ± 3.5 cm. latae, supra pilis mollibus sparsis vestitae, infra glabratae, ovatae vel ovato-oblongae, apice obtusae, basi subcordata vel cuneatae, marginibus crenatis vel crenato-serratis, ciliatis; subtus purpuratae, venis viridibus distincte reticulatae. Petioli ± 3.5 cm. longi, striati, sparsis albis pilis induti, vaginantes, bases versus latescentes. Inflorescentia maiuscula; rhachis ± 30 cm. longus, basi ± 1 cm. diam., erectus, nervatus, pilis brevibus vestitus, interdum purpurascens, plerumque 5-verticillatus, verticillis sursum deminuentibus in omnibus partibus. Bracteae verticillatae, plerumque 5 per verticillum; inferiorae breviter petiolatae, late lanceolatae vel lanceolato-oblongae, obtusae, crenato-serratae vel crenatae; superiorae oblongae, obtusae, sessiles. Flores 7—8 per verticillum. Pedicelli ± 4 cm. longi, patuli, paulo crassi, albis pilis vestiti. Calycis-lobi usque ad basim 5-partiti, extra pilosi; lobis oblongis vel obovatis, apice purpuratis, obtusis vel subacutis, marginibus ciliatis, venis purpuratis distinctis. Corollae ± 2.5 cm. diam., albae, in fauces luteae glandulosis pilis ornatae; tubi ± 6 mm. longi, 2 inferiori lobi subrotundi vel late oblongi, patentes; 3 superiori lobi obovato-oblongi, suberecti, 3—4 venis distinctis, apice rotundati vel prope truncati. Stamina 4, supra apicem corallae tubi paulo exserta; antherae cremeae, reniformes. Styli longi, graciles, papillati, stigmatis 2-lobatis. Capsula paulo compressa, ± 1 cm. longa, ± 5 mm. lata, mucronata, calyce persistente induta.

South Island: Fiord Botanical District—Shady banks of River Spey at about 210 m., and in open boggy ground towards Wilmot Pass at 670 m. altitude: L. C.

This species has affinities with Ourisia macrophylla Hook. and O. Colensoi Hook. f., but differs from the former in the rather thicker, smaller leaves, the absence of cauline leaves, the lower bracts lanceolate petiolate (not linear-oblong), the yellow-throated flowers, the oblong or obovate (not

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lanceolate or linear) calyx-lobes, and the rather larger hardly turgid capsules. From O. Colensoi it is at once distinguished by the stouter habit, bracts in whorls of ± 5 (not paired), more numerous larger flowers on longer petioles, in whorls of 7–8, and the calyx-lobes not linear.

The plant blooms for about two months, commencing under cultivation near Wellington in the middle of October, and continuing until the second week of December. It appears to be of easy cultivation, and in the southern rainy parts of New Zealand should grow with the greatest vigour. The tiers of large white flowers with yellow throats, one above another after the manner of Primula japonica, make it a most effective plant for the alpine garden. It will not tolerate drought.

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[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]

Fig. 4.—Leaves of Plagianthus betulinus (× ½), and of P. divaricatus and P. cymosus (× 1/1).

63. Plagianthus cymosus T. Kirk.

This so-called species invariably grows along the banks of tidal rivers not far from the sea where P. betulinus and P. divaricatus grow near one another. Mr. W. Martin, B.Sc., has recently collected it in Chatham Island. Cheeseman. (Illust. N.Z. Flora, vol. 1, t. 21, 1914) says, “If its characters are carefully compared with those of P. betulinus and P. divaricatus it will be recognized that it stands nearly half-way between the two species. The suspicion of a hybrid origin at

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once arises, and it must be confessed that such an assumption would go far towards explaining the peculiar rarity of the plant in some of its localities. On the other hand, fruiting specimens sent to me by Mr. H. J. Matthews from the Pelorus Valley have well-developed and well-ripened seeds with a fully developed embryo, and show no signs of the impaired fertility frequently seen in hybrids.” L. Cockayne, in the New Phytol. paper already cited, listed it as a probable hybrid, the case of which required further investigation. In view of the number of hybrids now known to produce fertile seed, Cheeseman's objection does not seem to have any weight.

We have examined specimens from several localities—Pelorus Valley, Kaitaia, Chatham Island—and the leaf-characters strongly suggest a hybrid origin. P. divaricatus (fig. 4, B1, B2, B3, B4, B5) has small linear obtuse leaves, quite entire, while P. betulinus has large ovate-lanceolate crenate-serrate leaves, acute, with more or less cuneate bases (fig. 4, A). Representative leaves taken from single shoots from each locality well show the mingling of parental characters, in their intermediate size and inconstant shape and toothing—Kaitaia specimen (fig. 4, C1, C2, C3, C4, C5); Pelorus River specimens (fig. 4, D1, D2, D3, and D4, D5, D6); Chatham Island specimen (fig. 4, E1, E2, E3, E4, E5). These should be compared with the figures given in Illustrations of the N.Z. Flora from specimens collected in the Pelorus Valley, which show still another somewhat different form. Leaves taken from among the inflorescences of herbarium specimens of P. betulinus are also shown (fig. 4, E, G, H). These show a reduction in size but retain the characters of P. betulinus, and it is to be remembered that in this species the leaves are not fully developed at flowering-time. P. divaricatus has flowers solitary or in few-flowered fascicles, while P. betulinus has numerous flowers in large panicles. Our Kaitaia specimen has small male panicles just as might be anticipated on the hybrid theory, while, according to Mr. Martin, in the Chatham Island plant the fruit-clusters never exceed four. The number of anthers in our Kaitaia specimen is about 15–20 per flower. There can hardly be any doubt that P. cymosus is a hybrid with the parentage suggested. *

64. Pterostylis confertifolia Allan sp. nov.

Herba terrestris glabra, ± 7 cm. alta; erecto caule e tubere pisiforme ± 7 mm. diam. Folia conferta caulem amplectantia, inferiora scariosa squamiforma, obtusa, superiora plerumque 3, erecto-patentes, pallide viridia, paulo crassa, in siccitate tenuiora; laminae 3–5 cm. longae, 1.5–2 cm. latae, elliptico-ovatae, obtusae, venis per vitam obscuris, marginibus paulo incurvatis, basim versus in vaginas latas albas angustatae. Flores solitarii, virides, venis rubris, vix folia excedentes, aliquanto in fructu increscentes, ± 2 cm longi, ovario obovoideo ± 1.5 cm. longo excepto. Galeae erectae usque ad ± 12 mm., deinde arcuatae, apicibus acutis. Petala lateralia in inferioribus dimidiis linearo-oblonga, in superioribus falcata, acuta. Sepala lateralia linearia, usque ad ± 8 mm. connata, deinde in lobos 2, ± 1.2 cm. longos, erectos, acuminatos producta.

[Footnote] * The Waimakariri station mentioned by Kirk (Student's Flora, p. 71, 1899) appears to be based on a mistake. There are no specimens thence in his herbarium. The original specimen from Dunedin was (according to information supplied by the Hon. G. M. Thomson, taken from his original notes of many years ago) gathered from a plant culti vated in a garden.

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Labellum subcrassum, latericium, linearo-oblongum, marginibus minute serrulatis; in canaliculatum, minute retusum, paulo exsertum apicem angustatum. Unguis curvatus, apice 2 longis et aliquis brevibus satis latis filamentis ornatus. Columna gracilis, galeam partem erectam aequans; auriculae decurrentes, superioribus lobis late triangularis, apice breviter acuminatis; inferioribus lobis late oblongis, obtusis, apice mollibus pilis vestitis.

North Island: Wellington Botanical District—On margins of subalpine scrub, and in lower subalpine herb-field, about 1,200 m. altitude, Ruahine Mountains, near Apiti: H. H. A.

Though coming into the group containing P. Banksii, P. australis, P. graminea, that have cauline leaves only, this species is very distinct. So crowded are the broadly elliptic-ovate, obtuse leaves that superficially the species more closely resembles the P. foliata group. The plant grows in small colonies, and when growing through moss cushions the stems may be much elongated.

65. Ranunculus maculatus Ckn. et Allan sp. nov.

Herba parva, caespitulosa. Folia numerosa, radicalia, maiusculis brunneis maculis notata; laminae ± 9 mm. longae, ± 10 mm. latae rotundato-cuneatae, supra paulo pilosae, nervis tribus distinctis; 3-lobatae usque ad media, mediis lobis parvioribus; lobi lati, obtusi, inaequaliter 2–3 crenato-dentati; petioli ± 16 mm. longi sine vaginis, ± 1 mm. diam., erecto-patentes, pilis longis sparsis appressis induti; vaginae ± 14 mm. longae, ± 3 mm. latae, marginibus paene hyalinis. Scapi ± 5 cm. longi, satis crass, ± 2.5 mm. diam., longis pilis induti. Sepala 5, fugacea, 3 mm. longa, 2 mm. lata, late ovata, obtusa, infra sparsis pilis obtecta, marginibus scariosis. Petala plerumque 5, ovata vet obovato-spathulata, 5 mm. longa, 2 mm. lata, flava, supra nitentia, basim versus alba. Nectarium breve, subulatum. Stamina non multa, filamentis ± 1.5 mm. longis; antherae ellipticae, 0.5 mm. longae. Carpella ± 20, glabra, aliquanto turgida. Stylus brevis, stigmate paulum recurvato.

South Island: South Otago Botanical District—In wet ground at 1,200 mm. on Rock and Pillar Range: L. C.

This seems to be a distinct species with no near relative.

66. Ranunculus rivularis Banks et Sol. ex Forst. f. var. glareosus Ckn. et Allan var. nov.

Ab varietatibus aliis speciei habitu robustiore, foliis 3-lobatis, lobulis inaequaliter 2–3 lobulatis, petalis obovato-oblongis haud linearo-oblongis, distinguitur.

South Island: Eastern Botanical District—(1) Ninety-mile Beach, L. C.; (2) mouth of Ashburton River, H. H. A. On shingly or sandy foreshore.

This variety is very closely allied to the other varieties of the species, but is apparently confined to coastal gravel or sand, a station very different from those usual for R. rivularis. It is of robust habit, with far-spreading rhizomes emitting tufts of dark-green leaves on long petioles, the blades being 3-lobed almost to the base, with the lobes again irregularly lobulate. In cultivation the plant becomes rampant, and in one case grew over and suppressed a common damp-ground form of the species.

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Ferns and Flowering-plants of Mayor Island, N.Z.

[Read before the Philosophical Institute of Canterbury, 1st October, 1924; received by Editor, 24th October, 1924; issued separately, 6th March, 1926.]

Plate 5.

Mayor Island (Tahua of the Maori) lies in the Bay of Plenty, north-north-east of Tauranga, about 32 km. (20 miles) from the mainland and beyond the 180 m. (100fath.) depth line. The island, of volcanic origin, is somewhat square, of 1,277 ha. (3,154 ac.) extent, with a high ridge running east and west across the south end, and north and south on the western side, rising to about 396 m. (1,300 ft.) at the north end. The centre of the island is occupied by a great crater, some 8 km. (5 miles) in circumference, in which are two lakes—one of 2.5 ha. (6 ac.), dark blood-red from a distance; the other 8 ha. (20 ac.), of a deep blue-black. The colour is apparently due to the abundance of algal growth. There is a small opening, Opo Bay, on the south-east corner, the usual landing-place. Seaward there are rough water-worn cliffs streaked with veins of obsidian and surmounted by ash-beds sloping into the crater.

Dr. L. Cockayne, F.R.S., has kindly forwarded us the notes taken by him on a very brief visit to the island in February, 1905, and the following account of the vegetation is based on these and upon the notes and specimens taken by one of us (K. W. D.) during a fortnight's stay in March, 1924. We have to thank Dr. Cockayne for critically examining certain of the specimens collected.

The list of species shows that Mayor Island belongs to the Thames Subdistrict of the South Auckland Botanical District of Cockayne. *

The dominant tree is the pohutukawa (Metrosideros tomentosa), many specimens of which are of great size, the largest noted measuring 9 m. in circumference, with a trunk clear of branches for 3 m., and surmounted by a magnificent spreading head. The pohutukawa forms great groves in the lower flats all over the island.

The slopes of the main ridges are a continuous succession of small, deep, dry gullies, the spurs of which are clothed in thickets of manuka (both Leptospermum scoparium and L. ericoides), amongst which Knightia excelsa, about 3.5 m. tall, is dotted about. The gully-bottoms have a dense growth of whau (Entelea arborescens), and the side-slopes contain large makomako (Aristoteha serrata) up to c. 0.7 m. in diameter, Litsaea calicaris, Dysoxylum spectabile, Suttonia australis, Brachyglottis repanda, Melicytus ramiflorus, and odd tree-ferns. The liane Rhipogonum scandens is frequent, and the undergrowth is mainly Coriaria sarmentosa, Rhabdothamnus Solandri, and a few ferns. The undergrowth is, however, much altered and depleted owing to the presence of pigs, while the unstable surface militates against regrowth. The high ridges contain a dense scrub in which manuka, Cyathodes acerosa vars., Leucopogon fasciculatus, Coprosma spp., Gaultherio oppositifolia are prominent.

[Footnote] * See Cockayne, L., 1921. Die Vegetation der Erde, xiv: The Vegetation of New Zealand, p. 300 et seq. Leipzig: Wilhelm Engelmann.

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There are small areas of swamp by the lakes, but otherwise the crater-basin is covered with pohutukawa and manuka. Much of the manuka shrubland is an indigenous-induced association on ground formerly cultivated by the Maori. In one place exists a clump of Pinus Pinaster, and this aggressive species threatens to extend its area rapidly. One old cultivation-ground is covered in Hibiscus trionum.

A small grassy flat at Opo Bay, containing scattered pohutukawa, has as its main constituents Sporobolus indicus, Holcus lanatus, Dactylis glomerata, Poa pratensis, Lolium perenne, Trifolium repens, and is thus an exotic-induced association.

There are a number of other exotic species established, especially on the old Maori clearings. Some of them are Ricinus sp., Rosa eglanteria, Phytolacca octandra, Oenothera odorata, Rumex obtusifolius, Solanum nigrum, Anagallis arvensis, Erigeron canadense.

List of Species of Pteridophyta and Angiospermae.

The list includes 98 species or distinct varieties, distributed among 78 genera and 50 families. The abbreviations “fl.” and “ft.” indicate that the species were observed to be in flower or in fruit in March, 1924.

  • Lycopodiaceae: Lycopodium volubile Forst. f.

  • Psilotaceae: Psilotum triquetrum Swartz.

  • Marattiaceae: Marattia fraxinea Smith.

  • Cyatheaceae: Cyathea dealbata Swartz; Cyathea medullaris Swartz; Cyathea Cunninghamii Hook. f.

  • Polypodiaceae: Adiantum hispidulum Swartz; Cheilanthes Sieberi Kunze; Pteris comans Forst. f.; Pteridium esculentum (Forst. f.) Cockayne; Blechnum capense (L.) Schlecht.; Asplenium flaccidum Forst. f.

  • Typhaceae: Typha angustifolia L. var.

  • Gramineae: Paspalum scrobiculatum L. (ft.); Oplismenus undulatifolius Beauv. (ft.); Spinifex hirsutus Labill.; Dichelachne crinita (Forst. f.) Hook. f.; Dichelachne sciurea Hook. f.; Agrostis Billardieri R. Br.

  • Cyperaceae: Mariscus ustulatus (A. Rich.) C. B. Clarke (ft.); Eleocharis sphacelata R. Br. (ft.); Scirpus nodosus Rottb.; Gahnia gahniae-formis (Gaud.) Heller; Uncinia sp.

  • Liliaceae: Rhipogonum scandens Forst. (ft.); Cordyline australis Hook. f.; Astelia Banksii A. Cunn. (ft.); Astelia sp. (epiphytic); Dianella intermedia Endl.; Phormium tenax Forst. (ft.); Phormium Colensoi Hook. f.

  • Piperaceae: Macropiper excelsum (Forst. f.) Miq. var. major Cheesem.; Peperomia Endlicheri Miq. (fl.).

  • Proteaceae: Knightia excelsa R. Br.

  • Polygonaceae: Muehlenbeckia complexa (A. Cunn.) Meissn.

  • Chenopodiaceae: Salicornia australis Sol.; Salsola Kali L.

  • Aizoaceae: Mesembryanthemum australe Sol. (fl.).

  • Ranunculaceae: Ranunculus hirtus Banks et Sol. (ft.).

  • Monimaceae: Hedycarya arborea Forst. (ft.).

  • Lauraceae: Litsaea calicaris (Sol.) Benth. et Hook. f. (ft.).

  • Cruciferae: Lepidium oleraceum Forst. f. var.

  • Pittosporaceae: Pittosporum Colensoi Hook. f.; Pittosporum crassi-folium A. Cunn.; Pittosporum sp. (probably P. umbellatum).

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  • Rosaceae: Acaena Sanguisorbae Vahl. var. (ft.).

  • Oxalidaceae: Oxalis corniculata L. var. (fl.).

  • Meliaceae: Dysoxylum spectabile (Forst. f.) Hook. f. (ft.).

  • Euphorbiaceae: Euphorbia glauca Forst. f.

  • Coriariaceae: Coriaria sarmentosa Forst. f. (ft.).

  • Corynocarpaceae: Corynocarpus laevigata Forst.

  • Sapindaceae: Dodonaea viscosa Jacq.

  • Rhamnaceae: Pomaderris phylicaefolia Lodd.

  • Elaeocarpaceae: Aristotelia serrata (Forst.) W. R. B. Oliver.

  • Tiliaceae: Entelea arborescens R. Br. (ft.).

  • Malvaceae: Hibiscus trionum L. (fl. and ft.).

  • Violaceae: Melicytus ramiflorus Forst.; Hymenanthera novae-zelandiae (A. Cunn.) Hemsl. (ft.).

  • Thymelaeaceae: Pimelea virgata Vahl.; Pimelea prostrata (Forst. f.) Willd. var. (fl.); Pimelea sp. (= P. Urvilleana Cheesem. non A. Rich.).

  • Myrtaceae: Leptospermum scoparium Forst. (ft.); Leptospermum scoparium Forst. var. (ft.); Leptospermum ericoides A. Rich. (ft.) [there also occurs a form with narrow-linear leaves and small fruits that is possibly L. ericoides × lineatum]; Metrosideros tomentosa A. Rich. (ft.).

  • Onagraceae: Epilobium cinereum A. Rich.

  • Halorrhagidaceae: Halorrhagis erecta (Murr.) Schindl. (fl.); Halorrhagis procumbens (Sol.) Cheesem.

  • Umbelliferae: Apium prostratum (DC.) Labill. (fl.).

  • Araliaceae: Nothopanax Sinclairii (Hook. f.) Seem.; Nothopanax arboreum (Forst. f.) Seem. (ft.).; Nothopanax sp. ? [the specimens gathered include a single remarkable leaf of what may prove to be an undescribed species—there are seven sessile leaflets, the larger ones broadly obovate with tapering cuneate bases and rather distant blunt teeth: see Plate 5]; Pseudopanax Lessonii (A. Rich.) C. Koch.

  • Ericaceae: Gaultheria antipoda Forst. f. var.; Gaultheria oppositifolia Hook. f. (ft.).

  • Epacridaceae: Cyathodes acerosa R. Br. [there are distinct forms prescnt, one of which is probably var. oxycedrus (R. Br.)]; Leucopogon fasciculatus (Forst. f.) A. Rich. (ft.); Leucopogon Fraseri A. Cunn.; Dracophyllum Sinclairii Cheesem.

  • Myrsinaceae: Suttonia australis A. Rich. (ft.).

  • Loganiaceae: Geniostoma ligustrifolium A. Cunn. (ft.).

  • Convolvulaceae: Calystegia Soldanella (L.) R. Br. (fl. and ft.); Dichondra repens Forst.

  • Verbenaceae: Vitex lucens T. Kirk (ft.).

  • Solanaceae: Solanum aviculare Forst. f. (fl. and ft.).

  • Scrophulariaceae: Hebe salicifolia (Forst. f.) Pennell var. (fl.).

  • Gesneriaceae: Rhabdothamnus Solandri A. Cunn. (fl.).

  • Myoporaceae: Myoporum laetum Forst. f. (ft.).

  • Rubiaceae: Coprosma grandifolia Hook. f. (ft.); Coprosma lucida Forst. f. (ft.); Coprosma lucida Forst. f. var. (ft.); Coprosma robusta Raoul (ft.); Coprosma robusta Raoul var. angustata Kirk ? [this is a very narrow-leaved form, and may be a hybrid; it does not belong to the × C. Cunninghamii group.) (ft.)]; Coprosma retusa Hook. f.

  • Campanulaceae Lobelia anceps L. (ft.).

  • Compositae: Brachyglottis repanda Forst.; Senecio Banksii Hook. f.

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A leaf of Nothopanax sp. Mayor Island.

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Vegetation of Mount Peel, Canterbury, N.Z.
Part 1.—The Forests and Shrublands.

[Read before the Canterbury Philosophical Institute, 1st October, 1924; received by Editor, 20th October, 1924; issued separately, 6th March, 1926.]

A. Introduction.

(a.) General.

The present paper owes its inception to a statement by Cockayne (1917, p. 62), when discussing his proposed botanical districts: “The actual boundaries of many of the districts are extremely hard to fix, and in no few cases must always be artificial, though that detailed research which must take place in due course as phytogeographic workers increase in number will eventually find out the most natural limits.” Observations were made at all seasons during the period 1917–21, some thirty weeks being spent in the field, in the attempt to provide one such detailed study. I am deeply indebted to my friend and master, Dr. L. Cockayne, F.R.S., F.N.Z.Inst, &c., for his unfailing interest and encouragement in all my botanical work, and for his help and criticism during the investigation.

For some sixty-five years the grasslands of Mount Peel have been carrying sheep, and for a long period the river-flats have been grazed by cattle. The forests have been partially milled, and large portions have been felled and turned into arable or pasture land. Burning has been practised extensively on the grasslands, and fire has not altogether spared the forest. The destruction of the forest still continues, but fortunately a considerable area has been created a scenic reserve. Obviously the present facies of the vegetation must differ considerably from its primitive condition, and successions now in progress must differ greatly in most cases from those which produced the primitive vegetation. Admirably suited to the needs of students of such vegetation is the classification of associations given by Cockayne (1919, p. 147): (1) primitive, (2) modified, (3) indigenous-induced, (4) adventitious-induced, (5) artificial. I also follow Cockayne (e.g., 1921, passim) in his usage of the terms “formation,” “association,” “subassociation,” “colony.” The term “relic” I apply to fragments of communities that persist in the midst of associations that have reached a further stage in the succession—e.g., relic river-bed communities in tussock-grassland, relic beech forest in rain forest. Individual plants may also be relics of a community that has passed by—e.g., Polystichum vestitum, a relic of forest. In some cases it is difficult to decide whether a community is relic or indigenous-induced.

(b.) Physiography and Climate.

The area examined is some 11,000 ha. (27,000 ac.), having its centre in about 43° 52′ S. lat. and 171° 12' E. long., and is included in the Eastern Botanical District of the Southern Botanical Province, as delimited by Cockayne (l.c., 1917, p. 65). The Rangitata River from just below its gorge forms the eastern boundary for some 12 km. (7 ½ miles), and the

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long gorge of the Orari forms the western boundary. The southerly-facing slopes arising from the plain, here 270 m. (886 ft.) above sea-level, reach 1,308 m. (4,291 ft.) in the peak of Little Mount Peel. Thence the crest of the ridge runs north-west, descending to 1,125 m. (3,691 ft.), and rising in the rounded mass here called Middle Peel to 1,585 m. (5,200 ft.), and culminating in the double peak of Big Mount Peel at 1,740 m. (5,709 ft.). The northern boundary is formed by streams running to the Orari and Rangitata from the saddle, 1,220 m. (4,003 ft.), immediately to the north of the summit. The southern boundary is formed by the remnants of forest on the plain. Numerous streams descend to both rivers from the ridge, several of the eastern ones joining to form the Lynn. On all these streams a series of waterfalls and miniature gorges occur. Small shingle-fans end the eastern streams, while the streams of the western slopes form small hanging valleys ending in waterfalls into the Orari. Both Big and Little Mount Peel are sharp in outline, with much-shattered rocky summits and numerous rock-buttresses, whereas Middle Peel is covered with finer debris and gives rise to fairly extensive screes (shingle-slips). The spurs leading to the main divide lie athwart the prevailing north-west winds, and have a higher average elevation as one proceeds along the ridge towards Big Mount Peel. The topography is thus of a youthful character, and offers varied habitats for plant communities. The main mass of the mountain is formed of the comparatively easily-weathered greywacke rock.

The outstanding climatic features affecting the vegetation are the rainfall, snowfall, and wind. Mount Peel lies within the rain-band of 760–1,020 mm. annual fall, which stretches throughout the South Island, narrow in Canterbury and broadening out in Marlborough and Otago. Peel Forest itself, however, has a somewhat higher mean annual rainfall, as shown in the following table, recalculated in millimetres from the Journal of the Canterbury Agricultural and Pastoral Association for 1918, 1919, and 1920. The mean value is stated to be derived from records of “a considerable number of years.”

[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]

Year. Jan. Feb. Mar. April. May. June. July. Aug. Sept. Oct. Nov. Dec. Total.
1917 50 0 67 1 111.8 72.6 148.6 44.7 121 2 98 8 243 3 102.1 79 0 275 1 1,414 3
1918 132 9 128 5 101 2 32.3 16.8 66 6 26.4 87.6 80.5 122.4 141 0 122 4 1,066.3
1919 165.0 6 4 28 2 101.3 12 4 62 2 114.6 85.3 190.0 40.4 130 6 121 2 1,138 5
Mean 116 6 110 7 126.7 95.3 63.0 77 0 83.3 57.4 94.5 97 1 106 7 118 9 1,147.2

The difference in rainfall over a period of nine years between Mount Peel Station, exposed to the dry north-west winds sweeping down from the Rangitata Gorge, and Peel Forest, sheltered in a basin of hills, is as follows: Mount Peel (mean), 960.15 mm.; Peel Forest (mean), 1,164.81 mm. Rain is fairly generally distributed over the year; any month may have a fall of over 150 mm., the average monthly rainfall being over 95 mm. In addition the number of cloudy days on the forested area is considerably in excess of that elsewhere. Speight, Cockayne, and Laing (1911) have sufficiently dealt with the influence of the Southern Alps on the rainfall and climate of the eastern ranges and plains. The influence of wind is dealt with later on.

From some 1,200 m. upwards on the Rangitata face of the mountain snow lies practically throughout the winter, though a strong north-west wind, even in midwinter, may temporarily clear the slopes in a remarkable manner. On the southern slopes the line of winter snow falls c. 200 m.,

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and the snow remains much later in the season. From 1,200 m. down to 650 m. snow lies for weeks at a time on the southern face, and for lesser periods on the Rangitata face. Below this elevation snow may fall at any season, but usually lasts only a few hours. The great influence of snow upon the vegetation at Mount Peel will be clear from the detailed descriptions which follow, especially in regard to the formation of the belts.

Of the past climatic history of the area, most importance attaches to the post-glacial period. This has been dealt with by Speight (1911), and as to its botanical significance by Speight, Cockayne, and Laing (l.c., 1911, p. 343). During the period of glaciation the higher peaks of Mount Peel probably rose above the glacier-field as nunataks, and the special features of the summit vegetation, as well as more general questions raised, are dealt with later on.

(c.) The Formations and their Physiognomy.

The broad features of the plant-covering are due to the interaction of the major factors briefly discussed above: (1) Topography, (2) climate, (3) the glacial period, (4) the greywacke substratum. Great modifications have been caused by the advent of man, but the belts of vegetation still persist with a semi-primitive appearance and character. The lower slopes of Little Mount Peel from c. 600 m. downwards to the south-west, south, and south-east are clad in rain forest, which extends on to the upper plains, where it is now broken into fragments. Above the forest is an extensive belt of tall-tussock grassland, merging, on the Orari and Rangitata faces, below into low-tussock grassland, above into fell-field or herb-field, and finally into the open formation of the summit peaks. On the southern face the tall-tussock grassland merges below into a belt of Phormium Colensoi with Blechnum capense, in which Aciphylla Colensoi is prominent, thence by way of an interrupted belt of shrubland into the forest. Scattered through the tall-tussock grassland on the southern and western aspects occur more or less extensive patches of scrub, which, with the brown of Dracophyllum or the green of Hebe, diversify the tawny grassland. Lining the narrow stream-valleys are fringing forests, dark green below, and sage-green above owing to the groves of Gaya ribifolia, To the north especially towards the Orari, the slopes contain occasional dark masses of subalpine beech forest, merging above into scrub and then the shingle-slips. Below the terrace of the Rangitata lies its broad river-bed clothed in great stretches of gorse.

B. The Forests.

(a.) General.

Both the main types of New Zealand forest occur at Mount Peel—the rain forest and southern-beech forest. The southern-beech forest belongs to the subalpine beech-forest association, with Nothofagus cliffortioides as its sole dominant and only tall tree. It is the climax forest of the eastern mountain-ranges of Canterbury. The rain forest groups naturally into two associations—that of the terrace lands, dominated by Podocarpus dacrydioides, P. spicatus, and P. totara in varying proportions according to the edaphic conditions, and that of the hill-slopes, characterized by the infrequence of the podocarps, by the lack of any dominance in general, and by the development of several distinct subassociations. This rain forest is in marked contrast with the southern-beech forest at similar

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altitudes on Mount Somers (mean rainfall 857 mm.) and Mount Hutt (mean rainfall 1,048 mm.), and may be correlated with the more favourable conditions of sheltered situation and rather greater rainfall.

The composition of the rain forest as a whole is as follows: Pterid phyta—8 families, 19 genera, 43 species; Gymnospermae—2 families, 2 genera, 4 species; monocotyledons—6 families, 12 genera, 19 species; dicotyledons—31 families, 53 genera, 89 species. The Polypodiaceae have 24 species, Hymenophyllaceae 8, Rubiaceae 9, Compositae 10. Represented by 1 species each are 17 families. Of the 155 species, 23 may be classed as abundant, 71 as more or less common, 45 as occasional to infrequent, 16 as rather rare.

The growth-forms include 6 small tuberous-rooted herbs (5 summer-green, 1 saprophytic); 3 simple rosette herbs; 41 tuft plants (including 18 tuft-ferns, 6 tree-ferns, 9 grass-like herbs); 2 trailing herbs; 16 lianes; 26 creeping plants (8 patch-herbs, 3 mat-plants, 15 sheet-plants); 61 bushy plants (including 3 herbs; 1 semi-woody plant; 30 trees—3 tall, 8 medium, 19 small; 27 shrubs—14 open-branching, 4 fastigiate, 9 divaricating). There are thus 80 herbaceous species, 3 semi-woody, 72 woody. There are 15 ferns that occur as epiphytes, and 3 hemi-parasitic shrubs.

(b.) The Podocarp Forest.

1. Composition.

The podocarp forest may best be described in two sections, though these merge into one another. On the flood-plain of the Rangitata at about 240 m. Podocarpus dacrydioides is dominant, with P. spicatus subdominant. Seen from above, this section presents a billowy appearance, P. dacrydioides reaching 27 m. and overtopping P. spicatus by some 4–5 m. P. totara is infrequent. Near the base of the terrace-slopes, where streams from the mountain and ooze from the terrace cause a development of swampy ground, we get true but limited P. dacrydioides swamp forest. Wintera colorata is abundant, often attaining 9 m. Below there is a dense under-growth, which may be mainly juvenile P. dacrydioides, or contain varying amounts of Wintera colorata, Coprosma rotundifolia, Melicytus micranthus, Myrtus pedunculata, Griselinia litloralis. These are largely covered with lichens, liverworts, and mosses. On the floor and scrambling over the shrubs are occasional Rubus schmidlioides. The floor-plants are Uncinia uncinata, Nertera dichondraefolia, Blechnum discolor, Microlaena avenacea, Hymenophyllum demissum, but only where there is not an excess of water. Climbing up the trees is much Cyclophorus serpens. Near streams other shrubs occur, notably Carpodetus serratus, Schefflera digitata, Fuchsia excorticata, Nothopanax arboreum, Hebe salicifolia var. communis. Ferns are more numerous, and include Blechnum fluviatile, B. lanceolatum, Asplenium bulbiferum, Polypodium diversifolium. Here, too, occur Ptero-stylis Banksii, Stellaria parviflora, Pratia angulata, and other herbs. Where the floor becomes drier the ground-plants are mainly Astelia nervosa, Poly-stichum vestitum, Asplenium bulbiferum. Suttonia australis becomes a common undergrowth shrub, and here P. spicatus is subdominant, while Elaeocarpus Hookerianus, Nothopanax arboreum, and Pennantia corymbosa become common; Asplenium flaccidum, Polypodium grammitidis, Cyclophorus serpens occurring on them. There will be odd Dicksonia fibrosa On the steep terrace-slopes P. totara enters in, and the association merges into the next section characteristic of the upper terrace-flats. In general P. spicatus is dominant, with P. totara, and P. dacrydioides in lesser

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amounts. Here and there, but little inferior in height to the podocarps, is Elaeocarpus Hookerianus. The second layer is composed of Grisedinia littoralis, Nothopanax arboreum, Schefflera digitata, Carpodetus serratus, Pittosporum tenuifolium, with the usual fern epiphytes. The shrub layer is mainly made up of Wintera colorata, Coprosma rotundifolia, C. rhamnoides, Fuchsia excorticata, Aristotelia serrata. Metrosideros hypericifolia creeps over the ground and climbs high among the trees. The chief floor-plants are Polystichum vestitum, Uncinia uncinata, U. riparia, Blechnum discolor, Polypodium diversifolium. There are numerous seedlings, especially par-sonsias and podocarps. On the outskirts is a tangle of various Rubi, Calystegia tuguriorum, Parsonsia heterophylla, Clematis indivisa, C. foetida, and Metrosideros hypericifolia, the last especially abundant.

Where the floor is low-lying and damp the association closely resembles the flood-plain section, and in gullies merges into the hillside forest. On the drier ground, even when the ground is only slightly higher, P. dacrydioides practically disappears, being replaced by P. totara. This becomes dominant on the driest areas. Here Suttonia australis, Pennantia corymbosa, Hoheria angustifolia, Plagianthus betulinus, Edwardsia microphylla become common in the second layer, with occasional Melicytus lanceolatus, and more or less Pseudopanax crassifolium var. unifoliolatum, Coprosma linariifolia, Melicytus ramiflorus. Melicope simplex, Nothopanax anomalum, and to a less extent Fuchsia Colensoi and Coprosma areolata enter the shrub layer. On banks and stony places occur Asplenium flabellifolium, A. Hookerianum, A. flaccidum, and occasionally Lycopodium rolubile, and sheets of Hydro-cotyle americana, H. moschata, Schizeilema Hookeri. On the forest-margin Senecio sciadophilus is added to the lianes.

2. Plagianthus betulinus Subassociation.

Plagianthus betulinus is known to form indigenous-induced associations, but the community at Mount Peel appears to be a remnant of a primitive subassociation, as the trees are adult, up to 15 m. tall, and reach 0.6 m. diameter at 1 m. from the base. The community now occurs only in certain fragments, and is much modified owing to the isolation of the fragments and the intrusion of stock. Besides the dominant tree, there are less amounts of large Griselinia littoralis and Hoheria angustifolia, with occasional adult Pseudopanax crassifolium var. unifoliolatum and Fuchsia excorticata. The undergrowth is made up of Coprosma linariifolia, Melicope simplex, Pseudopanax crassifolium (juvenile), with the usual lianes. The shrubs are sparse, and the floor is clothed mainly with exotic species, of which Poa pratensis, Holcus lanatus, and Dactylis glomerata are the chief. Where the undergrowth is dense the grass covering is replaced by Poly-stichum vestitum and odd seedlings, mainly parsonsias, Suttonia australis, Griselinia littoralis, Pittosporum tenuifolium.

3. Successions.

(1.) Milled Forest.—Forest that has been milled has two characteristic features : (i) The entry of various aliens, especially those of the “berried” type—e.g., Hypericum Androsaemum and Rubus fruticosus; (ii) the rapid growth of certain of the smaller trees and shrubs. Aristotelia serrata often forms thickets of slender stems 5–6 m. tall, Fuchsia excorticata also increases greatly, while Wintera colorata and Carpodetus serratus grow into small trees. Lianes often increase greatly and convert the forest into an almost impenetrable mass. Hoheria angustifolia does not increase markedly in such forest. Where cattle enter milled forest

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open spaces increase and the forest becomes separated into clumps. Of especial attraction to cattle are Uncinia spp., Carex spp., and to a less extent Microlaena avenacea. Certain shrubs—e.g., Schefflera digitata, Nothopanax arboreum—are also fed on. Where cattle do not penetrate, seedling podocarps and other species of the original forest become established, and the forest progresses towards its previous condition.

(2.) Succession to Podocarpus dacrydioides.—The succession from swamp to Podocarpus dacrydioides may be traced in places adjacent to the adult podocarp association. P. dacrydioides and Griselinia littoralis establish themselves in the swamp, which has Olearia avicenniaefolia, Coprosma spp. on its margin, and in some cases Olearia virgata var. and O. lineata are members of the swamp community. P. dacrydioides becomes dominant and forms thickets. The following description of the progress of such thickets is taken direct from my notes: The P. dacrydioides are here some 30–50 cm. apart, have a basal diameter of ± 12 cm., and a height of 9–14 m. The tall slender trunks are sparsely clad with short branches, and end in a small canopied head of foliage. The shrubs are few and poorly developed, and include Suttonia divaricata, Coprosma rotundifolia, Wintera colorata, C. parviflora, Pseudopanax crassifolium var unifoliolatum (juvenile), with rather larger Griselinia littoralis. These occur only on the patches of rather drier ground. Beneath is much Carex secta (Blechnum capense on it), Astelia nervosa, and, where the shrubs occur, some Polystichum vestitum. On the outskirts Rubus australis climbs to the top of the trees. Between the herbs of the floor are pools of water, slow-flowing streams, or boggy ground.

(3.) Succession to Hoheria angustifolia.—There are occurring on the felled areas two distinct successions—to Poa caespitosa grassland, and to IIoheria angustifolia thickets. The former is in a very early stage, the tussock grassland forming at present only clumps of varying size scattered here and there on the artificial grassland. The second is a characteristic feature of the grassland. The scattered clumps of Hoheria angustifolia are found in all stages from an open community of juvenile plants of the usual divaricating form to groves of adult trees in closed association. By the time the hoherias are assuming adult form above the community becomes so dense as to form thickets, in which seedlings of the following species become more or less common: podocarps, Edwardsia microphylla, Pennantia corymbosa, Elaeocarpus Hookerianus, Fuchsia excorticata, with various coprosmas. As these develop, the groves assume the form of young mixed forest. This stage is especially well developed as a defined margin to the existing forests in certain places. Polystichum vestitum, a relic from the original forest, increases greatly in amount, and other ferns and the lianes become plentiful. On the low-lying damper areas Podocarpus dacrydioides enters with the Hoheria, and early on becomes physiognomically important. On the drier stony ground sheep penetrate the clumps as the divaricating stage passes away, use them as camping-places, destroy the undergrowth, and induce a grassy ground-vegetation.

(c.) The Forest of the Mountain-Slopes.

1. Composition.

The mixed podocarp forest gradually merges into the forest that clothes the slopes and fills the gullies of Little Mount Peel up to about 600 m. As one ascends, Podocarpus dacrydioides disappears almost at once, soon followed by P. spicatus; but occasional examples of P. totara occur through-out—large by stream-sides, dwarfed on slopes and ridges. The general

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groundwork is made up of the following species in greater or less abundance: Griselinia littoralis, Nothopanax arboreum, Pittosporum tenuifolium, P. eugenioides, Edwardsia microphylla, Aristotelia serrata, Hoheria angusti-folia, Melicytus ramiflorus, Leptospermum ericoides, Metrosideros lucida, Fuchsia excorlicata, Pseudopanax crassifolium var. unifoliolatum, Suttonia australis, Olearia avicenniaefolia. The most common shrubs of the undergrowth are Wintera colorata, Nothopanax simplex, Suttonia australis, Suttonia divaricata, Coprosma rotundifolia, C. rhamnoides, C. linariifolia. All the lianes occur except Senecio sciadophilus, Rubus australis being usually in great abundance. The chief floor-plants are Polystichum vestitum, Blech-num discolor, B. capense, Asplenium bulbiferum, Polypodium diversifolium, Uncinia uncinata, U. riparia, Astelia nervosa. Seedlings of the common trees are usually numerous.

By stream-sides there is a greater variety and abundance of ferns, and Schefflera digitata, Aristotelia serrata, Fuchsia excorticata become prominent, the latter often being so plentiful as to be of distinct physiognomic importance, especially in its leafless winter condition. Hebe salicifolia var. communis and Gaya ribifolia are almost confined to the stream-side, and, as one ascends, Olearia arborescens, Nothopanax Colensoi, Senecio elaeagni-folius (as an open spreading small tree) become noteworthy. Pittosporum eugenioides is rare in or absent from the stream-side vegetation. Along the actual stream Hymenophyllum spp. are specially abundant, and here occur Epilobium rotundifolium, E. linnaeoides, Erechtites glabrescens, Myosotis Forsteri, Australina pusilla, Pratia angulata, Corysanthes macrantha. Steep banks are clad in sheets of Blechnum Patersoni, B. capense, Hymenophyllum demissum, H. multifidum, and H. peltatum. In places occur numerous Hemitelia Smithii, and logs, trunks of trees, and shrubs are clothed in epiphytes, mainly Hymenophylla and various ferns, liverworts, and mosses. Locally Alsophila Colensoi is prominent among great sheets of Blechnum capense.

The streams of the Lynn Valley are characterized by abundance of Helichrysum glomeratum. Hebe leiophylla, Veronica Lyallii, and the presence of Notospartium torulosum, Veronica linifolia, Olearia fragrantissima and in places much Senecio bellidioides. Stream-side rocks and rocky walls have Hymenophyllum pulcherrimum and Lycopodium Billardieri in addition to the ferns already mentioned. Where small rocky outcrops occur on the forest-floor Polypodium diversifolium, Cyclophorus serpens, and Rubus australis are common, and, if the crevices contain much soil, Asplenium Hookerianum, A. flabellifolium, A. Richardi.

Vertical rock-surfaces exposed to sun and wind have scattered Olearia avicenniaefolia, Helichrysum Selago, Hebe amplexicaulis, Danthonia setifolia, Coprosma brunnea, Notospartium torulosum, Dichelachne crinita, Agropyron scabrum var., not always all together. Some shaded rock-surfaces are clad in Angelica montana, Lycopodium Billardieri, Veronica Lyallii, and green tussocks of Schoenus pauciflorus, or breadths of Corysanthes macrantha among moss, where water drips.

Spurs and ridges are either occupied by the subassociations later described, or have as the most numerous plants Leptospermum ericoides, Suttonia australis, Metrosideros lucida, Nothopanax arboreum, Coprosma linariifolia, a large-leaved form of Myrtus pedunculata and Edwardsia microphylla. The undergrowth is mainly Suttonia australis, Coprosma rhamnoides, and in the upper portions Nothopanax simplex. The floor is rather sparsely clothed, mainly with Blechnum capense (stunted), Astelia nervosa, Uncinia uncinata, U. riparia, U. caespitosa, U.filiformis, Lycopodium volubile At the upper limits the association is invaded by members of the

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fringing scrub—e.g., Dracophyllum longifolium (reaching a stature of 1.5 m., with spreading branches clothed .at the tips with rosettes of leaves of juvenile form, in which condition it flowers freely), Astelia Cockaynei, Phormium Colensoi, Leptospermum scoparium, Cassinia Vauvilliersii, C. fulvida var. montana, Gaultheria rupestris, and various small herbs, notably Celmisia spectabilis, Senecio bellidioides.

Where the ridges broaden out as they descend on to the terrace lands, plants of the stream-side creep upwards, and the undergrowth is mainly Coprosma rotundifolia, Wintera colorata, and Fuchsia excorticata. Here occurs the chief development of the tree-ferns Cyathea dealbata, C. Cunninghamii, and Dicksonia squarrosa, with a wealth of Asplenium bulbiferum, Leptopteris hymenophylloides, and Blechnum discolor.

2. Subassociations.

(1.) Leptospermum ericoides Subassociation.—On many of the spurs on the southern slopes is developed a distinct subassociation with Leptospermum ericoides dominant and often the sole tree. Seen in the flowering season, these spurs stand out against the dark background of the general mass like the veining of a vast leaf. The Leptospermum averages 15 m. in height, and has a long trunk crowned by a small semifastigiate head, forming with its neighbours a dense canopy. While most of the shrubs may be present, Suttonia australis is usually very abundant, along with Coprosma rhamnoides and C. parviflora. In places Cyathodes acerosa, elsewhere rather rare at Mount Peel, predominates, with it being usually Gaultheriq, antipoda vars. At its upper limits the association has an undergrowth mainly of Nothopanax simplex, with stunted Griselinia littoralis and Olearia arborescens prominent. The floor-plants are the same as those for the ridges in general, but also include Blechnum penna marina, Cotula squalida, and Lagenophora petiolata.

In some places the association appears to be decadent, with many fallen trunks and dead or dying trees. In such places Blechnum discolor is plentiful, and juvenile Metrosideros lucida occurs in varying numbers. The association descends towards the streams of the western faces of the ridges, but on the eastern faces is rapidly succeeded by the general mixed forest.

(2.) Metrosideros lucida Subassociation.—This occupies considerable areas on rocky knolls and on slopes with a western aspect, and may be recognized from afar by its uniform sombre colouring, or in a good flowering season by its masses of red blossom. On the knolls the Metrosideros is youthful, and there is evidence that it is successional to the Leptospermum subassociation. The Metrosideros is the sole tree, c. 7 m. high, of a semifastigiate habit, much branched from the base, with slender naked branches each carrying a small canopied head of foliage, the whole forming a dense cover. The undergrowth is similar to that of the Leptospermum association, and is usually sparse. At the lower elevations Coprosma linariifolia is occasional, as tall as the Metrosideros, with trunks c. 13 cm. diameter. At the higher elevations occurs rarely small Libocedrus Bidwillii. On the slopes the Metrosideros is often adult, and the floor has a close cover of well-grown Blechnum capense with frequent Alsophila Colensoi, or uear streams the Blechnum may be replaced by sheets of luxuriant Gleichenia Cunninghamii, forming almost pure colonies.

(3.) Nothofagus cliffortioides Association Relic.—At an elevation of some 510 m.—i.e., near the upper forest-margin—and at stream-junctions at a lower elevation, in a few places, occur small patches of Nothofagus cliffortioides forest quite distinctly marked off from the surrounding forest-mass. The

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communities appear to be relics from a more widespread association that has been replaced by the rain forest. The association as developed on the knolls contains Nothofagus cliffortioides as the sole tree. The trees, averaging 12–15 m. high, with trunks up to 50 cm. diameter, form a dense canopy, under which the undergrowth is scanty, consisting of odd stunted Suttonia australis, Metrosideros lucida, Nothopanax simplex, Coprosma microcarpa, Melicope simplex, Gaultheria antipoda, Leptospermum ericoides, Cyathodes acerosa, scattered widely. At the junction with the rain forest many of the rain-forest shrubs occur. The floor-plants are also sparse and stunted, and include Astelia nervosa, Pteridium esculentum, Blechnum capense, Polypodium diversifolium, Celmisia spectabilis, Elytranthe flavida is occasional. Seedling beeches are absent. Other examples of the community are reduced to a few old decadent trees amidst the incoming rain-forest species.

In the examples occurring at stream-junctions the floor is often covered in Dicranoloma moss-cushions, the trees are more youthful, with odd old decadent trees, and a litter of old trunks and broken limbs. Seedling beeches are very infrequent. Coprosma rhamnoides, C. parvifiora, C. microcarpa, Aristotelia fruticosa, Suttonia australis, Cyathodes acerosa, Helichrysum glomeratum are the commonest shrubs. Elytranthe flavida is fairly common, and Botrychium australe is common in several forms. Other occasional floor-plants are Polystichum vestitum, Uncinia uncinata, U. riparia, Blechnum capense, B. penna marina, Polypodium diversifolium.

(4.) Melicytus ramiflorus Subassociation.—At the Lynn Valley Melicytus ramiflorus forms subassociations, especially on westerly-facing slopes. The Melicytus becomes a good-sized tree (some specimens reaching a height of 11 m. with a diameter near the base of c. 50 cm.), usually much branched from near the base. Scattered among the Melicytus are occasional Edwardsia microphylla, Plagianthus betulinus, Hoheria angustifolia, Griselinia littoralis, Nothopanax arboreum. Undergrowth is very scanty indeed, there being little else but Suttonia australis and Griselinia littoralis. As one ascends the gullies the association merges into the mixed forest. The floor is stony and bare. The herbs are distant and small, mainly Pellaea rotundifolia, Asplenium flabellifolium, A. bulbiferum, Polypodium diversifolium, Astelia nervosa. Seedling Melicytus and Edwardsia are frequent.

(5.) Gaya ribifolia Association.—At the upper margins of certain gullyforests, and as isolated patches at higher elevations c. 750–900 m., occur groves of Gaya ribifolia, which elsewhere is an occasional member of stream-side forest. The association occurs on stony ground by stream-sides where there is a certain degree of shelter from strong winds. The Gaya is the sole tree (averaging 5 m. in height), and beneath is usually little but scattered Hypolepis Millefolium, Polystichum vestitum, Blechnum penna marina, and sometimes Coprosma propinqua, C. parviflora, C. ramulosa, Hebe buxifolia var. odora. With a less dense cover the shrubs become more numerous. The groves form a marked physiognomic feature, bare of foliage in winter, white with blossom in summer, sage-green at other periods. The association is somewhat xerophytic in character, and is not a part of the rain forest, being treated here merely for convenience.

3. Successions.

(1.) Aristotelia serrata Succession.—This indigenous-induced association is well known as a successional stage towards regeneration of forest that has been destroyed by fire, and presents no special features at Mount Peel. It occurs in several places on the lower slopes where fire has run up from

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the felled bush below. The Aristotelia forms dense thickets of tall slender stems (5–8 m. high), and contains a good deal of Fuchsia excorticata of small size. As shrubs, occur Wintera colorata, Melicytus ramiflorus, Suttonia australis, Coprosma rotundifolia. Blechnum discolor and Polystichum vestitum are more or less abundant, sometimes replaced by Blechnum capense. Rubus australis and Muehlenbeckia australis are frequent. Old Griselinia littoralis is almost the sole survivor from the original vegetation. Seedlings of many of the mixed-forest plants are frequent. On the damper slopes Fuchsia excorticata is subdominant.

(3.) Leptospermum ericoides Succession.—Thickets of Leptospermum ericoides occur on the more sunny slopes, both after burnt forest, and after sown grassland on felled areas. In the first case the ground is rapidly clothed in dense young Leptospermum, in which various seedlings get a root-hold, notably Pittosporum tenuifolium, Griselinia littoralis, Nothopanax arboreum, Suttonia australis, Carpodetus serratus, Pseudopanax crassifolium var. unifoliolatum, Hebe salicifolia var. communis, with Rubus australis, Coprosma rhamnoides, C. parviflora, and various ferns. Examples are met with where these forest species are beginning to dominate the Leptospermum.

On the artificial grassland the succession takes a different course where Polystichum vestitum survives the burning, its tussocks gradually overshading the grasses and other exotic and indigenous herbs. Fuchsia excorticata, F. Colensoi, Hoheria angustifolia are early comers along with the Leptospermum (both L. scoparium and L. ericoides), and Rubus australis forms mounded heaps. With the dominance of Leptospermum, thickets resembling the first described are reached.

(4.) Minor Successions.—In the Melicytus ramiflorus subassociation not infrequently patches are broken down by winter snows. In such places Rubus australis, Muehlenbeckia australis, M. complexa, and Calystegia tuguriorum rapidly form dense tangles. Growing up through such tangles are to be noted Hebe salicifolia var. communis, Plagianthus betulinus, Edwardsia microphylla, and Griselinia littoralis, which gradually assume dominance.

In several of the gully-forests more or less extensive slips have started successional movements. The actual course of events varies somewhat according to the nature of the surface exposed, the aspect of the slope, and the surrounding vegetation.

On the drier sunnier slopes early comers are Epilobium pedunculare, E. microphyllum, Gnaphalium collinum, Erechtites quadridentata, and Pteridium esculentum. Poa caespitosa, Danthonia semiannularis, D. pilosa, Dichelachne crinita, and various exotics, especially Verbascum Thapsus, Hypochaeris radicata, Rumex acetosella, come in. Early shrubs to establish are Olearia avicenniaefolia, Fuchsia Colensoi, Hebe salicifolia var. communis, Pittosporum tenuifolium, and the exotic Leycesteria formosa. Lianes capable of forming mounded heaps—Muehlenbeckia australis, M. complexa, Rubus australis, R. cissoides, R. subpauperatus—-are more or less common.

On damper slopes the sequence is more rapid, and Coriaria sarmentosa and Griselinia littoralis are prominent, as are Acaena Sanguisorbae var. pusilla and Calystegia tuguriorum. Trees of the neighbouring forest establish more rapidly.

(d.) Subalpine Southern-Beech Forest.

This forest is quite isolated from the rain forest, and occurs at higher elevations on the slopes of ridges leading down to the Orari. The sole tree is Nothofagus cliffortioides, many of large size, with trunks up to 1 m. or more in diameter. The largest trees are decadent, with projecting dead branches and excessive amounts of Elytranthe flavida. Quite striking is the absence of seedling and juvenile beeches. As one ascends, the trees

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become smaller, and near the crests of the spurs are quite dwarfed owing to exposure to wind. The forests are accessible to sheep, which use them as shelter, and have probably caused the undergrowth to be sparser than in the primitive condition. There are occasional Coprosma linariifolia, C. parviflora, Aristotelia fruticosa, and more rarely C. microcarpa. On the floor are odd small patches of stunted Blechnum capense, Polystichum vestitum, Blechnum penna marina, and in rocky places Asplenium flabellifolium, Acaena Sanguisorbae vars., Celmisia spectabilis. At higher elevations occur Lycopodium fastigiatum and Hymenanthera dentata var. alpina. Along streams there is a more varied vegetation, including Hebe salicifolia var. communis, Aristotelia serrata, Gaya ribifolia, Griselinia littoralis, Olearia avicenniaefolia, and in damper spots Danthonia Cunninghamvi, Oxalis lactea, Anisotome aromatica var. incisa, Angelica montana, and other species, along with various ferns, among which Asplenium Trichomanes and A. Richardi are noteworthy.

(e.) Considerations Derived From The Study of The Forests.

From a study of all the examples of Nothofagus cliffortioides forest now existing it would appear probable that Nothofagus cliffortioides once clothed the sheltered sides of all the gullies. The greater rainfall in the area to the south of Little Mount Peel and in the deeper gullies has allowed a more mesophytic type of mixed forest to gain the ascendancy, restricting the beech to the more exposed knolls, and even threatening them with extinction there.

Supporting this view is the fact that various stages in the decadence of the beech forest on the knolls can be observed, one knoll at an elevation of c. 330 m. being now reduced to merely four or five beeches, among which is the ordinary mixed forest.

Some support is also given by the fact that the forest in almost equally sheltered places at Mount Somers, with a rainfall of only 857 mm., is almost entirely Nothofagus cliffortioides forest, while at Mount Hutt, with 1,048 mm., the forest is intermediate in character.

Such a view as to the character of the Nothofagus cliffortioides forest seems strongly supported by the arguments of Speight (l.c., 1911, p. 408). In this paper, Speight, in attempting to trace the climate of Canterbury since the glacial period, attaches great importance to the existence of former extensive forests in Canterbury. He says: “Apart from this coastal forest there were at the beginning of settlement considerable areas of standing bush, containing totara, black-pine, and white-pine, at Mount Peel, Geraldine, Waimate, and especially on Banks Peninsula, as well as in a few other localities in hilly places favoured by a good rainfall and a rich soil. At Mount Peel a considerable area still remains. These were in all probability remnants of a regional forest containing totara which covered extensive areas on the eastern slopes of the main range of the South Island.” He adds: “It is a remarkable fact, however, that the existing patches of bush occur in just those situations in which they might be expected to occur from ecological considerations had a slight desiccation of the climate come about.”

Using Speight's summary, we may make the following surmises as to the history of the forest at Mount Peel:—

The general sequence of events since the glaciation of the South Island in Pleistocene and post-Pleistocene times appears to have been the following:—

(1.) Glacial conditions, with probable steppe climate existing con temporaneously on the land to the east of the terminations of the

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glaciers, a condition which probably continued for some time, as the glaciers were retreating. (The Rangitata glacier overrides the lower slopes of Mount Peel, which projects as a nunatak. With the retreat of the ice, fell-field is established, developing gradually into tussock-grassland on exposed slopes and Nothofagus forest on sheltered slopes.)

(2.) Moist climate over the tract to the east of the main range, during which the forests were established or were widely spread and the rivers built up their fans. (Mixed podocarp forest develops at Mount Peel as part of an extensive regional forest. Nothofagus driven from all but the exposed knolls and less sheltered slopes.)

(3.) Modified steppe conditions over the belt to the east of the main range. (Podocarp forest persists at Mount Peel, owing to especially favourable rainfall, while succumbing elsewhere. Relicts of Nothofagus forest enabled to retain their position, but with difficulty, and largely succumbing in the Peel Forest area.)

C. The Shrublands.

(a.) Composition.

Shrublands of all degrees from very open communities to dense thickets and scrub occur in many parts of the area. Some bear a semi-primitive stamp, but the majority are more or less modified by fire or grazing, and many are reduced to fragments. Others, again, are indigenous-induced, and certain adventitious-induced associations are of importance. Although typical members of the shrub-communities described are frequent, there is sometimes a most bewildering admixture of various types. My treatment in this section is much condensed, as a full treatment would run to inordinate lengths. The following figures include all the members of indigenous communities, but omit many species that occur only as invaders or relics. Of the species that can be considered true shrubland members there are—Pteridophyta, 3 families, 10 genera, 14 species; monocotyledons, 5 families, 12 genera, 14 species; dicotyledons, 28 families, 49 genera 105 species. There are 18 species of Rubiaceae, 18 Compositae, 7 Scrophu-lariaceae. There are 13 families represented by one species each. Of the 133 species, 22 are abundant, 51 more or less common, 43 occasional or infrequent, 17 rather rare. The growth-forms include 4 small tuberousrooted herbs; 2 simple rosette plants; 20 tuft plants (4 ferns, 10 grass-like plants, 2 herbaceous, 3 semi-woody, 1 tuft tree); 5 trailing plants; 13 lianes (2 ferns, 3 scramblers, 5 twiners, 3 tendril-climbers); 19 creeping plants, including 2 turf-forming, 6 mat-forming, 10 sheet-forming; 70 bushy plants, including 6 leafless or scale-leaved shrubs, 20 spreading, 8 ball-like, 8 fastigiate, 22 divaricating, 4 depressed shrubs. There are 39 herbs, 8 semi-woody plants, 86 .woody plants. Hemi-parasites number 4.

(b.) The Associations.

1. River-terrace and Debris Shrubland.

(1.) Lowland.—The association, taken broadly, is dominated by Coprosma parviflora and Discaria toumatou, and may be modified, a relic of river-bed shrubland, or indigenous-induced. On concave slopes in the low-tussock grassland, exposed to much wind and insolation, and subject to burning, dry debris slopes develop colonies of Pteridium esculentum with various Rubi, and often a good deal of Paesia scaberula. Scattered among these are often Discaria toumaton, Coprosma parviflora, C. propinqua, Corokia Cotoneaster, Aristotelia fruticosa. Or a dense scrub of these divaricating

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shrubs may be formed, with Rubus australis, R. cissoides, R. subpauperatus, Muehlenbeckia complexa, Parsonsia capsularis, Clematis marata, some or all scrambling over the mass. Sometimes Hymenanthera dentata var. alpina, as an upright divaricating shrub, is included. Below the densest scrub there is little else but Asplenium flabellifolium. The shrubland of riverterraces and stream-fans may be almost identical with the above, or contain other species—e.g., various divaricating coprosmas, Carmichaelia subulata, Notospartium torulosum, Melicope simplex, Coriaria sarmentosa, Leptospermum scoparium, L. ericoides, Fuchsia excorticata, Myrtus obcordata, Hebe salicifolia var. communis, Hebe leiophylla, × Hebe Kirkii, Olearia avicenniaefolia, Cassinia Vauvilliersii. Floor-plants are more numerous, and include Hypolepis Millefolium, Pellaea rotundifolia, Helichrysum filicaule.

(2.) Montane.—At an elevation of c. 600 m. occurs on coarse debris near streams an association with Olearia nummularifolia and Coprosma propinqua as dominants. There is also much Coprosma rugosa, Cassinia fulvida var. montana, stunted Dracophyllum longifolium, Coprosma parviflora, and sometimes Aciphylla Colensoi, Phormium Colensoi. The floor-plants are chiefly Hypolepis Millefolium, Polystichum vestitum.

(3.) Subalpine.—On debris slopes at c. 1,000 m. on small stabilized shingle-slips occurs a scrub dominated by C. parviflora and C. propinqua, with large open mats of C. ramulosa between. Hymenanthera dentata var. alpina, as a dense semi-cushion plant, and decumbent Coprosma serrulata, Dracophyllum uniflorum, Aristotelia fruticosa, Hebe buxifolia var. odora, Aciphylla Colensoi, Discaria toumatou are more or less common. The chief floor-plants are Hypolepis Millefolium, stunted Polystichum vestitum, Blechnum penna marina, Lycopodium fastigiatum, Acaena Sanguisorbae var. pusilla, Trisetum antarcticum. Clematis australis is occasional as a liane. Gaya ribifolia occurs infrequently.

2. Hebe buxifolia-Coprosma parviflora Scrub.

This occupies large areas on shaded slopes at c. 700–1,000 m., and is sometimes apparently a development from the subalpine scrub already described. It was formerly still more extensive, but has been broken up by fires. Besides the dominants, Coprosma propinqua, C. cuneata, Dracophyllum longifolium, D. uniflorum, Hebe Traversii?, Cassinia fulvida var. montana are more or less frequent. Hebe buxifolia increases greatly near streams, and here Aciphylla Colensoi and Coprosma serrulata are common, the latter not developing the creeping form with underground stems, but merely becoming decumbent. Notospartium torulosum occurs in some examples of the scrub, up to 950 m. altitude. At the higher elevations Hebe lycopodioides is often very common. Of the smaller plants the chief are Gaultheria depressa, Anisotome aromatica, Senecio bellidioides, Helichrysum bellidioides.

3. Leptospermum Shrubland.

(1.) Leptospermum Thicket after Grassland.—At Mount Peel the Leptospermum shrublands are mainly indigenous-induced after burning of forest or grassland, but there are not wanting examples that appear to be of primitive stamp. They occur in all stages from open shrubland to dense thicket. Here the association developing on burnt grassland is described. Both L. scoparium and L. ericoides invade burnt areas, usually the former dominating. The raoulias, Leucopogon Fraseri, Lycopodium fastigiatum, L. scariosum, that also invade burnt areas, are more or less suppressed by the rapid growth of the Leptospermum, as are Dracophyllum spp. and Gaultheria antipoda. The sunnier slopes favour Leptospermum, the shadier Dracophyllum. The chief other plants occurring in the later stages are

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Botrychium australe, small Pteridium esculentum and Blechnum capense, Uncinia caespitosa, U. riparia, Microtis unifolia, Prasophyllum Colensoi, Lagenophora petiolata.

(2.) Leptospermum Swamp.—There are one or two examples of small swampy areas on grassland after felled forest with Leptospermum ericoides and L. scoparium dominant in varying proportions. Large mats of Nertera depressa surround the trunks, among which the water finds its way slowly downwards. On the mats are any of Blechnum penna marina, Epilobium nummularifolium, Forstera Bidwillii (slender and “drawn-up”), Luzula campestus vars., Thelymitra longifolia, Hydrocotyle asiatica, Uncinia riparia, Pratia angulata, Ranunculus hirtus, Anisotome aromatica var. incisa. Some of the following shrubs are usually present: Suttonia divaricata, Coprosma propinqua, Griselinia littoralis, Wintera colorata, Podocarpus dacrydioides (this often abundant but small). In shallow depressions Carex secta may be the chief plant below, and decadent Phormium tenax is common, the decadence partly due to the entry of cattle, which browse on the Phormium and Carex. Where the cover is not dense occur Scirpus inundatus, Ranunculus rivularis, Epilobium pallidiflorum; and with the conditions approximating to bog there is little below but sphagnum, with Blechnum penna marina and B. capense prominent on it.

(3.) Leptospermum scoparium-Exocarpus Bidwillii Association.—Broken rocky outcrops in low-tussock grassland at its higher limits often have an association dominated by a variety of L. scoparium and Exocarpus Bidwillii. The association may be open or closed. The Leptospermum is here a depressed shrub with dense, stout, spreading branches clothed towards the tips with thick broad leaves. Rather less abundant are the low, dense semi-cushions of Exocarpus Bidwillii. Other occasional shrubs are dwarfed Gaultheria antipoda, Dracophyllum longifolium, Hymenanthera dentata var. alpina, Hebe Allanii, trailing Suttonia nummularia. Where the association is not specially dense there are, as floor-plants, Lycopodium scariosum. L. fastigiatum, stunted Blechnum capense, Leucopo [ unclear: ] on Fraseri. Standing above the mass may be Metrosideros lucida, as a ball-like shrub, and stunted Podocarpus totara.

4. Dracophyllum Shrublands.

(1.) Dracophyllum longifolium Thicket.—Dracophyllum longifolium is an abundant plant in various associations up to c. 900 m., where it is replaced by D. uniflorum. It also frequently forms closed associations on rather steep, somewhat shaded slopes, and tends to reproduce itself after destruction by fire. Floor-plants are in greater number than in the scrub associations, more light penetrating to the ground, the chief species being Gaultheria depressa, Lycopodium fastigiatum, Anisotome aromatica, Senecio bellidioides, Leucopogon Fraseri, Blechnum capense, Poa Kirkii, Trisetum antarcticum. Some of the following shrubs are usually present : Hebe buxifolia, Coprosma parviflora, Cassinia fulvida var. montana, Gaultheria antipoda, G. rupestris, Pimelea sp., with Phormium Colensoi, Astelia Cockaynei, Aciphylla Colensoi.

(2.) Dracophyllum uniflorum Shrubland.—On rocky buttresses and running up steep spurs, thickets or more or less open shrublands dominated by Dracophyllum uniflorum are a well-marked feature high up among the tall-tussock grassland. D. Urvilleanum also occurs, along with Gaultheria rupestris, G. depressa, Coprosma ramulosa, C. parviflora, C. serrulata, all in small amounts. The commonest herbs are Celmisia spectabilis, Anisotome aromatica, Seneco bellidioides, Ranunculus Monroi var. dentatus, Forstera Bidwillii, and Microlaena Colensoi.

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(3.) Dracophyllum rosmarinifolium Dwarf Shrubland.—On the exposed slopes of Middle Peel an interesting dwarf shrubland occurs in more or less extensive patches amongst the fell-field and intergrading with it. The Dracophyllum is dominant, and associated with it are Pentachondra pumila, Phyllachne Colensoi, Drapetes Dieffenbachii, Gaultheria depressa, Raoulia grandiflora, Celmisia laricifolia, Lycopodium fastigiatum.

5. Cassinia Open Shrubland.

As noted above, Cassinia fulvida var. montana is a member, sometimes an abundant one, of various shrubland associations. In the Blechnum capense-Phormium Colensoi belt this Cassinia and C. Vauvilliersii frequently occur in marked amounts, forming an open shrubland. The association is indigenous-induced after burning. Burning, though only slightly affecting the Blechnum capense, sufficiently opens the association to allow of the free-seeding Cassinia to increase greatly, the seedling being tolerant of considerable shade. The remaining plants are those of the original association described later.

6. Scrub of the Upper Margins of the Rain Forest.

(1.) General.—The forest at its upper margin merges either into tall-tussock grassland by way of a narrow scrubby belt, or into one of the shrub associations above described, but there is no broad marginal belt. In the gullies the scrub consists mainly of dwarfed Griselinia littoralis, Nothopanax Colensoi, N. arboreum, Olearia arborescens, O. avicenniaefolia, Senecio elaeagnifolius, Nothopanax simplex, Fuchsia ercorticata, Hebe salicifolia var. communis. Rubus australis scrambling over these completes the scrub. On the ridges the following are prominent: Leptospermum ericoides, Suttonia australis, Senecio elaeagnifolius, Nothopanax simplex, Dracophyllum longifolium, Gaultheria antipoda.

(2.) Senecio elaeagnifolius Dwarf Forest.—On certain broad ridges the mixed forest is succeeded by a rather extensive association in which Senecio elaeagnifolius is dominant, the remaining shrubs being quite few in numbers. The Senecio is here a large shrub, up to 4 m. high, much branched from the base, the branches stout, wide-spreading, naked, with loose papery bark. Above, the branches divide several times at wide angles, the branchlets ending in rosettes of closely-placed large coriaceous leaves of oval form, dark-green above, with pale-buff appressed tomentum beneath. The shrubs are close together and form a continuous canopy, beneath which the vegetation is sparse. Here and there are slender sparingly-branched Leptospermum ericoides, Nothopanax simplex, N. Colensoi, Coprosma linarii-folia, Griselinia littoralis. Smaller still are odd bushes of Coprosma rham-noides and Gaultheria antipoda, open in habit. The floor-plants are small in size and scattered—Blechnum capense, Lycopodium volubile, L. Billardieri, Polypodium diversifolium. Seen from a vantage-point above during the flowering season this association is beautiful with its masses of golden flower-heads completely hiding the foliage.

Literature cited.

Cockayne, L, 1917. Notes on New Zealand Floristic Botany, including Descriptions of New Species. &c. : No. 2. Trans. N.Z. Inst., vol. 49, p. 56.

—, L, 1919. New Zealand Plants and their Story. 2nd ed. Wellington.

—, L, 1921. Die Vegetation der Erde vol. 14, Vegetation of New Zealand. Leipzig.

Speight, R., 1911. The Post-glacial Climate of Canterbury. Trans. N.Z. Inst., vol. 43, p. 408.

Speight, R., L. Cockayne, and R. M. Laing, 1911. The Mount Arrowsmith District: a Study in Physiography and Plant Ecology. Trans. N.Z. Inst., vol. 43, p. 315.

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A Die-back of Pinus radiata and P. muricata caused by the Fungus Botryodiplodia pinea (Desm.) Petr.

[Read before the Nelson Philosophical Society, 14th November, 1924; received by Editor, 26th November, 1924: issued separately, 6th March, 1926.]

Plates 6,7.

In August of this year the attention of the writer was drawn to the presence of what seemed to be an unrecorded disease in two species of Pinus in the Marlborough and Nelson Districts. In Nelson the disease occurred in trees in the town itself, while in Marlborough it was reported in the country, where valuable shelter-belts of P. radiata were said to be seriously threatened. On a visit being paid to Marlborough to investigate the matter it was found that, while in the case of certain shelter-belts it was necessary to take prompt steps to prevent extension of the disease, this disease was not responsible for the general dying-out of pines throughout the drier parts of Marlborough; this is rather due to the combined effect of drought, a deep shingly subsoil, and the inroads of various insect pests, notably the aphis Chermes pini Koch. The fungus disease, however, is sufficiently disastrous in the belts in which it is established to require attention, and a description of it is therefore now given.

General Appearance of Infected Trees.

The disease can readily be detected from a distance, its chief characteristic being the sharp contrast presented by the diseased and the healthy parts of the tree. The tip of the stem or of one or two of the higher branches dies, the disease then extending downwards for a length of as much as 20 ft. The stem is more often affected than the upper branches, and both are more often affected than branches low on the tree, although in one exceptional instance the lowest limb of a large tree was found to be wholly invaded by the fungus.

The surface of an infected area is not sunken or distorted in any way, but if sufficient time has elapsed since infection took place for reproduction of the fungus to have occurred the pycnidia formed in this process will be found studding the host-surface (Plate 6, figs. 1 and 2). The pycnidia may occur singly, but usually they are arranged in ranks or groups, the ranks being either directed straight up and down the stem or, more frequently, arranged in the shape of a narrow horseshoe, directed downwards and enclosing in the upper part of the loop the scar of a needle-bearing shoot. As a rule the ranked or curved arrangement is still indicated, even when the whole surface is densely covered with pycnidia. The continued growth of the stem eventually separates the shoot-scars and tends to straighten the sides of the horseshoe.

The pycnidia are minute and black, and as they push up the epidermis they cause it first to appear white, and then rupture it into more or less elongated slits. On the splitting of the papery-white cuticle the upper

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Fig. 1.—Tip of main stem of Pinus radicta, showing pycnidia of Botryodiplodia pinea (Desm.) Petr. on the surface. × 1.
Fig. 2.—As fig. 1, but an older specimen. Pyendia in rows and in groups. × 1.5.

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Cone of Pinus radiata bearing scattered pycnidia on the exposed surface of the scales. × 2.5.

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surface of the pycnidia lies exposed. The length of the ranks of pycnidia varies from that of the diameter of a single pycnidium to 2 cm., while the width is frequently only that of one pycnidium, although it may be condiderably greater.

Internal Appearance of Infected Tissue.

The simplest way to see the effect of the disease is to split the stem down the centre for a distance that includes several nodes. If the fungus is well established the bark and cortex are dark brown or practically black, the xylem has a cinereous tint, while the pith varies in shade from its normal rusty brown when only slightly invaded to pure black when the fungus is present in quantity. In nearly every case the tissue at the nodes where the cones are attached is more heavily infected than are the internodal regions, and it can often be seen that in a single stem infection has taken place at several successive nodes, a diseased stem-or branch-tip therefore frequently being the result of multiple infection.

The cones also become infected, but as a rule the fructifications of the fungus, instead of occurring in groups or rows, are scattered and appear singly as minute black dots piercing the epidermis (Plate 7).

The Fungus.

Vegetative Growth in the Host.—Whether a wound in the host-surface is necessary to enable the fungus to gain entry in the first instance is not known; but, once established, it is an active parasite. Larvae of Sirex juvencus L. have been found in wood infected by Botryodiplodia pinea, and the death of the wood has been attributed to these larvae, the fungus being regarded as merely saprophytic; but, as it is quite common to find branches suffering from fungus attack without there being any trace of the grub, this suggestion is untenable.

As already mentioned, the path of infection can usually be traced back to a node, and then out into the short stems of the cones. It would seem that in some instances cones or cone-stems when young have been the original parts infected, and that the fungus has then worked down into the stem or branch. In other cases, however, infection seems to have taken place directly in the stem or branch involved.

The hyphae, at first hyaline, soon change to a fuliginous shade, and finally become quite black, especially when near the surface of the stem. The hyphae vary in width, but the older, coloured hyphae are usually the largest, attaining an extreme width of more than 10 micromillimetres. Here and there may be seen hyphae bearing on their surface a close arrangement of small papillate outgrowths, suggestive of rudimentary haustoria. These hyphae are intercellular in position, and are usually hyaline. Growth in the cortex is vigorous, and it soon becomes a semi-stromatic mass of crushed dead cells bound together by numerous much-branched hyphae. Later, as the pycnidia are being formed, a true stroma is gradually developed beneath the epidermis, and eventually practically all the host-tissues outside the xylem become a black stromatic mass, which may extend for a considerable distance, parallel to the surface, beneath the seemingly still-intact bark.

The entry of the hyphae into the xylem takes place with few exceptions by way of the rays (text-figs. 1, 2). In radial sections of an infected area almost every cell of the rays, and particularly the median albuminous ones,

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will be found to be permeated with the stout, dark strands of the fungus, and throughout the length of the ray numbers of slender branch-hyphae are given off into the tracheids, down whose length they can be traced for long distances. The hypha in a tracheid usually runs straight, but it gives off, at irregular intervals, further branch-hyphae, which as a rule pass directly through the pits in the lateral walls into neighbouring tracheids. Here they in turn commence a straight and usually downward course. Although hyphae in the tracheids are not as a rule as wide as those in the cortex, here and there one of greater width may be seen, almost filling the tracheid containing it. The colour of these larger hyphae, like those in the cortex, deepens with age to a fuliginous shade or to black, eventually causing the wood to appear cinereous to the naked eye.

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Fig. 1.—Semi-diagrammatic representation of invasion of the wood by hyphae travelling by way of the rays and thence into the tracheids.
Fig. 2.—Portion of fig. 1 enlarged. The hyphae travel chiefly through the albuminous cells of the rays. × 352.

Reproduction.—When the disease was first noticed, in August of 1924, pycnidia were already present on the surface. The spores in some of them were light brown in colour, but in most they were still hyaline. Similar stages may still be found in the original material as well as in that gathered in mid-November. The production of pycnidia thus continues for some time. The spores are seldom mature when the pycnidia first appear through the ruptured epidermis. Even in pycnidia that are known to be several months old most of the spores are still pale if not actually hyaline. All stages in complexity of pycnidial structure are to be found (text-figs. 3, 4, 5). They may be simple, papillate, and more or less spherical, with the free surface perhaps a little flattened. Two or more may occur in a row or group and have their walls in contact. In the next stage of complexity, in which more pycnidia are usually present, the walls in contact are replaced by common walls, each pycnidium still having its own ostiole; a rudimentary or a well-developed stroma may or may not be present in this stage. In the final stage, which occurs in the largest pycnidial groups, the combined pycnidia are replaced by a stroma with a single locule, one or more ostioles, and one, several, or no partial dissepiments.

The simplest non-stromatic isolated pycnidia are to be found wherever pycnidial formation is only beginning. Later, adjoining these pycnidia, or

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in the region close by, stromatic pycnidia and chambered stromata are formed in close juxtaposition. An old row or group of pycnidia exhibits

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Figs. 3, 4, 5.—Pycnidia in different stages. In fig. 3 the pycnidium is single and has no stroma. In fig. 4 the pycnidia form part of a small group, and the beginning of a stroma is indicated in the area marked with parallel lines. In fig. 5 the pycnidia are part of a long row. A stroma is present, and one pycnidium is of the stromatic type with a single locule and a partial dissepiment. × 48.
Fig. 6.—Spores, young. × 352.
Fig. 7.—Spores, mature. × 352.

at one time every grade of pycnidial development, the one basal stroma being common to all the pycnidia of the group, with the exception of the

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simplest younger ones, which in the ranked arrangement occur usually at either end of the row, and in the group arrangement at the periphery. Even for the youngest isolated members of any group, however, it is usually only a matter of time before they too are linked to the main body of the group by an extension of the common stromatic base. As already mentioned, this stromatic linking has not been found on the cones, where the pycnidia as a rule remain isolated from one another. The smaller size and the isolation of the cone-pycnidia are probably due to the greater hardness of the cones preventing the free ramification of the mycelium evident in the cortex of stem and branch, where, moreover, the hyphae are in direct communication with the water-supply of the xylem.

The size of the pycnidia varies from 250 × 250 to 800 × 800 micro-millimetres. If they are prevented by mutual pressure from expanding, their height exceeds their width. On the other hand, in the complex type with the locular stroma the width is usually increased, and may be several times as great as the height. The ostiole is shortly papillate. The cells composing the walls of the pycnidium are, on the whole, thin-walled, angular to rounded, black on and near the surface, fuliginous in the intermediate region, quickly paling to hyaline in the interior.

The spores (pycnospores) are ellipsoid-obovate to cylindrical; the ends are rounded, the lower one being usually narrower than the upper, at times markedly so. The colour of mature spores is fuliginous to fuscous, but they may be discharged when immature, in which case the colour is often not deeper than a golden-fuliginous. The septation varies. In each row or group of pycnidia there are many in which only the non-septate Sphaeropsis type of spore is to be found, but if the material is kept until some of the pycnidia approach maturity, which may mean for several months, the older pycnidia show at least a small number of uniseptate spores. In certain cases the septate Diplodia type of spore can readily be found, but this is rare except when the material had been kept for a considerable time. As a rule it is unusual to find the septate spore in pycnidia on the cones, where the Sphaeropsis type predominates instead. Spores when mature range in size from 24–37 × 14–18 micromillimetres, the average being about 32–35 × 14–16 mmm. The sporophores are hyaline, and rather stout, attaining a size of from 7–12 × 2.5–3.5 mmm., with an average of 10 × 3.5 mmm.

Nomenclature.—The above description closely approximates that given by Kickx for Diplodia pinea (Desm.) Kickx * occurring on needles of Pinus montana and P. silvestris in France, Belgium, and Italy. Kickx, and Karsten also, reported a form of the same fungus in the bark of P. silvestris and of species of Abies. The only major point of difference between the fungus of Kickx and the present one is that Kickx, and also Desmazière, who originally described it, failed to notice the locular stromatic type of pycnidium occurring on stem and branch when the fructifications are advanced in age. But as the material described was on needles, that in the cortex being only noticed in passing, the significance of the omission is lessened, especially as the stromatic type does not occur regularly in the present disease. Moreover, it is only within the last few years that the stromatic type has been recognized to be a usual phase in the development of the more common Sphaeropsis or Diplodia phases: and, as Desmazière first described his organism as long ago as 1842, the omission of these

[Footnote] * As given in Saccardo, Syll. Fung. vol. 3, p. 359, 1884.

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characteristics of the pycnidium is not surprising. Recently von Hoehnel has claimed that a fungus known earlier as Phoma macrosperma Karst. is merely Diplodia pinea (Desm.) Kickx at the stage when its spores are still hyaline. More recently again, Petrak, after insisting on the validity of the genus Botryodiplodia for members with grouped pycnidia and spores over a certain size, gave a detailed description of Phoma macrosperma Karst., adding to it details of structure which had not been given in the original description, and including in particular a greater spore-size than had been given for Diplodia pinea (Desm.) Kickx. This discrepancy he has overlooked, or has perhaps considered it insignificant (seeing that this character may show considerable variation), and, on the strength of his own previous assertions concerning the validity of the genus Botryodiplodia, has transferred Diplodia pinea (Desm.) Kickx to Botryodiplodia pinea (Desm.) Petr. Since then, although the generic limits of the Diplodiae are being more closely examined as knowledge of the developmental variations exhibited by members of the group is acquired, the genus Botryodiplodia as Petrak understands it still stands unchallenged.

The fungus attacking Pinus radiata and P. muricata in New Zealand is therefore referred to Botryodiplodia pinea (Desm.) Petr.

Control of the Disease.

As the fungus as a rule works downwards from the tip of the stem or branches, control of the disease requires only the removal and destruction of the parts infected. The mycelium may reach some inches below the fructifications, and also below the lowest sign of unhealthiness in the tree; and the cut should therefore be made about 2 ft. below the lowest sign of the disease, the cut surface being then protected from parasites by a coating of Stockholm tar.

Literature Cited.

1884. P. A. Saccardo. Sylloge Fungorum, vol. 3, p. 359.

1913. T. Petch. Repr. from Annals Royal Bot. Gardens Peradeniya, vol. 4, p. 445. (Ref. consulted: Zeit. f. Pflanzenkrankheiten, vol. 23, p. 242, 1913.)

1914. J. J. Taubenhaus. ‘Phytopathology, vol. 4, p. 47.

1915. — Amer. Jour. Bot., vol. 2, pp. 324–31. Review in Centralbl. f. Bakt. Parasitenk. und lnfektionskr., 2 Abt., 51 Bd., p. 527.

1915. F. von Hoehnel. Sitzb. Ak. Wiss. Wien, vol. 123, p. 84.

1919. W. B. Grove. Journ. Bot., vol. 57, pp. 206–10. (Ref. consulted: Bot. Abstracts, vol. 3, p. 365, 1920.)

1922. F. Petrak. Annales Mycologici, vol. 20, pp. 306–8.

1923. — Annales Mycologici, vol. 21, p. 332.

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Three Fungous Diseases of Salix in New Zealand, and some Saprophytic Fungi found on the same Hosts.

[Read before the Nelson Philosophical Society, 27th August, 1924; received by Editor, 5th December, 1924; issued separately, 6th March, 1926.]

Plates 811.

Among the commonest and most beautiful shade and ornamental introduced trees in the Nelson district are willows of both the “weeping” (Salix babylonica) and “crack” (Salix fragilis) varieties. For some time it has been noticed that several diseases, probably of fungous origin, are becoming prevalent on many of these trees. In February, 1923, an investigation was commenced upon these diseases. It was found that, though in some cases both species of willow were attacked by the same fungus, the worst disease of each host was caused by a different fungus—e.g., by Marssonina on the weeping-willow and Macrophoma on the crack-willow.

On material collected for examination from many different localities from Wakapuaka to Tasman numerous saprophytic fungi were also found.

Three of the worst diseases of the willows in this district, and the fungi causing them, are described in the present paper, and notes on a number of the saprophytic fungi found on the same hosts are included.

Marssonina salicicola (Bres.) P. Magn. on Salix.

The most serious disease is caused by the fungus Marssonina, which, though it occurs on both species of Salix, is chiefly found on S. babylonica. The parasite attacks the tree with great vigour, causing considerable injury and loss, and throughout the Nelson district practically no tree of this species has escaped infection. Many have lost branches and are dying.

(In April, 1925, since this paper was in print, specimens of the disease on Salix babylonica were collected in Ward, Marlborough.)

The disease was also noticed by the writer in a mild form on weeping-willows at Mitcham, South Australia, in January, 1924, and at Aldgate, South Australia, in January, 1925.

Pathological Changes in Salix babylonica.—The effect of the parasite on this host is most, noticeable. In trees which are only mildly attacked the leaves become covered, principally upon the lower surface, with pale reddish-brown spots which have a purple tinge when fresh; a few brown cankers may be seen on the twigs. When badly diseased the tree almost completely loses its “weeping” habit; the leafy branches, instead of being long and pendulous, are much shorter and tend to grow more upright, until, at a distance, the tree might almost be mistaken for one of the upright varieties (Plate 8).

This effect on the host is due to the shortening of the internodes in the diseased “weeping” branches, the average distance between any adjoining two of the ten nearest the tip in a number of diseased branches being 0.8 cm., as compared with 2.4 cm. in an equal number of healthy specimens. This leads to an increase in the number of twigs, which are produced closer

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together and are shorter. Weeping branches may be 1 m., 1.5 m. long, or more, while on badly attacked trees there may be no true weeping branches, but in their place a number of small twigs, a few to 30 cm. long, bearing leaves at fairly crowded nodes. The twigs become distorted with cankers, and tend to curl upwards.

When the twig is first infected a small black elliptical spot appears upon it, which rapidly spreads (Plate 9, fig. 1). Upon this spot whitish acervuli are produced beneath the epidermis of the host, which they raise and finally rupture, setting free the hyaline two-celled spores. As the infected tissue of the twig dies, it dries, cracks, and is broken off by another layer of spores developing below. In this manner several layers may crack off in succession. The canker spreads along and round the twig, and may join other cankered areas, until a large part is involved (Plate 9, fig. 2). The infected leaves become covered with spots, described on pages 60 and 61 (see Plate 9, fig. 3). On badly diseased twigs the leaves, all of which are often infected to some extent, are usually very much smaller than those on healthy branches, averaging 5.9 cm. in length as compared with 12.5 cm. Instead of hanging pendulous against the twigs, they tend to curl and grow more at right angles to it; this is often due to infection of the petiole, causing it to blacken and shrivel. The midrib may also be infected, when elliptical or elongated black spots appear similar to those on the young twigs and petioles. The stipules are often attacked as the leaves unfold, when they drop off at a very early age, while normally they remain at the base of the leaf long after it is mature. Eventually the leaves become yellowish, and fall prematurely. It was observed in early spring that the young leaves were infected immediately on the opening of the leaf-buds (Plate 9, fig. 4), infection probably being caused by spores from cankers upon the living twigs of the tree, from bud-scales (Plate 9, fig. 5), and from dead leaves which had been diseased the season before and remained on the ground. Examination of these during the winter and early spring showed numerous acervuli still producing a large quantity of spores.

Spots similar to those on the leaves, bearing typical acervuli and spores, were also found on the catkins of the weeping-willow.

Pathological Changes in Salix fragilis.—The same fungus as the above occurs on the crack-willow, but much more rarely, and then with less disastrous effect. Blackened areas may be formed along the young twigs, and cankers are sometimes produced on the older ones; but the main effect is seen on the leaves, which become thickly covered on the upper surface with dark-brown or black circular or irregular spots, which differ from those on the weeping-willow by being smaller and more numerous. On the lower surface of the leaves of the crack-willow faint small reddish-brown irregular spots may sometimes be seen; these at times become confluent. The spots are more numerous on the upper surface of the leaf than on the lower. They produce acervuli and spores similar to those on the weeping-willow.

Identity of the Fungus.

Genus.—The fungus under discussion was named Marssonia by Fischer in 1874. But, as P. Magnus (1906, p. 88) pointed out, this name had already been used for the genus Marssonia Karst. of the Gesneriaceae, so he altered it to Marssonina P. Magn., under which title the genus is now known.

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Rabenhorst, Saccardo, Engler and Prantl, and other authorities classify Marssonina P. Magn. (= Marssonia Fisch.) as being “exclusively found parasitic on leaves,” and thus distinguish it from the similar genus Septomyxa, which is “saprophytic on stems and fruit.” But Diedicke (1913, p. 540) observed that this was not an accurate basis of classification, as species of Marssonina had been found on twigs also. He suggested that a more exact distinction would be made by limiting Septomyxa to those species with a stromatic conical acervulus in which the production of conidiophores occurs typically on the side of the acervulus, and by extending Marssonina to include twig-infecting forms with a flat subepidermal acervulus bearing conidiophores over the basal disc.

In the present instance the identity of the fungus upon the leaves of the willow with that upon the twigs, as is exhibited in the similarity of general appearance, shape of the acervulus, and shape, size, and structure of the conidia, serves to confirm Diedicke's statement as to the identity of leaf-and twig-inhabiting forms.

Species.—The only mention of a Marssonina on the twigs of the willow is made by Fukushi (1921, p. 1), who found a disease in Japan which he thought was “probably due to Marssonina.” If this should prove to be so, it may be found to be the same species as that occurring on the twigs of the willow in New Zealand. Furthermore, the local species does not agree with the single species of Septomyxa recorded on this host. As for the species of Marssonina found on the leaves, the present species does not agree in all respects with any recorded on the willow, though strongly resembling three.

The Fungus on Salix babylonica.

The spots may be on either surface of the leaf, but are principally on the lower surface, where they are circular, not sharply outlined, up to 1 mm. in diameter, light purplish-brown (reddish when dry), at first scattered, then crowded and confluent until in many cases the whole lower surface of the leaf is covered (Plate 9, fig. 3). Where present on the upper surface the spots are smaller, dark-brown, and scanty. Upon the spots are numerous white acervuli. These are at first circular, often becoming confluent, varying in size from 60 to 210 mmm.,* the average size when mature being between 150 and 200 mmm., lifting up and breaking through the epidermis and exposing the conidia on very short hyaline conidio-phores, which are produced over the surface of the flat acervulus. The acervulus may be slightly sunken in the centre, due to the collapse of the dead host cells below. The conidia are club-or pear-shaped, frequently curved, two-celled, the lower cell being much the smaller of the two, hyaline, contents granular or guttulate, 11–16 mmm. long by 3–7 mmm. broad (text-fig. 1). On the young twigs (text-fig. 2) the acervuli of the first layer formed subepidermally are similar to those on the leaves, but as the canker develops, splitting the cortex and producing new acervuli on the successive surfaces exposed, the acervuli are more irregular in shape and size. Throughout, however, the spores are identical with those on the leaves.

As mentioned, there are three species of Marssonina that distinctly resemble the species under consideration—e.g., M. salicicola, M. rubiginosa,

[Footnote] * In this article the contraction “mmm.” is used for micromillimetres.

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and M. nigricans. Briefly stated, the species found differs from Marssonina salicicola (Bres.) P. Magn. in the spots on the leaves being amphigenous, principally hypophyllous, and in having some spores smaller; from Marssonina rubiginosa (Ell. & Ev.) P. Magn. in the spots on the leaves not being “more definite on the upper surface,” and in often having larger spores which are not always acute below; and from Marssonina nigricans (Ell. & Ev.) P. Magn. in having larger spots on the leaves, the spots not being edged with a narrow tawny margin, and in the acervuli being numerous and white.

The species of Marssonina found upon Salix in Nelson agrees in all important features with Marssonina salicicola (Bres.) P. Magn., and is therefore referred to that species, in spite of the spots on the leaves being described as “epiphyllous,” whereas in the Nelson specimens they are amphigenous.

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Figs. 1, 2.—Marssonina salicicola (Bres.) P. Magn. on Salix babylonica L. Sections through an acervulus with spores. Fig. 1, on leaf; fig. 2, on young twig. × 330.

This species has hitherto been recorded only upon the leaves of willows, but it is most probable that in the present instance the diseases of leaves, twigs, and catkins are caused by the same fungus. The acervuli and spores produced are identical on the different parts of the tree. Slight differences in the size and colour of the spores can easily be accounted for by the differences in host-tissue in the different parts of the tree.

Marssonina salicicola on the leaves in New Zealand varies according to the species of willow upon which it is found. When occurring on Salix babylonica it varies towards Marssonina rubiginosa, the reddish-brown spots being more numerous on the lower surface, though the spores are usually larger than in M. rubiginosa and not always acute below. When it occurs on Salix fragilis it varies towards Marssonina nigricans, the spots

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being blackish and more numerous on the upper surface of the leaves, but the acervuli produced are neither scanty nor dark as they are in M. nigricans. However, in the two cases, on Salix babyloninca and Salix fragilis, the acervuli and spores resemble each other so closely that it is doubtless the one species of fungus occurring on both willows, the variation mentioned being due to the different hosts.

Since the three species resemble one another so closely, it will probably be found that M. rubiginosa and M. nigricans are merely variant forms of M. salicicola, the variation being dependent, as in the species under discussion, upon the species of Salix upon which they occur.

The fungus found in Nelson upon the different parts of both S. babylonica and S. fragilis is therefore referred to Marssonina salicicola (Bres.) P. Magn.

Macrophoma Salicis Dearn. & Barth. on Salix fragilis.

One of the more important fungus diseases of the crack-willow (Salix fragilis) in this district is caused by a species of Macrophoma. It was first noticed on the 8th February, 1923, at Appleby, and subsequently at Wakapuaka, Atawhai, and Nelson.

Effect on Host.—The infected leaf becomes discoloured at the tip. This discoloration, reddish-brown in hue, soon spreads over the upper end of the leaf, being somewhat sharply limited by a straight or semicircular line from the adjoining tissue, which, however, becomes rather yellow, showing that the fungus is encroaching and the cells dying. Examination of this brown tip shows very minute blackish-brown swellings, which indicate the position of the pycnidia (Plate 11, fig. 2). The tip of the leaf dies and becomes dry, as later does also the whole of the infected area, the remainder of the leaf becoming yellow, falling prematurely. Trees with this disease lose all their leaves when uninfected trees are still green (Plate 10).

It was noticed that the leaves on trees infected with Macrophoma were as a rule much smaller than those on uninfected trees, frequently being only from one-half to one-third the size. This may be due directly to the presence of the fungus, or indirectly to premature defoliation due to the disease weakening the vigour of the tree in previous seasons. It is quite common to see a branch every leaf of which has a reddish-brown tip (Plate 11, fig. 1).

On some of the tips of both living and dead leaves infected with Macrophoma, Pleospora herbarum was also found.

Identity of the Fungus.

The pycnidia of the fungus are distributed more or less densely, are very minute, 100–170 mmm. broad, globose or flattened-globose, deeply sunken in the tissue of the leaf, amphigenous, having a short broad papilla, which slightly raises the epidermis and opens by a circular pore, wall somewhat thin, concolorous with the dying tissue of the leaf, hyaline inside; spores (pycnospores) oblong with rounded ends, continuous, hyaline, granula, 16–20 × 6–8 mmm. (most 20 mmm. long), sporophores short, about 6 mmm. long, filiform. Parasitic on living leaves of Salix fragilis.

No record has been found of any species of Macrophoma parasitic on any part of the willow or saprophytic on the dead leaves. Of the

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several species recorded as saprophytic on dead wood of various Salix sp., the one found here most resembles Macrophoma Salicis Dearn. & Barth., in the description of which, quoted below, the points of dissimilarity to the Macrophoma in question are printed in italics.

Macrophoma Salicis Dearn. & Barth., Myc., vol. 9, 1917, p. 352.

“Pycnidia thickly distributed, sometimes gregarious or seriate, cortical, rupturing the epidermis in a cleft or stellate manner, 0.25–0.5 mm. in height and diam. Ostiola thick, short or longer up to 0.3 mm. Conidia hyaline, continuous, ovoid to oblong or fusoid, 12–16 mmm. × 6–9 mmm. On dead twigs of Salix exigua Nutt., Billings, Montana.”

It will be seen that the species found in Nelson differs from Macrophoma Solicis chiefly in the smaller size of its pycnidia, in lacking the (only occasionally) greater length of ostiole, and in the manner in which the pycnidia rupture the epidermis of the host. But in many cases in which a fungus occurs on different parts of a tree it is usual to find considerable variation in the size of the fungus fructifications according to the part in which they are formed, the pycnidia formed on leaves being smaller, as a rule, than those on twigs or branches of the same tree. Similarly, in pycnidia in which the ostiole is developed only until it reaches the surface of the host, as in the present case, it is to be expected that the length of the ostiole would, on the whole, be shorter in pycnidia formed on thin leaves than in those formed on a branch, where they may occasionally be more deeply buried. The manner of the rupture of the host surface as the pycnidia push through is again largely a local peculiarity, dependent chiefly on the arrangement of the resistant or less-resistant kinds of host-tissue in the immediate vicinity of the pycnidia, and also to a certain extent on the manner of grouping of the latter—i.e., whether they are in groups or series, and whether scattered or dense.

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Fig. 3.—Macrophoma Salicis Dearn. & Barth. on leaf of Salix fragilis L. Section through a pycnidium with spores. × 300.

Of greater significance is the difference in size, and sometimes in shape, of the spores; but as spore sizes in common occur in the two sets of material, and as there is no other species of Macrophoma on the willow

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which resembles so closely that found in Nelson, the latter is regarded as Macrophoma Salicis Dearn. & Barth. occurring parasitically, instead of saprophytically, as previously recorded.

Gloeosporium Capreae Allesch. on Salix fragilis.

A species of Gloeosporium on Salix was found by the writer in Nelson on the 21st February, 1923. This is the commonest fungus growing parasitically on the crack-willows (Salix fragilis) in this district, and it also occurs infrequently on Salix babylonica.

Effect on Host.—The fungus occurs only upon the leaves, forming upon the upper surface small round or irregular somewhat raised spots, greyish-white with a narrow dark-brown margin, varying from 0.5 mm. to 2 mm. in diameter, very often confluent and forming large irregular patches often up to 1.5 cm. long and exceptionally to 3.5 cm.; sometimes, especially when large and near the margin or tip of the leaf, a large part of the diseased area may drop away from the leaf; there is a tendency for the blotches to follow the veins of the leaf, so that long narrow patches may frequently be found along the midrib and larger veins (Plate 11, fig. 3). The sports are covered with minute black dots where the acervuli break through the cuticle. The under-surface of the leaf becomes discoloured with dark-brown areas corresponding to the spots above, but the acervuli are borne principally upon the upper surface.

Identity of the Fungus.

The acervuli are crowded, brown, small, 80–120 mmm. in diameter, often confluent, in which case they are larger, erumpent, lifting up and breaking through the cuticle; conidiophores light-brown, 15–20 × 1–2 mmm., bearing small oblong to oval, hyaline, one-celled conidia, 5–12 × 3–6 mmm., often 2-guttulate (text-fig. 4).

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Fig. 4.—Gloeosporium Capreae Allesch. on leaf of Salix fragilis. Section through an acervulus with spores. × 330.

Gloeosporium Salicis West. is the most common species on the willow occurring in America and Europe. The descriptions of this species given by Saccardo (Syll. Fung., 3, p. 711) and Rabenhorst (Bd. 1, Abt. 7, p. 500) are very scanty, but Potebnia (1910, p. 77) published a more detailed account describing two kinds of spores, macroconidia 14–16 × 5–7 mmm. and microconidia 4–5 × 0.5–1 mmm., while Dearness (1917, p. 360) gives the spore range as 15–23 × 7.5–10 mmm. The spores are usually slightly

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Weeping-willow (Salix babylonica L.) badly diseased with Marssonina salicicola (Bres.) P. Magn. Note loss of “weeping” habit, except in branch at top right hand, which is still more or less normal.

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Marssonina salicicola (Bres.) P. Magn. on Salix babylonica L.
Fig. 1.—Dark spots on young twig producing the first acervuli
Fig. 2.—Older twig covered with cankers.
Fig. 3.—Lower side of leaf covered with spots
Fig. 4.—Young twig showing the stage of first infection, which takes place immediately after the opening of the buds.
Fig. 5.—Part of twig in fig, 4 enlarged to show infected bud-scales and canker on the stem, from which infection of the young leaves has probably taken place.

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Macrophoma 'Salicis Dearn. & Barth. on a crack-willow (Salix fragilis L.) which is losing all its leaves early, while a healthy crack-willow in the left background is still green.

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Fig. 1.—Macrophoma Salicis Dearn. & Barth. killing tips of leaves of Salix fragilis L. The small round spots scattered over the leaves are caused by Gleosporium Capreae Allesch.
Fig. 2.—Macrophoma Salicis Dearn. & Barth. on tip of leaf of Salix fragilis L. An arrow indicates the position of some of the numerous small pycnidia in the infected area.
Fig. 3.—Gleosporium Capreae Allesch on leaf of Salix fragilis L. Note the tendency of the pustules caused by the fungus to follow the veins of the leaf, and of the diseased areas to fall away from the leaf. The minute black dots on the blotches indicate the acervuli.
Fig. 4.—Gleosporium bullata n. sp. on dead twig of Salix babylonica L. On the surface of the twig may be seen the pustules caused by the beaks of the perithecia raising and piercing the epidermis. The round bases of the perithecia some of which are dry, showing the concave manner of collapsing, are exposed on the underside of the bark, which has been partially stripped off; while the depressions left by them may be seen on the wood.

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curved and form short curved white cirri. In all these points this common, species differs from that which occurs in the Nelson district, while, on the other hand, a less common species, Gloeosporium Capreae Allesch., is very similar.

Gloeosporium Capreae Allesch. (Sacc., Syll. Fung., 14, p. 1010.)

Spots epiphyllous, large, indeterminate, covering almost the whole leaf, brownish-grey then becoming pale; acervulus epiphyllous, minute, in dense clusters of various shapes, covered by the epidermis, fairly often confluent, black; conidia oblong, straight or slightly curved, obtuse at both ends, hyaline, continuous, often guttulate, varying in size, 6–16 × 2–4 mmm.; conidiophores narrowed above, hyaline, continuous, 15–20 × 1–1.5 mmm.

On still-attached or fallen leaves of Salix Capreae, Munich, Bavaria.

If this diagnosis is compared with that given for the local species it will be seen that they differ in two points. In the first place, the conidiophores in the local species are light-brown in colour; and, in the second place, while the measurements of conidia of average size coincide in the two cases, those at the extreme of the range tend to be longer and narrower in Gloeosporium Capreae. The fact that the latter species occurs on S. Capreae; while the local species was found on S. fragilis and S. babylonica, suggests that the difference may be due to the effect of the different hosts. The coincidence in the size of the average conidia of the fungus in the two cases, and the fact that, although there is colouring in the conidiophores of the local species, it is not deep, suggest that these variations are not sufficient to justify the exclusion of the local species from Gloeosporium Capreae.

Fukushi (1921, p. 1) has recorded a disease of Salix purpurea var. angustifolia in Japan due to Physalospora Miyabeana n. sp. and its conidial form Gloeosporium. No Physalospora has been found in connection with the local species, nor have any twigs been seen here with the cankers which, are so marked a feature of the disease in Japan. The appearance of the spots on the leaves caused by the Japanese species differs from that of the species found here. Also, the latter, on the whole, agrees more closely with Gloeosporium Capreae Allesch., as which species, therefore, it is, in the opinion of the writer, to be regarded.

Gnomonia bullata n. sp.

Perithecia black, scattered or more often gregarious (occasionally two perithecia may be found with the beaks united), sunken in the bark, which when wet may be stripped off the wood, carrying with it the globose perithecia, the round shining black bases of which are then exposed, the bases contracting inwards when drying (Plate 11, fig. 4); beak straight or bent, 100–390 mmm. long × 230–330 mmm. broad, narrowed above, perforating the periderm and raising the epidermis to form a little round pustule through which the tip of the beak projects (text-fig. 5), thickened and swollen flask-shaped below, consisting of small blackish compressed angular cells which are smaller and arranged in parallel-fibred structure bordering the central canal, which is usually bent spirally or may be straight; wall of the perithecia of two layers, the outer consisting of several rows of blackish angular thick-walled cells, the inner of several

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rows of thin-walled hyaline parenchymatous cells which are often compressed laterally (text-fig. 6).

Asci 70–100 × 12–17 (most 14–16) mmm., clavate, narrowed at the apex, sharply attenuate below, having in the young ascus a long filiform stalk which in the mature stage breaks off or becomes swollen and shortened though usually still basally attenuate; wall of ascus very thin, except at the apex, where it is thickened and opens by a pore, 8-spored, distichous; paraphyses absent (text-fig. 7).

Spores oblong-elliptical, some obtusely rounded at both ends, others somewhat rounded-fusoid, straight, more often unequal-sided or slightly curved, uniseptate, slightly constricted by the septum which is at or near the centre, hyaline, 16–23 × 5–7 mmm., contents granular or guttulate.

On dead twigs and branches, still attached, of Salix babylonica and Salix fragilis, Nelson, 15th September, 1923.

Throughout the district this is one of the commonest saprophytes occurring upon dead twigs and branches still attached to the trees, particularly on those twigs which have died the season before It occurs more frequently and abundantly upon the weeping-willow than on the crack-willow.

Species.—The species differs markedly from Gnomonia pleurostyla Auersw., the one species of Gnomonia recorded by Saccardo (Syll. Fung., vol. 1, p. 569) for the willow. G. pleurostyla is found on dead leaves; the asci are six-spored, the spores filiform lance-shaped, 50–55 × 2 mmm. Winter, in Rabenhorst's Krypt. Fl. (1887, p. 589), gave G. apiculata (Wallr.) Wint. on the willow, but expressed a doubt as to whether the species really belonged to this genus; and later Petrak (1921, p. 176) showed that it was identical with a species of Cryptodiaporthe on Populus, and that it should therefore be called Cryptodiaporthe apiculata (Wallr.) Petr. Petrak (1921, p. 180) has also transferred G. salicella (Fr.) Shroet. to Cryptodiaporthe sahcella (Fr.) Petr. E. M. Doidge (1924, p. 56) records Gnomonia sp. upon Salix sp. in South Africa, but gives no diagnosis and does not state the species. Moesz (1918, p. 60) describes G. salicina n. sp. upon dead branches of Salix alba. Unfortunately, this paper is not available. These are the only records of species of Gnomonia found upon Salix, and a comparison with the other species of Gnomonia found on different hosts shows that the species found here does not agree entirely with any of them, and in the absence of Moesz' paper it seems justifiable to separate it as a new species.

Relationship to other Fungi.—As this fungus so frequently occurs on dead still-attached twigs which were infected with Marssonina the season before, the question arises whether this may be the ascigerous stage of the fungus of which Marssonina is the conidial stage. This is not unlikely, as Klebahn (1905, p. 336) has demonstrated by cultures and ascospore infection the connection between Gnomonia leptostyla and Marssonina juglandis (on the bitter-nut Juglans cinerea). On the other hand, other species of Gnomonia have been shown to have different conidial forms, chiefly Gloeosporium, Asteroma, and Leptothyrium. Of these genera, Gloeosporium Capreae has been found on Nelson willows. According to Klebahn (1918, p. 317), the conidial forms of Gnomonia belong to the Melanconiaceae, so either Marssonina salicicola or Gloeosporium Capreae may prove to be the conidial form of Gnomonia bullata.

The fungus germinates very readily in water (four hours), (text-fig. 8), in nutrient agar (turnip, willow, &c), and in a solution of soluble substances

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Figs. 5–8.—Gnomonia bullata n. sp.
5. Perithecia. × 10.
6. Perithecium (type with bent beak); epidermis and bark indicated. × 120.
7. Asci. × 330.
8 Spores germinated in water—(a) after four and a half hours, (b) after eight and a half hours. × 330.

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from the willow. Healthy cultures of vegetative growth were readily obtained and watched to see if conidial stages would be produced, with a view to proving whether or no this Gnomonia were connected with the Marssonina so common on the same trees, or with the Gloeosporium common on Salix fragilis. So far none of the cultures have produced spores. It is also intended to undertake inoculation experiments to see if one of the parasitic diseases may be produced by this common saprophyte.

Petrak (1921, pp. 176, 180) has proved that several recognized species of Gnomonia belong to the genus Cryptodiaporthe, and the question arises whether the one under consideration may also belong to the latter genus, as in some cases two perithecia were found with their beaks partially or completely joined and having a common mouth, and there is always a marked thickening round the beak, which suggests the possibility of a rudimentary stroma, of which, however, there is no other sign. A great quantity of material has been available, and careful examination has not furnished any evidence in favour of this view. Except in the instances mentioned, perithecia were separate, although crowded; the type of perithecium and the structure of the ascus have the marked characteristics of Gnomonia, in which genus the fungus has been placed.

Some Saprophytic Fungi Found on the Willows in the Nelson District.


Cryptodiaporthc salicella (Fr.) Petr.

Dead twigs of Salix fragilis. Toitoi Valley, Nelson, 25th October, 1923.

(?) Leptosphaeria Salicinearum (Pass.) Sacc.

Dead leaves of Salix babylonica. Nelson, 20th June, 1923.

Leptosphaeria Salicinearum is the only species of this genus recorded on willow-leaves, but no measurements of asci or spores are given in its description. The specimens found here agree essentially with Saccardo's description on such points as he mentions. The spores are distichous, 3-septate, not or hardly constricted, fusoid, mostly curved, 16–29 × 4–7 mmm., yellowish or light olive. The local species is placed tentatively under Leptosphaeria Salicinearum (Pass.) Sacc., although lack of detail in the description given by Saccardo prevents final confirmation.

Metasphaeria orthospora Sacc.

Dead wood, Salix babylonica. Wakapuaka, 3rd July, 1923; Maitai Valley, 10th August, 1923.

Dead wood, Salix fragilis. Maitai Valley, 17th August, 1923.

The spores of the local specimens are more constricted at the median septum, but not at the other two septa, than in those described by Saccardo.

Pleospora herbarum (Pers.) Rabh.

Dead fallen leaves, Salix babylonica and Salix fragilis. Common throughout the Nelson district.

Dead tips of living leaves of Salix fragilis, the tips having been first killed by Macrophoma Salicis. Appleby, 8th February, 1923.

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Macrophoma Salicaria (Sacc.) Berl. & Vogl. (Text-fig. 9.)

Common on dead twigs, Salix babylonica. Wakapuaka, 12th February, 1923.

Common on dead twigs, Salix fragilis. Maitai Valley, Nelson, 12th September, 1923.

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Fig. 9.—Macronhoma Salicaria (Sacc.) Berl. & Vogl. on dead wood of Salix babylonica. Section through pycnidium with spores. × 240

Diplodia salicina Lev.

Dead twigs, Salix babylonica. Maitai Valley, Nelson, 21st September, 1923.

Dead twigs, Salix fragilis. Toitoi Valley, Nelson, 19th September. 1923. The range of spores was 20–28 × 10–13 mmm. The Sphaeropsis stage was often present.


Coryneum Salicis Togn.

Decaying leaves, Salix fragilis. Appleby, 1st July, 1923.

Dead twigs still attached to the tree. Toitoi Valley, 19th September, 1923.

The specimens found differ from the type in having in most cases only the lower cell pale, the upper being the same colour, fuliginous, as the middle cells.

Hyaloceras Saccardoi (Speg.) v. Hoehn. (Text-fig. 10.)

Dead twigs and leaves of Salix babylonica. Appleby, 8th October, 1923.

As the only species of this genus recorded on Salix is Hyaloceras excipuliformis Bubak, which differs from the species found in Nelson in general appearance and in having only three septa and smaller spores, while the local species agrees with the description of

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Hyaloceras Saccardoi (Speg.) v. Hoehn. upon Quercus except in the smaller details of measurement of cilium and stalk, the local species is therefore referred provisionally, because of the lack of relationship between the two hosts, to Hyaloceras Saccardoi (Speg.) v. Hoehn.

The measurements of the Nelson specimens are as follows: Conidia 16–22 × 6–7 mmm. (most 18–20 mmm. long), 4-septate, constricted; one cilium 12–24 × 0.5–1 mmm. on upper cell; persistent conidiophore 20–30 × 0.5–1 mmm.

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Fig. 10.—Spores of Hyaloceras Saccardoi (Speg.) v. Hoehn. on Salix babylonica. × 330.

Pestalozzia funerea Desm.

Dead twigs, Salix babylonica. Nelson, 19th September, 1923.

As the local specimens of Pestalozzia differ markedly from the species of that genus already recorded for the willow, while they have all the characteristics of the common species Pestalozzia funerea Desm., they are referred to that species, which, although occurring on a wide range of hosts, has not hitherto been recorded on Salix.

The writer's thanks are due to Dr. Curtis, of the Cawthron Institute, for assistance and advice, and to Mr. Davies, of the Cawthron Institute, for the photographs.

Literature cited.

Dearness, J., 1917. North American Fungi. Mycologia, vol. 9, pp. 345–64.

Diedicke, H., 1913. Noch einige “Leptostromaceen,” die Nectroideen, Excipulaceen, und Melanconieen. Ann. Myc., vol. 11, pp. 528–45.

Doidge, E. M., 1924. A Preliminary Check-list of Plant-diseases in South Africa. Bot. Surv. S. Africa, Memoir No. 6, pp. 1–56.

Engler, A., und Prantl, K., 1897. Die naturlichen Pflanzenfamilien, 1 Teil, 1 Abt.*

—, A., und Prantl, K., 1900. Die naturlichen Pflanzenfamilien, 1 Teil, 1 Abt.*

Fukushi, T., 1921. Physalospora Miyabeana n. sp. and its Conidial Form (Gloeosporium) injurious to Salix purpurea var. angustifolia in Japan. Ann. Phytopath. Soc. Japan, vol. 1, No. 4, pp. 1–11. (Reference consulted: Int. Rev. Sci & Pr. Agr., 1921, vol. 12, p. 1492.)

Klebahn, H., 1905. Cent. Bak., vol. 15, p. 336.

“, 1918. Haupt-und Nebenfruchtformen der Askomyzeten, pp. 178–317.

Magnus, P., 1906. Notwendige Umänderung des Namens der Pilzgattung Marssonia Fisch. Hedw., vol. 45, pp. 88–91. (Reference consulted: Ann. Myc., 1907, vol. 5, p. 294.)

Moesz, —, 1918. (Botanikai közlemények, p. 60.)

Petrak, F., 1921 Mykologische Notizen, 3, Ann. Myc., vol. 19, pp. 176–223.

Potebnia, A., 1910. Beiträge zur Micromycetenflora Mittel-Russlands. Ann. Myc., vol. 8, pp. 42–93.

Rabenhorst, L., 1887. Kryptogamen Flora von Deutschland, Oesterreich, und der Schiveiz: Ascomycetes, vol. 1, Abt. 2.

—, 1901–3. Kryptogamen Flora von Deutschland, Oesterreich, und der Schweiz: Fungi Imperfecti, vol. 1, Abt. 6–7.

Saccardo, P. A., 1882–1911. Sylloge Funqorum, vols. 1–20.

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A New Genus of the Hysterangiaceae.

[Read before the Wellington Philosophical Society, 1st October, 1924; received by Editor, 31st December, 1924; issued separately, 6th March, 1926.]

Plates 12,13.

While collecting in the Whakatikei Reserve, Paekakariki, Wellington, Mr. and Mrs. J. G. Myers obtained abundant specimens of a peculiar white fungus, which they handed to the writer for identification. On examination this species proved unique, and calls for a detailed description of its structure.

The plants were found growing on rotting wood partially buried in the humus of open grassy places in the forest. Specimens are white in colour, and vary greatly in size and shape. The most characteristic feature is the presence of numerous sterile lobes, springing from the dorsal surface of the peridium. These lobes are dull-white, and may be clavate, capitate, or fan-shaped, solitary or branched, and are at their bases attached to a common pulvinate or globular body, the peridium. This also is white externally, and is attached to the substratum by several coarse white rhizoids.

When a mature plant is sectioned longitudinally the lobes are seen to be sterile and frequently hollow, the gleba being present only in the basal portion or peridium to which the lobes are attached. The function of these lobes is unknown. The gleba may extend for some little distance into their base. It is olivaceous in colour, and is traversed by numerous stout trabeculae, which arise from a distinct sterile basal disc. It is gelatinous in consistency, as is the greater portion of the interior of the lobes.

Structure of the Mature Plant.

Peridium.—This consists of two layers, an outer thin cortex or exoperidium, and a thick, gelatinous endoperidium. The exoperidium consists of a layer, 0.5–1 mm. thick, of loosely-woven, intricately-branched, colourless hyphae. The hyphae of which it is composed remain distinct during the lifetime of the plant, and at no time are gelatinized, nor do they assume the form of a pseudoparenchyma. This layer surrounds the endoperidium. The outer few layers of hyphae are somewhat evanescent, and give to the plant a pruinose appearance.

The endoperidium lies within the exoperidium, and is sharply delimited from it. It consists of a thick (3–5 mm.) layer of hyphae, which in the mature plant is partly gelatinized. It forms the whole of the internal portion of the lobes. At the base it forms a pulvinate sterile disc (basal disc), from which arise numerous stout trabeculae, which tend to divide the gleba into numerous locules.

Gleba.—This is traversed by the trabeculae, which give off secondary branches, which in turn are divided until an intricately-branched network of tramal plates is produced, the spaces between being occupied by lacunae lined with the hymenium. The lacunae are minute irregular spaces, closely compacted together save in the centre of the plant, where they are less crowded, and are frequently separated by large irregular interspaces. The

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tramal plates consist of a central gelatinized layer (often absent near the terminal ends of the ultimate branches), with a layer on either side composed of small polygonal cells—the subhymenium—bearing on their proximal surfaces basidia closely compacted into a palisade tissue. The basidia are long, cylindrical, frequently branched, and each bears, on short, almost obsolete sterigmata, eight elliptical continuous spores. The spores are minute, hyaline, and uninucleate.


As a sufficiency of developmental stages has not been obtained, the account given below is incomplete, and in consequence somewhat disconnected.

Developing plants first become noticeable as minute globular nodules on the upper surface of the rhizoids. Sections at this stage show the whole to consist of loosely-woven hyphae. When the plant has attained to a size of 3–4 mm., differentiation of the different tissues commences. The plant at this stage is globular, save where it is attached at the base, where it is slightly flattened. A definite exoperidium is marked off through the remainder of the internal tissues becoming more compacted. In the latter a more compact zone of hyphae appears near the base: from it strands of compacted hyphae arise, differentiation proceeding in an upward and outward direction until several primary trabeculae are formed. These are partly surrounded by irregular and poorly-defined lacunae, due, no doubt, to rupture of the hyphae in the vicinity. Next, branches are gíven off from the trabeculae, and in turn these branch so that the loosely-woven network of the gleba is formed. Within the margin of these branches small lacunae arise, and around the inner surfaces of these appear the subhymenial cells, which soon give rise to basidia. No specimens have been obtained showing the first formation of these lacunae, so that it is not known whether the formation of the regular hymenium is preceded by production of occasional basidia from irregular scattered papillae, as in Gallacea (1924). Growth of the trabeculae and smaller branches continues in this manner until a well-developed gleba is produced, this tissue appearing in plants of a diameter of 6–7 mm.

Later lacunae form within the peripheral zone (here termed the “endoperidium”) lying beneath the exoperidium. Gelatinization of the trabeculae, central portions of the tramal plates, and endoperidium commences shortly after glebal differentiation, until at maturity all the glebal tissue, save spores, hymenium, and subhymenium, is gelatinized.

Development of Lobes.—These do not appear until glebal differentiation is well begun. They arise as small outgrowths from the dorsal portion of the endoperidium, and enlarge until they appear as distinct clavate or capitate bodies. They arise from the portion of the endoperidium immediately underlying the exoperidium, and are thus in all probability not derived from potentially sporogenous tissue. While still small they may again produce secondary and even tertiary lobes. Lobes may arise from any part of existing lobes, but generally arise from the vicinity of their attachment with the peridium. In large specimens the lobes become hollow, but no further stage of development than this has been observed. The plants are indehiscent, spores being released by gradual decay of the plant.

As no plants possessing these peculiar lobed structures have been described, the writer believes the genus to be undescribed, and proposes to name it Phallobata, on account of its phalloid-like spores and basidia, and the lobed character of the peridium.

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Fig. 1.—Selected plants, natural size: showing variations in size and shape of plants, and especially of the lobes so characteristic of the genus.
Fig. 2.—Section of plant, × 2. showing structure of lobes and gleba.

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Section of plant, × 6: en. endoperidium; ex, exoperidium; l, sterile lobed portion; bas, sterile base; tr, travecula.

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Phallobata n. gen.

Peridium sessile, indehiscent, variously shaped, crowned with 1-several large single or branched sterile lobes; of two layers—an external loosely-woven intricately-branched layer of hyphae (the exoperidium), and an inner thick gelatinized layer forming the body of the lobes; attached to the substratum by numerous coarse rhizoids.

Gleba permanent, consisting of numerous anastomozing gelatinized tramal plates, enclosing minute lacunae, divided by several stout trabeculae springing from a sterile base into several loculae; hymenium lining the free surfaces of the tramae. Basidia 8-spored, subclavate or cylindricäl, branched; spores borne on short slender sterigmata (or subsessile), continuous, hyaline or tinted, smooth, uninucleate. Cystidia and other aberrant cells absent.

Habitat Saprophytic upon decaying wood.

Phallobata alba n. sp.

Peridium white or greyish-white, depressed-globose, globose, elliptical, or tuberous, up to 30 mm. wide, 35 mm. high, crowned with 1-several sterile lobes: attached to the substratum by numerous coarse white rhizoids. Lobes springing from the apical portion of the peridium, cylindrical, clavate, capitate, pulvinate, or irregular, white, smooth or rugose, pruinose, dry, frequently hollow within.

Gleba olivaceous, traversed by numerous gelatinized hyaline trabeculae, arising from the sterile pulvinate basal disc, enclosed within the broad gelatinized endoperidium. Lacunae elliptical, ranging in size from 0.1 mm. to 2 mm. or more.

Spores smooth, continuous, almost hyaline, rounded at both ends, 2–3 × 1 mmm.

Habitat: Gregarious on rotting wood in grassy places in the forest.

Distribution: Forest reserve, Whakatikei, Paekakariki, Wellington, 45 m.; J. G. Myers, Mrs. Myers! 2nd June, 1924. J. C. Neill! J. G. Myers! 16th June, 1924.

Type collection. Herb. No. 1187.

Systematic Position of the Genus.

The earlier stages of development and glebal characters would place the genus in the family Hysterangiaceae, as defined by Fisher (1900). The 8-spored branched basidia and subsessile minute uninucleate spores would place it in the Phallales. It more closely resembles the genus Phallogaster Morg. than any other, but is separated on account of its being indehiscent, and in the possession of the queer lobes, so characteristic a feature of the plant. This genus, together with Phallogaster, would serve as a connecting-link between the Phallales and the Hysterangiaceae, and for this reason should be placed in an intermediate family. As such does not exist, it is retained in the Hysterangiaceae for the present.

Literature cited.

Cunningham, G. H., 1924. The Development of Gallacea Scleroderma (Cke.) Lloyd, Trans. Brit. Myc. Soc., vol. 9, pp. 193–200.

Fischer, Ed., 1900. Hymenogastrineae in Engler and Prantl. Nat. Pflanzenfam., 11*, pp. 296–313.

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Third Supplement to the Uredinaceae and Ustilaginaceae of New Zealand.

[Read before the Wellington Philosophical Society, 1st October, 1924; received by Editor, Editor, 31st December, 1924; issued separately, 6th March, 1926.]


Additional Species.

The collecting of the following species adds a fourth family to the Uredinaceae of New Zealand.


Teleutosori subepidermal. Teleutospores unicellular, catenulate, apedicellate, united laterally into columns or pulvinate masses; epispore coloured or hyaline, smooth; basidium external. Uredosori with or without peridia. Uredospores catenulate or borne singly on distinct pedicels. Aecidia present or absent.

This family, although not recognized by certain workers, is, I believe, sufficiently characterized by the catenate teleutospores to warrant retention.

Sydow (Mon. Ured., vol. 3, p. 502, 1915) places Chrysomyxa and Barclayella in the Chrysomyxeae, a subfamily of the Melampsoraceae.

Chrysomyxa Unger.
Ung., Beit. verg. Path., p. 24, 1840.
Melampsoropsis (Schroet.) Arth., Res. Sci. Congr. Bot. Vienne, p. 338, 1906.

Heteroecious. Cycle of development includes 0, I, II, III.


Spermogones immersed, flask-shaped.


Aecidia erumpent, with definite peridia. Aecidiospores globose to elliptical; catenulate; epispore hyaline, covered with coarse deciduous warts; germ-pores indistinct or wanting.


Uredosori with or without peridia, pulverulent, erumpent; peridia, when present, delicate, evanescent. Uredospores catenulate, apedicellate; epispore hyaline, verrucose; germ-pores indistinct.


Teleutosori erumpent, waxy, pulvinate, becoming velvety. Teleutospores catenulate, in simple or branched chains, unicellular, elliptical; epispore hyaline, smooth, thin; germ-pore apical, obscure.

Distribution: Europe; Asia; North America; India.

The aecidia, when present, occur on species of Picea; the uredo-and teleuto-spores on Empetrum, Pyrola, Rhododendron, and Ledum.

The uredospores occur in chains, and peridia may be present or absent; when present they are usually evanescent and difficult to detect, save when sections are made through nearly mature sori. Between the uredospores in situ large interstitial cells are present. The teleutospores are catenulate, and become compacted into pulvinate sori. They germinate without a period of rest, giving to the sori a characteristic velvety appearance.

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1. Chrysomyxa Rhododendri (DC.) de Bary. Ericaceae.

DB., Bot. Zeit., p. 809, 1879.

Uredo Rhododendri DC., Fl. Fr., vol. 6, p. 86, 1815.

Caeoma Rhododendri Link., Sp. Pl., vol. 2, p. 16, 1825.

Coleosporium Rhododendri Schroet., in Cohn's Beitr., Bd. 3, Heft 1, p. 56, 1879.


Spermogones amphigenous, flask-shaped, honey-yellow.


Aecidia hypophyllous, seated on elongate yellow spots. Peridium irregular, laterally compressed, 0.5–3 mm. long, 0.25–0.75 mm. high, margins hyaline, lacerate. Spores globose, subglobose, or elliptical, 20–44 × 14–20 mmm. * (Sydow); 17–45 × 12–22 mmm. (DB.); epispore hyaline, closely and finely verruculose, save for a smooth longitudinal strip along one side; 2–2.5 mmm thick; germ-pores indistinct.


Uredosori hypophyllous, seated on discoloured spots which are visible on the upper surface, minute, 0.2–0.5 mm. diam., circular or irregular, bullate, solitary, or more frequently in scattered groups of 3–15 or more, orange-yellow, ruptured epidermis inevident; peridium delicate, evanescent, hyaline. Spores catenulate, interstitial cells evident, obovate, elliptical or subglobose, 18–26 × 15–21 mmm.; epispore hyaline, closely and finely verruculose; 2–2.5 mmm. thick; cell contents orange-yellow, granular, with one prominent central nucleus; germ-pores indistinct.


Teleutosori hypophyllous, seated on small discoloured spots which are visible on the upper surface, minute, 0.2–0.5 mm. diam., circular, pulvinate, brownish-red, ruptured epidermis inevident. Spores catenulate, chains 80–130 mmm. long, shortly cylindrical or prismatic, 20–30 × 10–14 mmm.; epispore hyaline, smooth, 1 mmm. thick, with an annular thickening at the summit of the uppermost cell; germ-pore apical, obscure.

Host: Rhododendron sp. cult. On leaves. Herb. No. 1636. II. Stratford, Taranaki, W. Pettigrew! 4 Nov., 1924.

Distribution: Europe; Asia.

This species has been introduced from Europe with the host. Only the uredo stage has been collected, and the determination has been made from this. It agrees closely with the published descriptions and with the uredo stage of herbarium specimens in the possession of the writer.

The aecidia occur on Picea excelsa and P. obovata in Europe, the uredo-and teleuto-spores on numerous species of Rhododendron. The life-cycle is but imperfectly known; the teleutospores germinate in early summer, producing basidiospores which may infect the leaves of Picea. The resultant aecidiospores may infect Rhododendron leaves, and in these the mycelium apparently overwinters, producing uredo-and teleuto-spores the following summer.

Pucciniaceae (Tribe Phragmideae).
Kuehneola Magnus.
Mag., Bot. Centr., vol. 74, p. 169, 1898.

Autoecious. Cycle of development includes 0, II, III.


Spermogones subcuticular, flattened, hemispherical.


Uredosori erumpent, pulverulent, without peridia; of two kinds, primary and secondary. Primary uredosori usually associated with the

[Footnote] * In this article the contraction “mmm.” is used for micromillimetres.

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spermogones, aparaphysate; secondary uredosori frequently paraphysate. Uredospores borne singly on distinct pedicels; epispore rough, hyaline; germ-pores equatorial (?), obscure.


Teleutosori erumpent, aparaphysate. Teleutospores divided by transverse septa into 3-several cells; epispore coloured or subhyaline, smooth: germ-pores solitary, apical; pedicels hyaline, short, fragile.

Distribution: Europe; North and South America; India.

Members of this genus are in Europe confined to the Rosaceae, but in North and South America they occur in addition on the Malvaceae. The solitary New Zealand species has been introduced with the host.

The teleutospores characterize the genus, and appear as if several single teleutospores of some coronate species of Uromyces were superimposed one upon another, the lowest being attached by the pedicel to the sorus.

1. Kuehneola albida Magnus, I.c. Rosaceae

Chrysomyxa albida Keuhn, Bot. Centr., vol. 16, p. 154, 1883.

Uredo Muelleri Schroet, Krypt. Fl. Schles., vol. 3, p. 375, 1887.

Phragmidium albidum Lagerh., Mitth. Bad. Bot. Verh., p. 44, 1888

Knehneola Uredinis (Link.) Arth., N. Am. Fl., vol. 7, p. 186, 1912.


Spermogones epiphyllous, in small groups, seated on discoloured spots, depressed-globose, 0.1–0.2 mm. diam.


Primary uredosori epiphyllous, surrounding the spermogones, in crowded groups, 0.5–1.25 mm. across, usually somewhat elongate, often arranged in a circinate manner, bullate, pulverulent, orange-coloured, fading to pallid cream, surrounded by the ruptured epidermis. Secondary uredosori hypopyllous and caulicolous. frequently seated on discoloured spots which are visible on the upper surface, scattered, or more frequently in crowded irregular groups, circular, small, 0.1–0.5 mm. diam., bullate, pulverulent, lemon-yellow, bleaching to a pallid cream-colour with age, aparaphysate. Spores elliptical, obovate, or more rarely subglobose, 20–30 × 17–24 mmm.; epispore hyaline; moderately and bluntly echinu-late, 1–1.5 mmm. thick; cell-contents lemon-yellow, vacuolate; germ-pores indistinct (probably 3 or 4, equatorial—Arthur).


Teleutosori hypophyllous, scattered or in small groups, circular, 0.2–0.5 mm. diam., pulvinate, tinted yellow, ruptured epidermis inevident, Spores 5–13-celled, commonly 5–6, cylindrical or cylindrical-clavate, 85–110 × 18–24 mmm., each cell 17–40 × 15–24 mmm., trapezoidal, somewhat coronate above; epispore thickened from below upwards, 1.5 mmm. below, apex 3–5 mmm., hyaline or tinted yellow, smooth; pedicel persistent (or wanting), short, hyaline, delicate; germ-pore solitary, situate in one of the projections at the apex of each cell.

Host: Rubus fruticosus L. On leaves and stems. Herb. Nos. 246, 1637.


Tokomaru, Wellington, G.H.C., 6 Dec., 1920. Kauri Gully, Auckland, G.H.C., 2 Mar., 1921. Helensville, Auckland,E. Bruce Levy! 3 Feb., 1921. Opotiki, Auckland, M. Darey! 25 Oct., 1920. Te Puke, Auckland, R. Waters! 18 Feb., 1922. Wairoa, Hawke's Bay, E. H. Atkinson! 12 Oct., 1924.

Distribution: Europe; North America.

The uredo stage only has been collected in New Zealand; this, in a previous paper (Trans. N.Z. Inst., vol. 54, p. 690, 1923), was in error referred to Gymnoconia Pcckiana, the caeomata of which it resembles closely. Dr. B. O. Dodge (United States Department of Agriculture

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Washington), to whom specimens were submitted, wrote pointing out this error, and in addition forwarded material for comparative purposes. On examination and comparison of the specimens of Kuehneola albida kindly, forwarded by Dr. Dodge with New Zealand material I find the two to be identical. Gymnoconia Peckiana should therefore be deleted from the record of Uredinaceae occurring in New Zealand.

Additional Hosts.

Puccinia pedatissima G. H. Cunn. (Trans. N.Z. Inst., vol. 54, p. 673, 1923)

Host: Ourisia macrocarpa Hook. f. Herb. No. 1633. On leaves:


Roaring Creek, Arthur Valley, Otago, E. H. Atkinson! 18 Jan., 1920.

This host is endemic and confined to the South Island (Cheeseman, Fl.N.Z., p. 549, 1906).

Puccinia Elymi Westnd. (Trans. N.Z. Inst., vol. 55, p. 1).

Host: Bromus sterilis L. Herb. Nos. 1638, 1639. On leaves. II-III. Roadside, Ashburton, Canterbury, J. C. Neill! 18 Oct., 1924.

This host is an introduced plant, common throughout the agricultural areas of New Zealand.


Additional Species.

Tolyposporium Woronin.
Wor., Abh. Senck. Nat. Ges., vol. 12, p. 577, 1882.

Sori forming a granular spore-mass at maturity, usually in the inflorescence, commonly in the ovaries, formed of numerous spore-balls. Spore-balls consisting of few or many spores, opaque, dark-coloured. permanently united, sterile cells absent.

Spores bound together by ridged thickenings or folds of the epispore, dark-coloured, irregular in shape, smooth or roughened on their free surfaces; germination similar to that of Ustilago.

Distribution: Europe; Asia; North and South America; Ceylon; India; Africa; Australia.

The following endemic species is the only one that has been collected in New Zealand. The genus is fairly well represented in Australia, for McAlpine (Smuts Aus., p. 186, 1910) records seven species. Species of the genus are confined to the Gramineae, Cyperaceae, and Juncaceae. The genus is characterized by the very firm spore-balls, their dark colour, and the amon of individual spores by folds of the epispore. It is separated from Thecaphora by the more compact nature of the spore-balls, and chiefly on account of the much darker colour; from Sorosporium, in the balls being permanent, although in this latter genus certain species occur in which the balls are also permanent. In such a case they are separated by the presence of (in Tolyposporium) the peculiar ridged folds of the epispore.

1. Tolyposporium littorale n. sp. (Fig. 1.) Cyperaceae.

Sori in ovaries, usually destroying all in the inflorescence, concealed within the glumes, subglobose to elliptical, 1–2 mm. long, black, at first agglutinated in firm masses, becoming apparent only upon rupture of the

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perigynium, when pulverulent and granular. Spore-balls of 20–50 or more spores firmly and permanently united by ridged folds, irregularly elliptical, subglobose or angular, 48–110 × 40–70 mmm., black, opaque.

Spores irregular in size and shape, obovate, cuneate, or angular, 15–25 × 14–20 mmm.; epispore dark brown, smooth on the united faces, covered with coarse deciduous tubercules on the free, 1–2 mmm. thick where smooth, up to 8 mmm. thick on free surface, often with lateral winged projections at margins of free surfaces.

Host: Cladium Huttoni T. Kirk. In ovaries. Herb. No. 1640. Seashore, Tauranga Harbour, Auckland, J. C. Neill! G. H. C., 20 Jan., 1924. (Type.)

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Fig. 1.—Tolyposporium littorale G. H. Cunn., on Cladium Huttom T. Kirk: spore-ball (× 320) and spores (× 500).

The host is endemic and confined to the North Island, where it is apparently common on the margin of certain lakes (Cheeseman, Fl. N.Z., p. 787, 1906).

I have been unable to germinate the spores. The spore-balls of this species are so firm that separation of the spores is not possible unless the balls are first boiled in lactic-acid solution, or treated for some time with caustic-soda solution. The spores appear as if covered on their free surfaces with large, irregular, deciduous tubercules, the crevices between giving to the spores a decidedly areolate appearance The spores are not unlike those of T. lepidosperma McAlp., in that they are thickened and much roughened on their free surfaces, but the spore-balls are quite distinct, being much larger in size, more irregularly angular in shape, and spores are much larger.

Farysia Raciborski.

In a former paper on the Ustilaginaceae of New Zealand (Trans. N.Z. Inst., vol. 55, p. 414, 1924) the name Elateromyces Bubak was used for a genus characterized by the presence of “elaters” in the sori. I find that the name Farysia was used by Raciborski for a species (F. javanica Racib.) possessing this same character, and, as Raciborski's name has priority, the three species included in Elateromyces should be cited as under:—

Farysia Raciborski, Bull. Acad. Sci. Cracovic, p. 354, 1906.

  • F. nigra n. comb.

    Syn. Elateromyces niger G. H. Cunn.

  • F. olivacea (DC.) Sydow, Ann. Myc., vol. 17, p. 42, 1919.

    Syn. Elateromyces olivaceus (DC.) Bubak.

  • F. endotricha (Berk.) Sydow, l.e.

    Syn. Elateromyces endotrichus (Berk.) G. H. Cunn.

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When in England recently I examined the type of this last species (Herb. Kew., No. 4748), and find that the description given in my former paper is inaccurate, the spores especially being wrongly described. The following amended description has been drawn up from the type specimen:—

Farysia endotricha (Berkeley) Sydow, l.c. (Fig. 2.) Cyperaceae.

Sori on peduncles and main axes of the inflorescences, not in ovaries, jet-black, compact, elliptical, up to 20 mm. long, 8 mm. wide; elaters short, up to 5 mm. long, stout, black, curled, giving to the sorus a decidedly velvety appearance.

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Fig. 2.—Farysia endotricha (Berk.) Syd.: a. spores from Gahnia pauciflora T Kirk; b, spores from type material at Kew; both × 500.

Spores globose or shortly elliptical, 12–18 mmm. diam.; epispore dark-olivaceous or dark chestnut-brown, 1 mmm. thick, covered with numerous flattened irregular tubercules, the crevices between giving a distinct areolate appearance to the spores.


Gahnia sp. On peduncles. Type: Herb. Berk., Kew, No. 4748.

Auckland, Sinclair!

Gahnia pauciflora T. Kirk. On peduncles. Herb. No. 2190. Silverstream, Upper Hutt, Wellington, H. Hamilton! Jan., 1924.

The latter host is endemic, and not uncommon throughout both Islands (Cheeseman, Fl. N.Z., p. 793, 1906) With this emended description it will be seen that the areolate nature of the epispore markings separates this from the other two species recorded from New Zealand. The figure given by Berkeley is inaccurate in that the elaters are much exaggerated, and the spores do not arise from them as his figure represents.

In the Farysia (Ustilago) endotricha folder at Kew (labelled “U. tricho-phora Kunze”) are numerous collections. In working over these I found that only the type collection, consisting of one specimen from which Berkeley prepared his illustration, is of this species, all others being Farysia olivacea (DC.) Syd. This applies also to the “co-type” specimen of Cooke, which was acquired when his herbarium was bought by the Kew authorities.

Additional Host.

Ustilago Avenae Jens. (Trans. N.Z. Inst., vol. 55, p. 405, 1924).

Host: Avena sativa L. In inflorescences. Herb. Nos. 1496, 1501 Tapanui, Otago, J. C. Neill! G. H. C., 12 Feb., 1924. Winton, Southland, J. C. Neill! 15 Feb., 1924.

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Urocystis Anemones = Tuburcinia Anemones.

In a recent paper Liro has shown, as a result of examination of the original material, that Tuburcinia Fries and Urocystis Rabh. are identical. As Tuburcinia has priority, this name should replace Urocystis. The genus should therefore be cited as under:—

Tuburcinia Fries, Syst. Myc., vol. 3, p. 439, 1832; and the species on p. 430 of my previous paper,—

T. Anemones (Pers.) Liro, Ann. Univ. Fennicae Aboensis ser. A, tom. 1, No. 1, p. 55, 1922.

Liro has split the original into several species; but to me, judging from the descriptions given, all are races of the one species.

Botanical Notes, including Descriptions of New Species.

[Read before the Auckland Institute, 25th November, 1924; received by Editor, 28th November, 1924, issued separately, 6th March, 1926.]



During recent years many alterations have been made by European pteridologists in the classification and nomenclature of ferns. As far as New Zealand genera are concerned, various species of Aspidium and one Nephrodium are now classed under Polystichum, the remainder of the Nephrodia. and one Polypodium being known as Dryopteris. Polypodium tenellum is now Arthropteris tenella; Davallia novae-zealandiae is Leptolepia; Asplenium umbrosum is restored to its place as Athyrium; A. japomcum is Diplazium japonicum; Lomaria is Blechnum; and so on.

For some time I have been in correspondence with Dr. Carl Christensen, of Copenhagen, who is the European authority on ferns, and from his notes much of the following information is compiled. Quotations from his notes are followed by his initials (C. C.).

1. Dryopteris punctata C. Chr.

Syn. Polypodium punctatum Thunb (Manual, 1008); P. punctatum var. rugulosum (Thomson).

“It is a true Hypolepis, not at all a Dryopteris as listed in my Index, where I followed Engler and Prantl. In the Supplement to the Index it is

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placed under Hypolepis. The typical Hypolepis punctata is very different from your plant, which is Polypodium rugosulum Labill. (not rugulosum, as commonly spelled). I call attention to the fact that the genus Hypolepis totally lacks true scales'—i.e., flattened trichomes; the vesti-ture is always formed by hairs—i.e., trichomes formed by one cell or a single row of cells. By this character Hypolepis may always certainly be known from Dryopteris.”(C. C.)

2. Dryopteris parasitica (of New Zealand authors).

Syn. Nephrodium molle (Manual, 1006).

“The specimen sent [gathered at Kaitaia, North Auckland] resembles closely a common form widely spread in tropical countries. It is, however, scarcely typical parasitica (L.), and should probably be named D. dentata (Forsk.) C. Chr. The typical parasitica of Linnaeus is Chinese and, inter alia, characterized by the frond not being narrowed downward.” (C. C.)

3. Diplazium japonicum Bedd.

Syn. Asplenium japonicum Thunb.

“The form from Kaitaia corresponds closely with the original description and figure of D. congruum Brack, and I do not hesitate to refer it to that species, but I am rather in doubt how it may be distinguished from D. japonicum without having access to a large number of specimens from central Polynesia. I should prefer to name your form provisionally D. japonicum, suggesting that it is presumably that form described as D. congruum Brack.” (C. C.)

There is another form occurring in the far north with longer pinnules which are more or less deeply toothed, and a form gathered at Te Whaiti, between Rotorua and Hawke's Bay, is still more deeply lobed. These Te Whaiti plants were gathered by Dr. Petrie and Mr. H. B. Matthews. A plant which they brought to Mr. H. B. Dobbie has developed pinnules more deeply lobed and a more open character. It appears to me to correspond very closely to the description of the late T. Kirk's Asplenium umbrosum var. tenuifolium. Dr. Christensen has not yet seen this form, but I hope to forward specimens to him shortly.

4. Asplenium lamprophyllum sp. nov.

Rhizoma breviter repens 7–10 mm. longum, vel densum, pallide viride, squamis paucis fugitivis. Stipes 15–20 cm. longus, teres, alte canaliculatus a parte superiore, a parte postica atro-purpureus. Frondes 10–55 cm. longae, 10–15 cm. latae, oblongo-lanceolatae, pallide virides, subacuminatae, supra lucentes, erectae, submembranaceae, bipinnatae; rhachides teretes, a parte superiore canaliculatae. Pinnae numerosae, 5.15 cm. longae, 2.5–5 mm. latae, lanceolatae a parte inferiore pinnatae, ¾ pinnatifidae; rhachis secundaria late alata. Pinnulae 15–30 mm. longae, late cuneatae, alte lobatae, lobis oblongis obtusis. Sori obliqui angusti 3–9 mm. longi, costis propinqui, raro prope margines.

A beautiful light-green brightly-shining fern, often growing in close masses 3–10 ft. in diameter. Rhizome shortly creeping, 7–10 mm. long, or stout and compact, pale green with a few scattered dark linear-lanceolate scales. Stipes 15–20 cm. long, terete, deeply grooved in front, usually

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purplish-black and scaly at the back, green in front. Fronds 10–55 cm. long, oblong-lanceolate, slightly acuminate. light green, brightly shining on upper surface, erect, submembranous, bipinnate; rhachis terete, grooved in front, with a few deciduous scales. Pinnae numerous, ascending below, horizontal above, 5–15 cm. long, 2.5–5 cm. wide, lanceolate, sub-acuminate or acuminate, partly pinnate at base, the greater part pinnatifid: secondary rhachis broadly winged, with a few dark scales. Pinnules 15–30 mm. long, broadly cuneate, deeply lobed, lobes oblong, obtuse. Sori oblique, narrow, 3–9 mm. long, close to the midrib, seldom near the margin.

Habitat.—North Island: Maungataratara, Whangaroa County; Aponga, Maungatapere, Whangarei County; Great Barrier Island; vicinity of Auckland City; Mauku, Papakura (H. B. Dobbie!) Franklin County; Te Aroha, Piako County.

This fern has for many years been accepted as “a form of Asplenium bulbiferum.” Not being satisfied with this determination, I submitted specimens to Dr. Christensen, who remarks: “This is a very interesting fern, and it is a wonder that till now it has escaped the attention of pteridologists, for it certainly is not A. bulbiferum but a species of Asplenium, as far as I can see, undescribed. A close comparison of the specimens with several dried and living specimens of A. bulbiferum (in our Botanical Garden) show that scarcely one character is common to them. I conclude, then, that it is an undescribed species. If you agree in this I think you will describe and name it.”

Asplenium lamprophyllum usually occurs on or among rocks in shady woods; occasionally it is found on the lower parts of the trunks of trees, and sometimes on clay banks. Especially fine specimens are to be seen in the small woods among the lava blocks at Mount Wellington, near Auckland I have not yet seen specimens from farther south than Te Aroha, but probably it will be noted in other districts.

5. Blechnum capense (of New Zealand authors).

Syn. Lomaria capensis (Manual, 980); L. procera (Thomson's Ferns, 67).

“The three varieties received are certainly very interesting. A close study in the field is very necessary before finally deciding that these forms really belong to a single species. Is your fern really conspecific with Blechnum capense, a South African fern ? B. capense is said to be distributed over the whole Tropical and the South Temperate Zone, but I have little doubt that a comparative study based upon rich material will affirm my idea that several distinct species of more definite geographical range can be distinguished. This being the case, your fern should probably be named B. procerum (Forst.).” (C. C.)

6. Hypolepis.

“The three forms sent as Hypolepis tenuifolia, H. Petrieana, and Dryopteris punctata C. Chr. are as under:—


H. tenuifolia Bernh.


H. punctata (Thunb.) Mett. Syn. H. Petrieana Carse.


H. rugosula (Labill.) J. Smith. Syn. Dryopteris punctata (see No. 1 of this paper).

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“H. punctata differs from the other two in its pale, almost glabrous stipes and rhachises. Hooker and Baker united H. punctata and H. rugosula under Polypodium, hence comes this confusion.” (C. C.)

Owing to this confusion, no New Zealand student of ferns appears to have been aware that H. punctata was indigenous. It was classed as “merely a form of H. tenuifolia.” I was not satisfied with this view, so that when Dr. Petrie drew my attention more closely to this fern I did not hesitate to describe it as a new species (Trans. N.Z. Inst., vol. 50, p. 64, 1918).

“H. rugosula is distinguished from H. punctata by its red rhachises and its viscosity.” (C. C.)

Of H. tenuifolia there are at least two quite distinct forms, which may eventually have to be separated. One is a tall robust plant with a rhizome and stipe 10 mm. or more in diameter. The other is much smaller.

7. Polypodium dictyopteris Metten.

“Rhizome small, short, knot-like … emitting woolly rootlets, some of which creep and produce new tufts of fronds.” (Manual, 81).

On lava rocks on the Auckland Isthmus the juvenile form of this fern produces a creeping rhizome 3–15 in. long, with tufts of fronds, or single fronds at intervals. As a rule by the time the plant attains maturity the long rhizome has disappeared, leaving the tufts or single fronds as separate plants.

Mr. J. W. Brame, of Aucklamd, who has devoted many years to the study of New Zealand ferns referring to this fern, writes,—

“Our last outing resulted in my obtaining a suite of specimens better than any I have before seen. Among them I have noted as follows:—

“(1.) Roots branching and producing single fronds or tufts of fronds at nodes of both main roots and branches of same; these all juvenile. Roots upwards of 12 in. long.

“(2.) Two or more tufts of fronds connected by a root; these both adult and juvenile.

“(3.) Adult or mature fronds, well spored, from 2 ½ in. to 14 in. long, and some of them 1 in. wide when fresh.

“(4.) In some instances fronds, instead of being ‘tufted at top of rhizome,’ seem to be produced in series, with short intervals between, along the rhizome, as in some of the dwarfed forms of P. australe.”

8. Gleichenia.

“I call attention to my Index Fil. Supp. [113], where I, after examination of type specimen of G. circinata Sw. (Herb. Stockholm), have changed the names thus, and, no doubt, correctly:—

“1.G. circinata Sw. (1806), not authors. Syn. G. dicarpa R. Br. (1810).

“2. G. microphylla R. Br. (1810). Syn.G. circinata of authors.

“Your series should thus be named—

“1. G. circinata with its var. hecistophylla.

“2. G. microphylla (G. circinata of yours).

“May G. alpina be maintained as a distinct species ?” (C. C,)

Much confusion has been caused through the mixing-up of these two species, and many writers in New Zealand appear to infer that there are

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great difficulties in distinguishing them. After a considerable amount of study in the field I find that they have very distinct characters which will enable the student to tell the one from the other almost at a glance.

The following are some of the main differential characters:—

[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]

G. circinata (syn. G. dicarpa). G. microphylla (syn. G. circinata).
Scales and hairs Reddish-brown, often very dense Dark reddish-brown, less dense
Colour Pale green, upper surface dull, usually very glaucous below Dark green, upper surface polished, pale-yellowish below, less often glaucous.
Shape Suborbicular. apex truncate or rounded, pouched Triangular on a broad base, inner margin curved, giving apex an outward direction, apex subacute.
Sporangia 1—2, rarely 3, dull yellowish-white 1—4, rarely 5, bright shining yellow.

Environment does undoubtedly affect the outward appearance of these two species, as suggested by Dr. Cockayne in Some Noteworthy New Zealand Ferns, p. 55. In the shade the pouch-like segments of G. circinata (formerly known to us as G. dicarpa) are almost flat, the scales and woolly hairs much less developed. But the shape remains. In circinata the segments are invariably suborbicular, in microphylla triangular with the inner side curved outwards; the sori of the former always show same with 2 sporangia of a dull yellowish-white, the latter almost invariably some with 4 sporangia of a bright-yellow hue.

The majority of writers on New Zealand ferns appear to infer that the type form of G. circinata Sw. (= G. dicarpa of authors) does not occur in the Dominion. This I have long held to be a mistake. When I sent specimens of what I took to be the type form and variety hecistophylla to Dr. Christensen he had no hesitation in recognizing the one as the type, the other as the variety. I sent specimens to the Sydney Botanic Gardens. Mr. E. Cheel, to whom they were submitted, writes: “The specimen of G. circinata marked ‘Type?’ seems to me to closely approach our Port Jackson plants of G. dicarpa R. Br., and may not be specifically different from that species, but the specimen labelled ‘G. circinata var. hecistophylla (A. Cunn.)’ seems to me a well-marked form.”

A specimen received from Sydney labelled “G. dicarpa” can readily be matched in this country, so that I think we may take it that G. circinata (G. dicarpa of authors) is here represented by the type plant and its two varieties.

G. circinata var. hecistophylla differs from the type in being smaller and more densely woolly. This seems to me to be the result of environment, as the variety is found in cold, sour land, exposed to sun and wind. The form known as var. alpina seems to pass by gradual steps from var hecistophylla, some of the subalpine forms of which it is almost impossiblc to distinguish from alpina.


9. Lepyrodia Traversii F. v. Muell.

The habitat of this very local plant is given in the Manual as “Swamps between Hamilton and Ohaupo, middle Waikato district. Chatham Islands: Abundant in peaty swamps.”

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Since this was written the plant was discovered on a peat bog near Kaitaia, in the far north of Auckland.

Recently I found it occurring very plentifully in peat bogs at Tauhei, in the Morrinsville district, about twenty miles east from the Ohaupo Swamp. Here it was very much larger than I had seen it elsewhere. The Manual gives it “2–5 ft. high.” The Tauhei plants, growing among tall Leptospermum scrub, are 4–10 ft. high, the stems being thick in proportion.

Lepyrodia will probably be extinct before very long. The Kaitaia Swamp has been drained and the plant is no longer seen. I understand that drainage and fires have destroyed the Ohaupo plants, and doubtless the same fate will shortly overtake those at Tauhei. It is more than probable that other colonies of Lepyrodia may exist among the thousands of acres of still undrained swamp in the middle Waikato area. The distribution of this plant at stations so widely separated as the Chathams, the middle Waikato area, and Kaitaia is noteworthy.

10. Pittosporum umbellatum Banks and Sol.

In woods near Whangaroa Harbour the leaves of juvenile plants are deeply ternately lobed, passing gradually on the upper part of the branch to the ordinary elliptic-oblong or lanceolate-oblong form. I have not noticed this heterophyllous form elsewhere.

In the same woods I found two mature trees, in fruit, on which the majority of the leaves are rather narrower than usual and are deeply lobed ol bluntly toothed.

11. Pittosporum pimeleoides R. Cunn.

In Trans. N.Z. Inst., vol. 53, p. 365, 1921, is an article on P. cornifolium A. Cunn. by D. Petrie, in which it is shown that that plant is in all probability unisexual, that the male flowers are produced in 6–8-flowered umbels, and that the female flowers are solitary or rarely in pairs.

In studying P. pimeleoides I find that the same thing occurs. The male flowers are produced in terminal umbels of 4–8 flowers, each flower on a long slender pedicel. The female flowers are solitary or in pairs on much shorter pedicels. It is to be noted that binate female flowers and capsules are much more common in this species than in P. cornifolium.

12. Pseudopanax Lessonii C. Koch.

In the Manual this species appears under the heading “Leaves of young plants not markedly different from those of old ones.”

Some years ago the late Mr. Cheeseman told me he suspected there was a form of this plant (or possibly a distinct species) in which the juvenile leaves differ materially from the mature ones. Since then I have kept a lookout, and here give the result of my observations.

In the juvenile plants of what I take to be typical P. Lessonii, as described in the Manual, the leaflets hardly differ in shape from those of mature plants. In my specimens, from shrubs 2–5 ft. high, the leaflets are 2–4 in long, rather more deeply toothed.

The other form is very-different. In the mature state the leaves are usually 5-foliolate, leaflets oblong-lanceolate, linear-lanceolate, to linear-oblong, 3–6 in. long, more coriaceous than in the type form.

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The juvenile form of this plant is often difficult to distinguish from the trifoliolate form of P. crassifolium, though, as a rule, the leaflets are less rigid.

The plants from which these specimens were obtained are 1–6 ft. in height, none showing signs of having flowered at any time. I have divided them into two sets—(a) those having 3–5 or 5–7 leaflets, and (b) those having 1–3 leaflets.

(a.) Shrubs 4–6 ft. high. Leaflets 3–5, 5, or 5–7, broadly or narrowly linear, 4–10 in. long, ¼–1 in. wide, distantly, bluntly toothed, an incurved hook at the end of each tooth.

(b.) Shrubs 1–4 ft. high (possibly merely younger states of above). Leaflets 1–3, narrow linear, 3–10 in. long, ¼–1 in. wide, toothed as in (a).

So far I have seen two only of this form in flower. My specimens were gathered in various places between the North Cape Peninsula and Coromandel Peninsula.

Mixed with these is a juvenile form which the late Mr. Cheeseman suggested to Mr. John Bishop, of Titirangi, might perhaps prove to be the Chatham Island P. macrocarpa.

13. Dracophyllum Matthewsii sp. nov.

Frutex D. latifolio A. Cunn. affinis, sed in partibus omnibus minor. Truncus ± 4.5 m. altus. Rami graciles non verticillate. Folia 10–30 cm. longa, 12–25 mm. lata. Panicula 10–28 cm. longa, semper pendula. Flores quam in D. latifolio minores et minus densi, rubro-purpurei vel nigropurpurei. Capsula 2 mm. in diam. Floret tempore verno.

A shrub or small tree allied to D. latifolium A. Cunn., but smaller. The branches are not whorled as in that species, and are much more slender. At the extremities, where the leaves occur, they are only ⅛–¼ in. in diameter. Leaves 4–12 in. long, ½–1 in. wide at broadest part, tapering to an acute point. Panicles 4–12 in. long, always drooping. Rhachis and branches yellowish-green in colour, as also sepals. Petals purplish-red or almost black; anthers white tinged with pink.

D. latifolium A. Cunn. var. Matthewsii Carse, Trans. N.Z. Inst. vol. 43, p. 238, 1911.

Hab. North Island: Dry ridges in forests from Mongonui County to Great Barrier Island and Waitakerei Ranges. September-October.

This plant bears a general resemblance to D. latifolium, but the smaller size in all its parts, the pendulous panicle, the difference in colour of its flowers, and the differing flowering-period (September instead of January) distinguish it clearly from that species.

In the preparation of this paper I am greatly indebted, for specimens, information, and assistance, to Dr. Carl Christensen, of Copenhagen; Dr. Petrie, of Auckland; Mr. E. Cheel, of Sydney; and to Messrs. H. B. Matthews, H. B. Dobbie, J. W. Brame, and J. Bishop; to all of whom I gladly tender my thanks for their valuable assistance.

Literature consulted.

Brown, R. Prodromus.

Cheeseman, T. F. Manual of the New Zealand Flora. Wellington, 1906.

Illustrations of the New Zealand Flora. 2 v. Wellington, 1914.

Cockayne, L. Some Noteworthy Ferns.

Dobbie, H. B. New Zealand Ferns. Auckland, 1921.

Petrle, D. Trans. N.Z. Inst., vol. 53, p. 365, 1921.

Thomson, G. M. The Ferns and Fern Allies of New Zealand. Melbourne, 1882.

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Dactylanthus Taylori.
Order Balanophoreae; Tribe Synomorieae.

[Read before the Hawke's Bay Philosophical Society, 17th July, 1924; received by 31st December, 1924; issued separately, 6th March, 1926.]

Plates 1417.

The root-parasite Dactylanthus Taylori, although known in a slight way since the year 1855 (1), has hardly received that attention from botanists which it deserves. The plant is fairly widely distributed over the North Island, to the north of latitude 40°c S., ranging in bush country from near the sea-coast to inland, where it is found at an elevation of over 3,500 ft. The paucity of references connected with the life-history of the Dactylanthus in this country must be set down to difficulties in the way of a systematic study of this plant, as until quite lately the date of flowering was not known, and few botanists have the opportunity of seeing the growths in situ.

The illustrations given in (3), pl. 30, figs. 1 and 2, show attachments of tiny Dactylanthus rhizomes to the host, also the form of the host attachment in the case of well-grown host attachments on separation from a rhizome. So also pl. 31, fig. 1, illustrates pistillate spadices as they appear when growing on a bank in the gloom of the bush, and fig. 2 shows rhizomes with pistillate flowers as found attached to a host; pl. 32 further shows the spadices of a staminate flower surrounded by a perianth made up of sepals and petals in different whorls.

In Cheeseman (5), vol. 2, pl. 178, there is a rhizome with pistillate spadices only, attached to a host: fig. 1 illustrates a single staminate spadix, and fig. 10 gives a spadix showing both male and female flowers. There is no capitulum at the end of a scaly stem terminated by a perianth containing from 18 to 28 spadices, each spadix crowded with anthers, as illustrated in the margin, and as shown in pl. 32 of my paper (3).

As illustrated by Dactylanthus, by Cordyceps, by Bagnisia, and by another undescribed specimen of a parasite in my possession, the study of parasitism presents many attractive features that offer suggestions for inquiry in several directions in relation to animal and vegetable life. These parasites have allured me to Taupo and the inland bush country adjacent to Opepe on many occasions, owing to the fact that the time of their flowering was unknown. In the case of Dactylanthus only isolated specimens of staminate inflorescences have been seen, and much remains to be learnt of the other parasites named.

Since the issue of Cheeseman's Illustrations of New Zealand Flora (1914) up to March of this year it had not been possible for me to go to Taupo

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except in early spring, midsummer, and midwinter. Each time Taupo was visited, a visit was also made to Opepe, in the hope of finding something new and rare in connection with parasitism. Pistillate spadices of Dactylanthus with fruit were usually obtainable, but never the staminate spadices, although the evidence was plain enough that the staminate flowers had bloomed.

Having decided to spend three months at Taupo, commencing in late December, 1923, it became possible to visit Opepe and the surrounding bush country where Dactylanthus and other rare plants are to be found. January and February, however, were unusually dry, and there was not a trace of fungoid growth or of active parasitic life. Towards the close of February, however, the weather broke, and soaking rain continued off and on during March until the date of leaving for Napier, on the 23rd. Although the weather was unfavourable, it was decided to visit Opepe on the 21st March with my native companion, Tamati te Kurapae, chief of the Ngatituwharetoa, who knows the bush country well. We arrived at our destination, eleven miles from Taupo, at 8.30 a.m.

Although hundreds of spadices of Dactylanthus have been examined by me from time to time, I have not seen a spadix with staminate and pistillate flowers as shown in pl. 178, fig. 10, of Cheeseman (5). This must have been sent to Cheeseman by Mr. Frank Hutchinson, jun., from the Puketitiri Bush, although rhizomes and inflorescences were sent by me to Mr. Cheeseman on many occasions.

The forenoon was spent in the bush seeking for Bagnisia and Cordyceps, but our quest for the former was fruitless, though Cordyceps were abundant.

Rain drove us from the bush in the early afternoon, and we sought the shelter of a camp of native rabbiters, where we obtained information about Cordyceps and also obtained a number of live specimens, which the Maoris call the makaroa. The rain having ceased, we started for the gully where in former years the late Mr. A. Hamilton, of the Dominion Museum, and I had spent many delightful hours in the collection of botanical specimens, &c., and where also the late Mr. Cheeseman, F.L.S., of the Auckland Museum, had accompanied me in our search for Bagnisia and Dactylanthus. On approaching the glen I confess that my thoughts were not unmixed with sadness, for only a few weeks earlier Dr. Donald Petrie, M.A., F.L.S., an ardent botanical friend had visited me at Taupo but he was not well enough to go to Opepe.

When the Maori in whose whare we had taken shelter from the rain learnt that we were seeking for waewae-te-atua, by which name Dactylanthus is known to them, they asked to accompany us, for, although they had heard of the plant from others, they had not seen it growing. They knew the pua-te-reinga, which Taylor says was the native name of the Dactylanthus, but which the Taupo natives apply to Bagnisia Hillit, family Burmanniaceae: see (5), vol. 2, p. 198.

The country extending from the plateau near Opepe, 2,300 ft. above sea-level, gradually descends in the direction of Lake Taupo in a series of deep canal-like valleys for 1,000 ft. The valleys run lakeward like the spokes of a wheel from the circumference towards the centre, and each valley in its upper part represents a glen, sometimes wider and sometimes deeper than others, but all are waterless. On the banks grow many kinds of shrubs, such as Aralia, Pittosporum, Weinmannia, Aristotelia, Carpo-detus, &c., among ferns and lycopods, and in summer most of them are truly fairy dells. On entering the glen there was a strong perfume noticeable,

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which was at first set down to the moistness of the air; but it grew stronger as we moved down the valley towards the banks from which specimens of Dactylanthus had hitherto been collected. Although the place was so familiar to me, I had never had the slightest suspicion that the valley floor was a veritable garden-meadow of Dactylanthus. Opening out, along the dry floor of the valley for a chain or more, appeared hundred of flowers in clumps. Some were in bud, some half-opened, some in full bloom, measuring 5 in. across, the whole forming a picture that was quite new to us all. As shown (Plates 14,15, and16, fig. 1), the flowers are all staminate, and are raised from 1 in. to 3 in. above the surface. They formed a picture never before seen by us, and expressions of pleasure and of wonder were frequent. I was reminded of the bed of Bagnisia plants found less than half a mile away; but the Dactylanthus flowers were far more numerous, and much larger, and covered a considerable area. The perfume was overpowering, and reminded me of my first find of Dactylanthus, more than twenty-five years ago, at Matarau, near East Cape. Near-by small rhizomes attached to small host-roots were found. These had pistillate flower-heads, which are much shorter than those attached to the rhizomes carrying staminate flower-heads, but are not buried in the ground as is the case with the staminate-bearing rhizomes. All the rhizomes with staminate flowers are buried some inches in the earth, and are much larger and different in shape from the rhizomes producing pistillate flowers. A number of clumps of the flowers with rhizomes and host attachments were carefully dug up and packed for removal as specimens, and the following is a description of the various parts:

Host Roots.

Roots of Pittosporum and Aralia are the ones most frequently attacked. Those with staminate rhizomes, which are buried some inches below the surface near the place of attachment, are sometimes 1 in. or more in thickness. The rhizome when fresh can easily be separated from the host by means of boiling in water. The expanded surface of the host very closely resembles the petals of a pansy (Plate 16, fig. 1), and suggests a powerful suction as between host and rhizome. There is no trace that the host suffers from the attack of the parasite, but from specimens collected tiny rhizomes attach themselves to a root not at the end but on the side. When fully grown, however, the rhizome always represents the terminal end of the root. Plate 17, fig. 1, shows the appearance of a host attachment after separation from the rhizome.


The rhizomes containing pistillate flower-heads are much smaller and flatter, and possess warty-looking shoots over the whole surface. The flower-stem is much shorter than in the case of staminate flower-heads. The staminate rhizomes are buried in the ground and grow to a much larger size than the disc-like rhizomes bearing pistillate flowers. When dug up most of the specimens of rhizomes containing staminate flowers had shoots growing in bundles not unlike bundles of young asparagus. These do not appear to reach maturity, but die away following the decay of the spadices forming the staminate inflorescences. From the inside of the rhizome there radiates from the centre, where it is attached to the host, a series of smooth ridges corresponding to the depressions in the petal-like surface of the host. These extend to the cortical layer on the outside, where tiny shoots are formed. Under the microscope these shoots are of two kinds

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one containing tiny leaves, the other a pollen-like material having the appearance of resin. When fresh, some of the rhizomes can be cut with a knife and are not unlike pith. The colour is purple, and the taste is neutral. The rhizome hardens by exposure, and when soaked in water gives it a deep-golden colour.


A. Staminate.—A flower-shoot from a staminate rhizome varies in length from 4 in. to 8 in. Scales appear sparsely along its whole length, increasing in number and form up to the perianth. As shown in Plate 16, fig. 1, the perianth consists of sepals and petals, the former being wider and shorter, the latter linear-oblong, the upper tip showing enlarged corners like a cat's ears. The petals vary in colour from a light to a dull purple, and the sepals are streaked here and there with a faint purple colour. The number of leaves in the perianth (see Plate 16, fig. 2) varies from 18 to 28, as seen. The perianth encloses 20 or more spadices. Each spadix is not unlike a canoe-paddle or beaver's tail, the upper part of which is crowded with anthers as shown in the illustration. The anthers are very numerous, and are attached to the spadix in a small curve from left to right. The anthers split lengthwise, and are full of pollen, which resembles under the microscope the resin-like material in the warty shoots of a rhizome. At the place of attachment of spadix and petal there is a tiny claw (Plate 16, fig. 1, c). In the centre of the spadices is a small abortive pistil.

B. Pistillate.—A pistillate inflorescence consists of a scaly stem shorter and less robust than the staminate one. It is terminated by a perianth enclosing 30 or more spadices (Plate 16, fig. 3, c). Each spadix consists of a number of pistillate flowers along its whole length. The style and stigma persist up to the time when the seed is ripe, as do also two tiny scale-like attachments to the ovary. The fruit is a small nut, hard and dry. On the removal of the skin-covering the white portion has the appearance of boiled rice, or coconut. Under the microscope no embryo could be found. In all the specimens examined no single spadix was found containing both male and female flowers as shown in Cheeseman's illustration (5). The flowers are dioecious, sometimes monoclinous.


1. R. Taylor. New Zealand and its Inhabitants. London, 1855.

2. T. Kirk. Trans. N.Z. Inst., vol. 28, p. 493.

3. H. Hill. Trans. N.Z. Inst., vol. 41, p. 437.

4. J. Grant. Trans. N.Z. Inst., vol. 43, Proc. 3, p. 98.

5. T. F. Cheeseman. Illustrations of New Zealand Flora. Wellington, 1914.

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Dactylanthus Taylori: Rhizome and pistillate flowers.

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Bed of Dactylinthus at Opepe, Taupo, showing opening staminate flowers. Rhizcmes below the surface.

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Fig. 1.—Staminate flower showing perinanth: a, stamens closed: b, stamens open, with mature anthers; c, single spadix with sepal and petal attachments.
Fig. 2.—Staminate flower and spadices with perianth: a, perianth; b, pollinated stamens
Fig. 3.—Pistillate flower: a, flowers; b, seeds; c, spadices.

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Fig. 1.—Host attachments of rhizomes. la, a single attachment.
Fig. 2.—a, pistillate flowers, showing attachments of flowering-stem to rhizome; b, pistillate spadix; c, seeds.

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Geology of the Rakaia Gorge District.

[Read before the Canterbury Philosophical Society, 4th June, 1924; received by Editor, 30th July, 1924; issued separately, 6th March, 1926.]

Plate 18.




Physiographical Description.




Permian or Triassic Sediments.


Cretaceous Igneous Rocks.


Cretaceous Sediments.


Tertiary Igneous Rocks.


Pleistocene Sediments.





Literature cited.

1. Introduction.

The district dealt with in this paper is that immediately surrounding the gorge through which the Rakaia River passes before issuing from the mountainous region of the Southern Alps on to the Canterbury Plains—or that part of the valley of the Rakaia River which lies between the Mount Hutt Range, Fighting Hill, and Round Top, the southernmost peak of the Rockwood Range. The general structure of the area is that of an aggraded, glaciated valley, into the floor of which the river has re-entrenched itself. In the central part of the district the river in thus lowering its bed has encountered a barrier of resistant Tertiary and pre-Tertiary rocks, and has cut through it a winding gorge.

References to the Rakaia Gorge district are found in many of the reports of the early geologists of this country. The most comprehensive accounts of the stratigraphy and general geology of the area are those of Haast (1871), in a report on the Malvern Hills, and Cox (1884), in a report on the Selwyn and Ashburton Counties. Since the work of Cox, however, there has been no further investigation. An attempt is there fore made in this paper both to give a more detailed account of the district than has hitherto been published, and to examine some of the problems which the outcrops present in the light of the newer conceptions of the geological history of New Zealand which have evolved during the last forty years. [ unclear: ]

Thanks are due to Professor Speight for assistance both in the field and in preparation of this paper; to Mr. P. G. Morgan for his kindness in obtaining for me the rock analyses included below; and to Mr. S. Sylvester and Mr. T. A. Phillips for facilities while engaged in the field-work.

2. Physiographical Description.

The map (fig. 2) indicates the arrangement of the drainage of the district. The river-bed is flanked on both sides by extensive series of

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terraces. Above and below the gorge these terraces are cut out of the Pleistocene deposits, and may be of considerable longitudinal extent. About the gorge they may be cut from older rock or may be of composite structure; these terraces are ill general of much more restricted extent and of more complex arrangement than those above or below the gorge.

The valley where it issues on to the plains is broadly U-shaped in cross-section. To the south the slopes of Mount Hutt (6,810 ft.) show the typical smoothed surfaces with truncated spurs of a glaciated region. On the other side of the valley Round Top (2,917 ft.) is also smoothed and rounded, and Fighting Hill (2,393 ft.), lying to the north of the district, forms a typical roche moutonnée. Smaller roches moutonnées in this district are Bryant's Hill, to the north of the lower end of the gorge, and a low hill composed of rhyolite lying about a mile to the northwest of Bryant's Hill.

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Fig. 1.—Geological sketch-map of the Rakaia Gorge district.

The topmost terrace, which forms the main valley-floor, is throughout this district covered with irregular masses of morainic material. In

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places these moraines, which are composed of fluvio-glacial conglomerate with occasional large angular erratics of greywacke, are not more than a few feet in thickness; while elsewhere, as against the south-eastern slopes of Fighting Hill, they give rise to undulating country which indicates a thickness of 50 ft. or more. A prominent effect of these morainic accumulations is the formation of swamps and lakelets, such as Lake Constance, where erosion has failed to establish a complete drainage. The farthest extent of the moraines to the south-east is shown by an arc of deposits of somewhat greater thickness than those immediately behind it, which stretches from about the Glenroy Saddle, through Woolshed Hill, to the south-eastern end of the Mount Hutt Range.

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Fig. 2.—Topographical sketch-map of the Rakaia Gorge district.

A prominent physiographical feature of this district is what is known locally as “The Railroad,” which consists of two roughly parallel ridges running from Bryant's Hill in a north-westerly direction to the river-bed west of Fighting Hill. Since a full description and discussion of the origin of this feature has been given by Speight and Dobson (1923), further mention of it will be omitted from this paper.

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3. Stratigraphy.

The following table shows the succession of rocks observed in the Rakaia Gorge district, with their approximate age correlations :—

[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]

Sedimentary. Igneous. Correlations.
European. New Zealand.
7. River silts, gravels, loess, &c. Recent Recent.
6. Conglomerates, silts, boulder-clays Pleistocene Notopleistocene.
5. Teschenite and allied rocks Tertiary Notocene.
4. Coal-measures Upper Senonian Piripauan.
3. Andesites Rhyolites and pitchstones Cretaceous Pre-Notocene and post-Hokonuian.
1. Greywacke, shales, &c. Permian or Triassic Maitai or Hokonuian.

(a.) Permian or Triassic Sediments.

The oldest rocks exposed in the Rakaia Gorge district consist of intensely folded and faulted beds of greywacke, indurated grits, and dark-coloured finely-laminated slaty shales. They outcrop on both banks of the river at the upper end of the gorge. On the left bank they are overlain by the lower members of the Pleistocene deposits (Plate 18, fig. 1), and meet the rhyolite in an almost vertical junction—possibly due to faulting. On the right bank they are overlain by both rhyolites and Pleistocene deposits with marked unconformity. The rocks near this junction show intense weathering, which has resulted in the formation of masses of limonite irregularly distributed within a zone of about 10 ft. thickness. Fighting Hill and the Mount Hutt Range are also composed of these rocks.

The average strike of these beds throughout the eastern part of the alpine area of Canterbury appears to be in a north-easterly direction. In the small outcrops of the Rakaia Gorge inlier, however, folding has been intense, and small overthrust faults further obscure any general orientation of the strata. The dip is always at a steep angle, and the following represents a series of observations of the strike taken at intervals of about 20 ft. along the left bank of the river: E. 18° S.; E. 61° S.; E.; E. 10° S.; N. 20° E.

The only organic remains found in these rocks in the Rakaia Gorge are some indistinct annelid tubes similar to Torlessia McKayi Bather. They consist of straight or slightly curved and usually flattened tubes; jointing of the body-segments is visible with a pocket-lens, each segment being about 2 mm. long. The width of the tubes varies from 2 mm. to 4 mm. These occur in a grey coarsely-laminated shale in the outcrop on the left bank, as noted by Haast in a geological map of the Malvern Hills, dated 1871, and now in the Canterbury Museum, Christchurch.

With the criteria at present available it seems probable that these rocks should be referred to the restricted Maitai series—i.e., should be approximately correlated with the Maitai rocks of Nelson, which Trechmann (1917) has shown to be of Permo-Carboniferous or Permian age. Jaworski's (1915) opinion, however, should be noted—that Torlessiia McKayi should be referred to the genus Terebellina and indicate Triassic age.

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(b.) Cretaceous Igneous Rocks.

Following upon the orogenic movements of early Cretaceous times, and preceding the deposition of the coal-measures in this district, there was a period of volcanic activity which gave rise to flows of rhyolite, pitchstone, and andesite, and some andesitic fragmental deposits. The intrusion of a dyke of andesitic character into the older sedimentaries probably accompanied these eruptions.


At the upper end of the gorge rhyolites occur on both banks of the river, lying with marked unconformity upon the denuded surface of the above-described shales and greywacke. On the left bank they are overlain by andesite-flows and andesitic breccias; on the right bank by the lower beds of the coal-measures. The relation of the andesites on the left bank to the coal-measures and rhyolites on the right bank is somewhat obscure; this is probably due to an abrupt thinning-out of the former, such as is illustrated in fig. 3.

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Fig. 3.—Cross-section at upper end of Rakaia Gorge (line AB in fig. 1). 3, rhyolites; 4, andesites; 5, coal-measures; 7, Pleistocene deposits.

Separated from the junction of the rhyolite with the older sedimentaries on the left bank by a few feet of normal rhyolite, there is a mass of pitchstone extending 2 chains along the river-bank. On the same side of the river there is a smaller mass of pitchstone about 6 ft. thick lying between the rhyolite and the andesite.

On the right bank no outcrop of pitchstone was found, though fragments of that rock occur amongst the talus derived from the slope on which the rhyolite meets the Maitai rocks; this probably indicates an extension of the pitchstone mass which lies near to that junction on the opposite bank.

The andesites overlying the rhyolites on the left bank are overlain farther down-stream by the coal-measures.

At the lower end of the gorge rhyolites occur on both banks, and are penetrated by pitchstones near their junction with the andesites. Lying between the pitchstone and the andesite there is a narrow layer of much-weathered rhyolite. The base of the rhyolite is hidden by the Pleistocene gravel deposits, but on both sides of the river there are clear sections showing it to be overlain by andesite-flows with interbedded andesitic breccias. The andesites, in turn, on both sides of the river are overlain by coal-measures.

An intrusion which was doubtless associated with the eruption of the andesites occurs at the upper end of the gorge on the left bank of the river, in the form of a dyke from 1 ft. to 1 ft. 6 in. in width, which penetrates the Maitai rocks. It is clearly shown in a section exposed about 7 chains up-stream from the junction of the older rocks with the rhyolites, and extends up the cliff with very uniform thickness, though with somewhat

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irregular walls. The beds on either side do not show any displacement, and there is little sign of contact metamorphism.

Besides these outcrops exposed in the gorge itself, there are numerous smaller occurrences of the Cretaceous volcanic rocks in the immediate neighbourhood. In Camping Gully rhyolite has been exposed in several places below the Pleistocene deposits. Bryant's Hill, and a lower ice-worn hill lying about a mile to the north-west of it, are also composed of rhyolite. Another outcrop of this rock occurs on the terrace slope leading up to the main level of the valley-floor about 20 chains to the west of the upper end of the gorge.

In Round Top, of the neighbouring Rockwood Range, a mass of andesites and associated breccias form the upper and north-western part of the hill. These rocks apparently form a capping with a north-westerly dip of about 15°, and overlie the rhyolites of which the south-eastern slopes are composed. Here, in the gully marked P on the map (fig. 1), pitchstones also occur with the rhyolites.


Rhyolite.—A hard, compact rock, usually jointed into massive rectangular blocks. Through the action of weathering agents the rock generally presents a surface which is stained to a light-brown colour. When fresh it is either white or may vary in colour from green to grey or black. Megascopic crystals of quartz and garnet (almandine) are locally abundant.

Microscopic examination of specimens of this rock taken from various parts of the gorge showed that, while the texture of the groundmass varied considerably, both the mineral content and the structure of the rock was very uniform throughout, the chief mineralogical difference between the various specimens being the presence or absence of garnet. Moreover, apart from that shown in a single flow as described below, there seems to be no progressive change throughout the mass of the rock either in degree of crystallinity of the groundmass or in abundance of garnet. Almost holocrystalline flows are irregularly interstratified with flows whose groundmass may be almost completely glassy. Types rich in garnet also appear apparently irregularly arranged amongst types poor or lacking in these phenocrysts.

The minerals present as phenocrysts are quartz, orthoclase, plagioclase (andesine to oligoclase with occasional albite), garnet, and biotite. Minerals of the groundmass are quartz, feldspar, apatite, magnetite, and rarely zircon. The groundmass varies in texture in different slides, and in different parts of the same slide, from a dark-brown glass to a microfelsitic (Iddings, 1909) matrix. In a few slides indistinct micrographic intergrowths of quartz and feldspar are apparent in the more crystalline parts of the groundmass, together with patches showing microspherulitic structure. In general, however, spherulitic intergrowths are absent. Flow-structure is nearly always present to some extent, and may be very prominent. A feature of nearly all the rhyolites examined was the development of “flow-breccia” structure (Iddings, 1909, p. 331).

Pitchstone.—A brittle, easily-weathered rock, showing either (as in the outcrop at the upper end of the gorge) very perfect rectangular jointing, or (as at the lower end of the gorge) massive outcrops with little jointing. The rock is pitch-black in colour, with the characteristic vitreous lustre and conchoidal fracture of volcanic glasses. Phenocrysts of clear or slightly discoloured quartz are visible, and garnets are abundant in all of the outcrops in this district.

Under the microscope the pitchstones are markedly uniform in character and closely resemble the more glassy types of rhyolite; the occasional presence of small phenocrysts of hypersthene is the only difference in mineral content, and, except that no microspherulitic intergrowths were observed in the pitchstones, their micro-structure is similar. Perlitic structure, especially in the rock of the outcrops at the lower end of the gorge, is perhaps more marked than in the glassy rhyolites.

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Fig. 1.—Pleistocene deposits overlying Maitai beds at upper end of Rakaia Gorge (left bank).
Fig. 2.—Terraces at lower end of Rakaia Gorge (right bank).

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Andesite.—The rock of which the flows are composed is a hard dark-grey or black rock, which, however, is very susceptible to the action of weathering agents. Fresh specimens of the rock are difficult to obtain except where erosion has been so rapid that the products of decomposition have not been able to remain in place. In the field the rock generally appears as a reddish or greenish mass, presenting a somewhat incoherent and crumbling surface. The only crystals visible megascopically are of feldspar, and these occur as numerous, evenly-distributed, rectangular prisms. The presence of these phenocrysts is very characteristic of the rock, and they remain visible to the naked eye or under a pocket-lens even in most weathered specimens. A very general feature of this rock is the abundance of secondary silica present as quartz of different varieties—chalcedony, amethyst, agate, and opal. These minerals, together with chloritic decomposition-products derived from the ferromagnesian minerals of the rock, are found filling veins, druses, steam-vesicles, and other cavities throughout the whole mass. In places the rock becomes scoriaceous, and the vesicles are filled with bright-green amygdaloids composed chiefly of chlorite. The presence of these minerals has led to the popular supposition that copper-ores and gold may be found in these rocks, but prospecting has failed to detect any minerals of economic value in payable quantities. The amethysts, which have attracted some attention, are of a pale colour, usually occurring in druses as clusters of small hexagonal prisms terminated by pyramids. Specimens have been collected from the face opposite the Mount Hutt homestead up to 4 in. in length, but these are exceptionally large for this locality

Calcite is also common as a vein-mineral in the andesites. On the southern slopes of Round Top there are veins varying in size up to 2 ft. which have been partially filled with calcite, quartz being subsequently deposited as a coating on the calcite, and giving rise to negative pseudomorphs.

The breccias associated with the andesite-flows are typical volcanic breccias consisting of angular fragments of the andesite rock varying in diameter up to 2 in. or 3 in. with occasional larger blocks cemented together in a matrix of finer material. There is no apparent regularity in arrangement of the flows and fragmental deposits, but the former are in far greater abundance. The breccias occur merely as occasional strata, never more than a few feet in thickness, interbedded between the flows. The secondary minerals associated with the andesite-flows are also abundant in the breccias.

The microscopic character of the andesites is very uniform in all specimens collected from this district. The chief variations observed were in the proportion of ferromagnesian minerals present and in the texture of the groundmass. Plagioclase (acid-labradorite to andesine), hypersthene, and augite occur as phenocrysts in a groundmass which is typically composed of minute feldspar laths, some pyroxene in granular masses, magnetite, and a brown glass. The texture of this matrix varies in different slides: in some there is little glass and the feldspar laths attain a larger size, when they may be recognized as andesine or labradorite; in other slides the glassy material predominates, and the feldspars appear as scattered microlites. The amount of pyroxene present in the groundmass also varies, being in some slides apparently absent. In general the structure is hyalopilitic as defined by Rosenbusch, the “felted” character being prominent in the more crystalline

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varieties. Throughout the rock there is much secondary quartz and opal, filling small veins and cavities. Small round vesicles frequently show a border of radiating, fibrous, chalcedonic quartz, with the centre filled in with opal.

The rock forming a dyke in the greywacke at the upper end of the gorge is much weathered, and presents a brown sandstone-like appearance. Plagioclase (acid-labradorite), augite, ilmenite, apatite, and a large amount of secondary quartz and calcite are recognizable under the microscope. The texture is of a fine, even grain, the bulk of the rock being composed of feldspar laths with occasional patches of glassy residuum. It would seem more probable that the intrusion of this dyke was associated with the Cretaceous volcanic activity than with the Tertiary basic intrusions, solely on account of the andesitic character of the rock.

The following are the results of analyses made in the Dominion Laboratory of specimens of each of the effusive rocks of this group, with their classification according to the C.I.P.W. system :—

[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]

(1.) (2.) (3.)
Silica SiO2 75.68 69.54 58.57
Alumina Al2O3 12.45 13.18 17.03
Ferric oxide Fe2O3 0.44 1.04 4.85
Ferrous oxide FeO 0.40 0.92 1.33
Magnesia MgO 0.07 0.31 1.17
Lime CaO 0.66 1.06 5.45
Potash K2O 6.41 2.25 2.70
Soda Na2O 2.14 4.16 3.20
Water lost above 105° C. 0.66 4.98 1.23
Water lost below 105° C. 0.58 2.16 2.60
Carbon dioxide CO2 0.11 Trace 0.09
Titanium dioxide TiO2 0.19 0.21 1.38
Zirconium dioxide ZrO2 0.01 0.01 0.01
Phosphorus pentoxide P2O5 0.20 0.11 0.32
Sulphur S None None None.
Chromium trioxide Cr2O3 None None None.
Nickel oxide NiO Trace Trace 0.02
Manganous oxide MnO 0.01 0.02 0.11
Strontia SrO None None None.
Baryta BaO 0.06 0.10 0.05
Lithia Li2O Trace Trace Trace.
100.07 100.05 100.11

Rhyolite: the island, lower end of Rakaia Gorge.


Pitchstone: lower end of Rakaia Gorge, north side.


Andesite: lower end of Rakaia Gorge, south side.

[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]

In (1) and (2) the analyst reports that the carbon dioxide is not present as calcite, as it is liberated only on heating to about 100° C with dilute hydrochloric acid.

(1.) (2.) (3.)
Quartz 37.98 34.14 17.16
Orthoclase 37.81 13.34 16.12
Albite 17.82 35.11 27.25
Anorthite 1.95 4.45 23.91
Corundum 1.43 2.24
Diopside 0.46
Hypersthene 0.20 1.33 2.80
Magnetite 0.70 1.39
Ilmenite 0.46 0.46 2.74
Haematite 4.96
Apatite 0.34 0.34 0.67
Calcite 0.20 0.20

I 3(4) 1″ 2″ (Magdeburgose).


I 3(4) (1)2 4 (Alsbachose).


I (II) 4 3 (3)4 (Yellowstonose).

– 99 –

The chief points of interest regarding the composition of these rocks arise out of a comparison of the rhyolite and pitchstone. As is suggested below, the pitchstone may represent merely certain of the acidic flows of these eruptions, which, either owing to rapid cooling or other causes by which free molecular movement was hindered, have consolidated in a more vitreous form. If this is the case the rocks should show similar chemical compositions. The difference in relative proportion of the alkalis is, however, marked: the potassic nature of the rhyolite shows itself mineralogically in the abundance of orthoclase relative to the soda-bearing feldspars, while in the pitchstone the reverse occurs. The chemical similarity of the two rocks, however, seems sufficient to admit of their being successive flows of the same eruption, a conclusion to which the field evidence points.

Comparison of these analyses with those given by Speight (1922, p. 79) of Banks Peninsula rhyolites and pitchstones shows a general similarity.

Owing to the abundance of secondary silica contained in the andesite, the C.I.P.W. quantitative classification, which the authors definitely limit to fresh specimens, does not illustrate the character of the rock. Classified as “yellowstonose,” it is grouped with dacites and rocks corresponding in composition with quartz-diorites, whereas its microscopic character clearly shows that it belongs to a more basic group.

The Origin of the Pitchstones.—Haast (1871 and 1879) considered the pitchstone to be merely a facies of the rhyolite—the first flows to be erupted on to a cold land-surface. Cox (1884, p. 40), however, considered the pitchstone to be dykes belonging to the same system as the dolerite and basalt intrusions found penetrating the coal-measures and older rocks. The following evidence seems to confirm Haast's view that in this and neighbouring districts they are merely glassy facies of the rhyolite :—

1. No instances have been reported of the pitchstones associated with the volcanic rocks here considered penetrating any rocks other than the rhyolites.

2. In every outcrop observed the directions of the pitchstone masses conform to those of the apparent flows of the rhyolite. The presence of jointing frequently makes it difficult to determine the orientation of the flows of the rhyolite, but where determination is possible the pitchstone is found to be in parallel arrangement. This is especially true of the occurrence at the lower end of the Rakaia Gorge, where the direction the rhyolite-flows is clear.

3.There are certain undoubted rhyolite-flows which may be considered as intermediate in character between the normal pitchstone and rhyolite types. They are black in colour, show a somewhat vitreous lustre, and microscopically appear as very glassy rhyolites. Other flows which are chiefly composed of normal rhyolite at the margin grade into a pitchstone type as shown above.

While it thus seems more likely that the pitchstones do not form dykes, yet Haast's inference that they represent those flows resting directly upon the older rocks, and therefore cooled more rapidly than the succeeding flows, requires some modification. The presence of normal rhyolite between the pitchstone and the older (Maitai) rocks shows that, whatever may have been the cause of the glassy facies, it was not due to conditions attendant only upon the first flow to be put out on to the land-surface. Furthermore, the pitchstones occurring near, but here also not next to, the junction with the andesites, may only be due to this rapid cooling of the first flow if the andesites represent the older of the two volcanic series. As shown below,

– 100 –

however, the weight of evidence seems to point to the andesite being of younger origin than the rhyolite. Thus while it is possible that the first flows of the rhyolitic series in places may have assumed a glassy character, yet the pitchstone facies is by no means confined to that horizon.

In the Rakaia Gorge it must be considered as accidental that each of the pitchstone outcrops occurs close to, though, as careful examination shows, not in actual contact with, the junction of the rhyolites with other rocks.

Order of Eruption of the Rhyolites and Andesites.—Haast, in treating of this and the adjacent Malvern Hills and Mount Somers districts, always considered the andesitic eruptions to have preceded the rhyolitic. Cox, who was the last to discuss at all fully the mutual relations of these two series of rocks, agrees with Haast that the andesites were the first erupted rocks, but in view of inconsistencies arising from an adoption of this theory in the Malvern Hills and Rakaia Gorge he postulates a special type of eruption for the rhyolites. These rocks he considered to have been erupted in a very viscous condition, the accumulation taking place by a process of endogenous growth, as described by Judd (1881, p. 134). Of recent years, however, there has been some doubt cast on this hypothesis, and a close examination of the Rakaia Gorge district, supplemented by more rapid observations in the Malvern Hills and Mount Somers districts, would seem to show that there is a balance of evidence in favour of the rhyolite being the first erupted rock.

Haast does not state definite reasons for considering the andesites to be older, but Cox bases his endogenous-growth theory upon the following observations :—


Micro-structure of the rhyolite indicates a viscous lava.


On the flanks of Mount Somers the andesites dip inwards towards the main rhyolite mass.


“The fact … that the melaphyres are lying on the liparites [in the Rakaia Gorge] at the angle of dip which they assume is in itself a proof of greater age, when we consider that beds of tufa are interstratified with the solid floes.” (Cox, 1884, p. 39.)

Of these lines of evidence, (1) is of merely accessory value; (2) and (3) may both be criticized in the light of later advances in the science of New Zealand geology. At the time of Cox's report the conception of a Pliocene period of crustal movements had not been developed. The work of McKay, Cotton, and others, however, has since shown that the present topography of New Zealand is independent of that of pre-Notocene times. Thus the fact that the andesites flanking Mount Somers dip towards the centre of that mass must be considered as possibly due to the presence of faults. The third argument is also open to criticism. The coal-measures overlie the andesites in this place with a dip of not less than 35°; so that, taking Cox's angle of dip of the andesites (45°), and allowing that no movements have disturbed the igneous rocks relative to the coal-measures, the angle of slope at which the tuffs and breccias were laid down could not have been greater than 10°. This occurrence therefore seems insufficient as “a proof of greater age” of the andesites.

A further point which doubtless influenced both Haast and Cox in considering the relative age of these rocks is that they supposed that there were beds of rhyolitic tuffs interstratified with the coal-measures at Mount Somers and the Rakaia Gorge. As shown below, however, the coal-measures in the Rakaia Gorge are probably entirely of cataclastic origin.

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With regard to the possible objection that if the andesites overlie the rhyolites the solutions which deposited the silica, calcium carbonate, &c., in them would be expected to have penetrated the underlying rhyolites, two suggestions may be made. Such solutions were probably closely connected with the magma itself, and deposition of the secondary minerals would follow closely upon the consolidation of the rock, and be to a large extent confined to it. Secondly, the rhyolite, being of a compact nature, would not easily be penetrated by these solutions. The fact that small amethysts, with some veins of chalcedony, occur in the rhyolite on the right bank at the lower end of the gorge shows that these solutions did to some extent affect this rock.

The evidence in favour of the rhyolites having preceded the andesites is—


The andesites in each outcrop of the Rakaia Gorge occur between the outcrops of rhyolite and coal-measures, and in Round Top rest upon the rhyolite. The possibility of faulting having caused an inversion of the older rocks over the younger is unlikely.


The nature of the contact of the andesites with the rhyolites at the lower end of the gorge. The rhyolite which lies between the pitchstone and the andesite is much weathered and decomposed to an easily eroded white mass.

Where the rhyolite rests upon the older Maitai rocks there is little evidence of alteration of the erupted rock. This seems to indicate that the weathered rhyolite represents an old land surface upon which the andesites were erupted.

The supposition that either the andesites or the rhyolites of the Malvern Hills and Rakaia Gorge were not contemporaneous with those of the Mount Somers district would clear away the chief difficulties involved in this problem. The rhyolites of both districts, however, in megascopic and microscopic character are almost identical, and both have similarly associated pitchstones. The andesites of both districts are also of a closely similar nature, and since both series are definitely of Cretaceous age it must be considered very improbable that the eruptions of each type were not contemporaneous in both districts.

(c.) Cretaceous Sediments.


The coal-measures overlying the rhyolites and andesites in the Rakaia Gorge are the only members of the Notocene group of sediments exposed in this district. They outcrop as part of a basin-shaped fold which is tilted towards the south-west, so that one-half of the basin is obscured below the Pleistocene deposits on the south side of the river. The maximum thickness exposed is about 1,000 ft. At the upper end of the gorge on the right bank they rest on a denuded surface of the rhyolite, and strike W. 30° N. with a southerly dip of 30°. On the left bank the coal-measures occur in the central part of the gorge, resting on the andesites. At the upper end of the outcrop the strike is roughly north and south, but in Chasm Creek it has swung round to N. 15° E., and the beds dip at 30° to the west. Following the outcrop down-stream, the strike continues to swing round, following the edge of the basin, until, where again the beds may be seen resting on the andesites, it has assumed a N.N.E.-S.S.W. direction, and the beds dip at an angle of 35° to the W.N.W. On the right bank, below the Mount Hutt homestead, these beds occur again resting on the

– 102 –

andesites with a north-east strike and dipping at 35° to the north-west. Thus the southern wing of the exposed segment of the basin is completed; further evidence of the form of the beds to the south-west is buried beneath the later deposits of gravel and silts lying between the river and the foot of the Mount Hutt Range.

In both outcrops of coal-measures on the right bank the topmost bed exposed is overlain by the teschenite rock of a sill; elsewhere the Pleistocene deposits follow unconformably.


The coal-measures consist for the most part of coarse conglomerates with interstratified beds of shale and sandstone and seams of brown coal. Current-bedding is a prominent feature of the deposits wherever exposed. Associated with these sediments, and more especially with the coarser conglomerates exposed on the right bank near their junction with the teschenite sill, there is a considerable quantity of fossil wood which has been preserved by partial petrifaction. Large pieces of tree-trunks and boughs often several feet in length occur associated with limonite masses, and usually containing radiating tufts of fibrous haematite at the nodes.

In two places, marked S on the map (fig. 1), there are deposits of siliceous sinter together with silicified grits and sandstones. These deposits occur at about the same level on opposite banks of the river, and probably represent a period of thermal activity following upon the above-described volcanic activity. The partial silicification of the wood-remains and the formation of haematite are doubtless largely due to the action of waters associated with the deposition of this sinter.

There is little regularity in the arrangement of the beds of different texture throughout the sequence. Conglomerates are the predominant type, and occur throughout. In general the lower beds exposed are of finer material than those of the higher parts of this inlier. The seams of coal are also distributed irregularly, the thickest seams occurring in the upper half of the sequence.

1. The Coarse-grained Beds.—The conglomerates consist of closely packed pebbles of rhyolite varying in diameter up to 6 in. Together with the rhyolite debris there are occasional pebbles which may possibly be composed of much-altered greywacke, but close searching failed to show the presence of any andesite pebbles. In view of the close association of the andesites with the rhyolites in the district, this feature of the deposits requires some explanation. The possibility of the andesites being of later eruption than the deposition of the coal-measures is precluded by their undoubted mutual position. That the andesite pebbles, being more easily decomposed than the rhyolite, have been completely removed by weathering agents, either during deposition or subsequently, is also unlikely. An explanation is more probably to be found in a consideration of the locality from which the waste forming these beds was derived. The well-worn and rounded condition of the rhyolite pebbles suggests that they have been transported for a considerable distance. It is possible that the rhyolites may have extended to the east of the present outcrops over a wide tract of country, from which the pebbles of the coal-measures were derived, whereas the andesites were confined to within the area of deposition. This tract of rhyolite country would now be buried beneath the gravels of the Canterbury Plains, which (Speight, 1915) doubtless occupy the position of an infilled, down-faulted block. The facts that parts of the greywacke surface (now “fossil peneplain”) are found to the

– 103 –

west of this district with coal-measures lying directly upon them, and that rhyolite pebbles are found in the Notocene rocks of the Broken River and Big Ben outliers, supports the idea of an early Notocene land-surface to the east rather than to the west of the Rakaia Gorge district.

2. The Fine-grained Beds.—The finer sediments of the coal-measures consist of normal sandstones, grits, mudstones, and shales. Being doubtless derived from the same landmass that supplied the rhyolite pebbles of the conglomerates, these rocks are usually light in colour and of a siliceous character. In many cases there is much finely-divided carbonaceous matter scattered throughout.

There does not seem to be any evidence of volcanic tuffs in these coal-measures, as assumed by Haast and Cox. The section referred to by Cox (1884) as showing tuffs at the upper end of the gorge shows the normal succession with no brecciated material, volcanic bombs, or lapilli, and the sandstones and mudstones are interbedded with conglomerates of well-worn pebbles. The firmly-cemented character and light-grey or white colour of the finer-grained beds certainly give rise to a superficial resemblance to some volcanic tuffs, but the presence of finely-comminated plant-remains scattered irregularly throughout many of them disposes of this possibility. The colour, moreover, is readily understood, since these beds doubtless represent the detritus from a landmass which was largely composed of rhyolite.

3. The Coal.—The coal-seams vary in thickness from a few inches to about 10 ft. The thickness of any one seam may also vary considerably when traced laterally, the masses of coal being usually in the form of lenses of slight convexity. The coal itself when unaltered is a brown, hydrous coal, very similar in character to that being worked at Homebush, White Cliffs, Glenroy, Mount Somers, and elsewhere in Canterbury. On the right bank, near the junction of the coal-measures with the teschenite sill, it assumes a semi-anthracitic character, due to thermal metamorphism induced by the intrusion.

In the Rakaia Gorge the coal has been worked privately by the various owners of the land for their own consumption. On the left bank, in Chasm Creek, drives have been put in to a distance of about 100 ft. at two levels, to work a seam of 10 ft. thickness. Farther down-stream, on the same bank, a shaft was sunk near the junction of the coal-measures with the andesites, in order to pick up this seam again. Though doubtless it was passed through, it had here thinned out, and nothing of profitable thickness was found. These workings, which were carried out by Mr. George Gerard, of Snowdon, have been discontinued for the last twenty years, since all the easily-accessible coal has been removed. There is no record of the amount of coal extracted.

On the right bank of the river certain of the holders of the Mount Hutt Station have extracted small quantities of coal from the outcrops near the junction of the coal-measures with the sill described below, and near the junction with the andesite below the homestead. In these two places drives have been put into seams of from 4 ft. to 6 ft. in thickness for a distance of about 80 ft.

Age.—Since the fossil wood referred to above is the only recognizable organic material as yet found in the coal-measures of the Rakaia Gorge, the age of these beds can only be inferred by correlation with neighbouring districts. The lithological similarity and the identity of relations with the rhyolite leave no doubt, however, that the Rakaia Gorge coal-measures are the correlatives of those of the Malvern Hills, which (Trechmann, 1917; Woods, 1917; Wilckens, 1922) have been determined as Upper Senonian (Piripauan).

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(d.) Tertiary Igneous Rocks.


Overlying the topmost beds of the coal-measures exposed on the right bank in the central part of the gorge is a mass of igneous rock in the form of a sill. There are two outcrops of this rock: one, forming a cliff with a maximum height of 200 ft., extending in an east-and-west direction to the north of the Mount Hutt homestead; the other a bank varying up to 10 ft. high and about 5 chains long which lies about 20 chains south of the homestead. The junction of this rock with the underlying coal-measures is clearly visible; near it the normal sediments have been altered to a hard spilosite-like rock, and in an adjacent coal-seam the normal brown coal has been changed to a semi-anthracite analogous to similar altered coals in the Acheron River and Malvern Hills. From the nature of the contact and the lithological character of the whole mass it is evident that this body of rock represents a sill intruded into the coal-measures. The upper part of the intrusion and any overlying Notocene sediments have been either removed by erosion or buried beneath the deposits of Pleistocene silts and conglomerates.


The texture of this rock varies markedly at different levels. At its lower margin, where it is little weathered, it appears as a dark, hard rock of fine grain. Proceeding upwards it becomes coarser in texture, until at the highest point of its exposure it appears as a coarse-grained rock much weathered to a dark-green incoherent mass. In this part of the sill there are numerous small veins, varying from ½ in. to 2 in. in thickness, and arranged roughly parallel with the floor. The rock of which these veins are composed is of lighter colour and finer grain than the enclosing country.

The following are the chief types which occur as various facies of this intrusion :—


Coarse-grained or teschenite type. Coarse even-grained rock containing plagioclase (labradorite), titan-augite, olivine, analcite, prehnite, biotite, ilmenite, and apatite. Ophitic structure was not observed.


Non-porphyritic or dolerite type. Even-grained but of much finer texture than (1). Plagioclase (acid-labradorite), titan-augite, aegirine-augite, olivine, biotite, apatite, and ilmenite present. Ophitic structure slightly developed.


Porphyritic or basalt type. Very fine grain with small phenocrysts of olivine and a slightly greenish augite. The groundmass consists of a fine-grained though holocrystalline matrix of small labradorite laths together with some granular augite and a little olivine. Ilmenite is also abundant.


Vein rock. Of typically granulitic texture and containing orthoclase. nepheline, analcite, pyroxene (both titan-augite and aegirine), barkevicite (occasional granular crystals), biotite, and ilmenite. Slides made from the margin of these veins show an abrupt transition, though with no definite line of division, into the normal country rock by decrease in the nepheline and orthoclase with increase in the mafic elements.

Of these types, (1), (2), and (3) grade insensibly into each other, and depend upon distance from the margin of the intrusion. The whole group

– 105 –

is a typical suite of teschenitic rocks, and bears a close resemblance to many of the Scottish analcite rocks (Tyrrell, 1923; Walker, 1923).

The vein rock, which corresponds in mineral composition to a nepheline syenite, is of interest as showing a leucocratic differentiate of the magma from which the intrusion was derived. The process of differentiation was doubtless of the type in which the veins represent the product of a fluid residuum which has been squeezed along lines of weakness in the mass and has crystallized after the consolidation of the main bulk of the rock. The rock may thus be compared with the aplites of acidic rocks; the almost panidiomorphic structure of the vein-rock of this sill is noteworthy in making this analogy. Tyrrell (1923) shows that differentiation of a teschenitic magma in a syenitic direction will result in a theralite. The vein-rock of this sill does not, however, contain plagioclase, and thus represents a still further stage in the differentiation process—a nepheline syenite.

The age of the intrusion of this sill cannot be determined within close limits. It is post-Senonian and pre-Pleistocene. Probably it was intruded contemporaneously with the numerous other similar basic intrusions of Canterbury, and may have been associated with the Kaikoura orogenic activity, which reached a maximum in Pliocene times. It is, however, involved in the same folding and faulting that was then induced in the Notocene sedimentary rocks.

(e.) Pleistocene Sediments.


Overlying the Notocene and older rocks of the Rakaia Gorge inlier there are deposits of more or less unconsolidated sediments, including fluviatile conglomerates, lacustrine silts, glacial and fluvio-glacial deposits. These sediments surround the inlier, and form the material with which the Rakaia Valley is aggraded. They extend from the base of the Mount Hutt Range to Fighting Hill and the Rockwood Range, and form a continuous sheet between these two elevations into the High Peak basin. Upon the slopes of Round Top and the adjacent members of the Rockwood Range greywacke boulders representing glacial deposits of this age occur up to a height of 1,340 ft. above the level of the water at the lower end of the Rakaia Gorge. To the south-east of this district these Pleistocene deposits extend outwards to form the Canterbury Plains. In this district these deposits attain a great thickness above and below the gorge, where the river has eroded to a depth of 700 ft. below their surface without reaching the underlying rock.


The chief types of sediments represented in this series are:—

(1.) Conglomerate: coarse to medium grained, the pebbles having an average diameter of 3 in. or 4 in. Greywacke is the chief rock of which the pebbles are composed, but limestone, dolerite, gabbro, and, below the gorge, rhyolite and andesite, are sparsely represented. These gravel-conglomerates form by far the greater part of the Pleistocene deposits, and of them two distinct types may be recognized: one is stained to a light-brown colour through oxidation of the iron-bearing minerals under the action of weathering; the other is of the normal grey colour of the greywacke pebbles.

(2.) Silts: fine-grained, of a yellow or white colour. Usually show “varve” banding of very fine and slightly coarser material, each band

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being from ¼ in. to ½ in. thick. Occasional pebbles of greywacke are scattered throughout the deposits of this silt, suggesting transportation by floating ice during the time of deposition.

(3.) Fluvio-glacial conglomerate: composed of angular and striated boulders together with normal rounded pebbles of fluviatile gravels. Large angular boulders varying up to 4 ft. or 5 ft. in diameter occur in these beds. Greywacke, as in the normal fluviatile conglomerates described above, is the chief rock of which the boulders and pebbles are composed. The whole is cemented together with a very fine glacial silt.

(4.) Boulder-clay: a typical glacial boulder-clay consisting of angular boulders of variable size, some of which are smoothed and scratched, embedded in a very fine-grained silt. The boulders in this case also are almost exclusively composed of greywacke.

Correlation of the various sections of these deposits exposed in and about the gorge is rendered difficult by lack of continuity of the beds. The following represents the general succession observed:—


Loess, sands, and other recent superficial deposits.


Upper fluvio-glacial conglomerate.


Grey fluviatile conglomerate.


Brown fluviatile conglomerate.


Lacustrine silts.


Lower fluvio-glacial conglomerate.



The complete succession is not, however, shown in any single section. The following may be taken as representative exposures illustrating the stratigraphical relations of the several deposits:—

(1.) At the upper end of the gorge, resting upon the Maitai rocks on the left bank (Plate 18, fig. 1):—

[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]

7. Blown. sands, river-gravels, &c. 10 ft.
3. Fine-grained lake-silts 30 ft.
2. Silts with interstratified fluviatile conglomerates; conglomerates increasing with depth 30 ft.
Fine silts with occasional boulders 10 ft.
1. Boulder-clay 10 ft.
0. Greywacke.

(2.) At the lower end of the gorge, on the right bank, up-stream of the bridge. The sequence of beds, though somewhat obscured below the road by talus, appears to be as follows:—

[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]

7. Superficial deposits 2 ft.
5. Grey fluviatile conglomerate 30 ft.
3. Lacustrine silts 10 ft.
Conglomerate with limonite 3 ft.
Lacustrine silts 80 ft.
2. Fluvio-glacial conglomerate 50 ft.
1. Boulder-clay 10 ft.
0. Rhyolite.

The chief point of interest in this section is the presence of an over-thrust fault which has involved all the beds except the veneer of gravel forming the top of the terrace. In the road-cutting the effect of this fault is clearly shown where the silts have been thrust across the grey conglomerate; in the latter a bed of sandy material shows considerable distortion in the neighbourhood of the fault-plane. Below the road the line of fault becomes less clearly marked, and probably resolves itself into a series of parallel displacements; where, however, it has been thrust across the fluvio-glacial conglomerates the rhyolite shows a typical slickenside surface. The line of fault has an approximately N.N.E.-S.S.W. direction,

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and the beds to the east have been tilted so that they dip at an angle of about 15° to the south-east. The throw of this fault cannot be estimated with precision owing to the absence of any recognizable horizon on both sides of the line of movement. On the west side of the fault there is no sign of the easily recognizable bed of limonitic conglomerate which appears about half-way up the whole section, so that the throw cannot have been less than 40 ft. Estimating from an average level of the junction of the silts with the fluvio-glacial conglomerate, the throw would appear to be of about 70 ft. This junction, however, is not a definite line, since the silts pass gradually down into the conglomerate.

(3.) Below the gorge the Pleistocene deposits are represented by grey fluviatile conglomerates (5), and upper fluvio-glacial beds (6), overlying the brown conglomerates (4). Sections exposed on the right bank show à thickness of 150 ft. of brown conglomerate overlain by 350 ft. of grey conglomerate, which is overlain by fluvio-glacial beds varying in thickness up to 50 ft.


Tectonic Features.


Structurally the Rakaia Gorge inlier is an intermontane basin analogous to the Trelissick basin, Big Ben outlier, High Peak basin, or any of the numerous remnants of Notocene beds preserved amongst the oldermass blocks of the alpine regions of Canterbury. Owing to the presence of the Pleistocene deposits, however, the exact positions of the bounding fault-lines cannot be accurately determined. Some such major fault doubtless passes in a north-westerly direction between the gorge inlier and the Mount Hutt Range. In the northern part of the district there is also probably some continuation of the southern fault boundary of the High Peak basin, extending in a roughly south-west direction in front of Fighting Hill.

Two minor faults are apparent in the beds of the inlier:—

(1.) As described above, the rhyolite at the lower end of the gorge has been thrust across the Pleistocene deposits. This line of displacement probably continues across the river, so that the cliff forming the north-western boundary of the old ferry reserve may be a fault-line scarp. No definite evidence of displacement of the beds on the left bank of the river is, however, available. A similar example of Recent overthrusting is that described by Morgan (1908, p. 72) in North Westland; he attributes the movement to ice-pressure in Pleistocene times.

(2.) As shown on the map (fig. 1), faulting has occurred between the rhyolite at the upper end of the gorge and the adjacent rhyolite, coal-measures, and andesites. The downthrow side is to the south, and there is no evidence of its extension on either side of the gorge. The throw is indeterminable, but probably not great.


Small earthquakes, not recorded by the seismograph at the Christchurch Observatory, are frequently felt in this district. The shocks are of a sharp character, short in duration, and without appreciable preliminary tremors. Their intensity is about IV on the Rossi-Forel scale. These movements are probably due to settling-down of the loosely compacted sediments of the Canterbury Plains, which causes slipping at their junction with the older rocks rather than to disturbances of a deeper-seated nature.

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The Gorge.

The presence of Pleistocene silts and conglomerates above the gorge to a depth of at least 500 ft. below the level of the top of the barrier of resistant rocks now forming the gorge inlier requires that either (1) the bed of the river before the deposition of these beds was eroded to that depth, and therefore the barrier cannot have occupied its present position; or (2) this basin in the older rocks represents a trough hollowed out by the glacier which at one time occupied the valley, and whose deposits form the base of the Pleistocene series.

That glacial troughs analagous to the latter do occur has been amply shown by De Martonne (1911) and others. Such troughs, however, show gently sloping floors in an up-stream direction near the termination of the trough. In this case, allowing for later erosion by the river (which it should be noted would not greatly tend to steepen this face), the floor must have risen 300 ft. in 10 chains—i.e., possessed a grade of 1 in 2.

It would therefore seem that the later geological history of the district has been somewhat as follows:—

1. After emergence caused by the Kaikoura orogenic movements, erosion continued until a topography little less mature than that of the present was produced.

2. An extension, followed by retreat of the Rakaia valley glacier, took place, and the above-described boulder-clay was deposited.

3. Through upward movement of the block which now forms the gorge inlier, at a rate that was quicker than the river could keep pace with in cutting down its bed, a lake was formed in which were deposited the silts and conglomerates now overlying the lower boulder-clay. It should be noted that the part of the barrier now forming the south-eastern part of the inlier—i.e., the rhyolite of the island and surrounding parts—at one time must have been the highest part of the barrier, since the silts are found as far down-stream as Pipeclay Gully.

4. Aggradation continued until silts and gravels filled the valley to about the level of the present highest terrace.

5. About the time when this point of maximum aggradation was reached the glaciers again advanced, and the second glaciation of the district as described above took place. Upon the retreat of the glacier, erosion of the valley-floor set in, and the present gorge of the river was carved out with the terraces as described below.

A feature of morphological interest shown in the gorge is the influence of the structure of the beds upon the form of the course of the river. At the upper end it runs parallel with the strike of the coal-measures along the line of their junction with the older volcanics. Upon leaving the igneous rocks it makes a right-angle turn and follows the dip of the coal-measures until diverted by the less easily eroded igneous sill; from here it follows roughly the strike of the coal-measures as it swings round in the basin-shaped fold until a point is reached where it breaks across the volcanic rocks forming the lower wing of the fold.

The Glaciation.

Evidence that the valley of the Rakaia River has during Pleistocene times been occupied by an extensive valley-glacier is afforded by (1) the presence of glacial deposits, (2) the character of the existing topography. Some descriptive account of these features has been given above, and this

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section will be confined to reconstructing that part of the geological history of the period which will throw light upon their origin.

As shown when dealing with the Pleistocene deposits, there are two independent glacial deposits in this series which are separated by a considerable thickness of normal fluviatile or lacustrine sediments. It therefore follows that there must have been at least two periods in which ice, extended as far down the Rakaia Valley as the present gorge.

Evidence of fluctuations in the severity of the glaciation in New Zealand is extremely scanty, and it is generally considered that there has been but one great advance of the glaciers, followed by a gradual retreat to within their present limits. Speight (1921) shows that the notched spurs of the Upper Rakaia and Waimakariri regions give evidence of only one period of ice-advance. The extent of this intervening retreat may not have been great, but the thickness of intervening sediments, which may be, as above the gorge, more than 700 ft., shows that the duration of the interglacial phase was not short.

Of the extent of the first advance of the ice little can be said, since its deposits, which are visible only about the gorge (where they have doubtless been elevated to an exposed position by subsequent earth-movements) are the only traces of its existence. Results of the erosion of the ice of this period upon the topography have been completely obliterated by the action of the subsequent extension. The thickness of the lower glacial deposits, moreover, gives little indication of the intensity of that advance, since it is impossible to say whether they represent accumulations at a terminal face during a stationary period of the glacier, or whether they are merely the results of deposition along a steadily-shrinking ice-front.

Of the second extension more may be deduced. As shown by Haast (1879), the ice probably reached, when at its maximum, as far as Woolshed Hill, and spread out in piedmont form across the Canterbury Plains from the neighbourhood of the Glenroy Saddle to the south-eastern end of the Mount Hutt Range.

It is of interest, in considering the character of this glacier in its final extension, to notice how the loosely consolidated Pleistocene deposits which form the floor of the valley have resisted the erosion of this mass of ice. If the erratics found on the north-western slopes of Round Top are due to this extension, the ice in this place must have had a thickness of at least 500 ft. Even if these erratics were due to the earlier advance, a considerable thickness of ice must have been present to cause its extension as far as the Woolshed Hill morainic deposits. Despite this mass, however, the sediments of the valley-floor were not entirely scooped out. This occurrence seems to lend support to the theory that the action of glaciers on the floors of their valleys exerts a protective rather than erosive influence.

The Terraces.

The terraces occurring above and in the gorge are most easily explained as due to the slow reduction of the barrier of resistant rock. Their heights roughly alternate on either side of the river, as is characteristic of barrier terraces. Of especial importance in the preservation of the terrace-remnants about the gorge are the effects of bluffs and ridges of the more resistant rock, which have acted as turning-points for the stream in the course of its entrenchment. Illustrations of this preservation are shown in Plate 18, fig. 2.

The terraces above the gorge barriers may thus be explained without the assumption of any change in base-level due to change in position of the

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land relative to sea-level; the numerous terraces below the gorge, however, present some difficulties. The explanation advanced by Hutton (1873) and supported by Marshall (1912), that they are normal river-terraces due to emergence, cannot be accepted without question.

Speight (1907) shows that there is a lack of supporting evidence for a theory of recent elevation of the central Canterbury area. Such movement would also necessitate the deposition of the upper beds of the gravels of the plains under marine conditions. As yet, however, there has been no evidence adduced to show that any part of the plains has been formed of delta or marine deposits. Speight, concluding that these terraces are not due to uplift, suggests that the effect of a diminution in the supply of waste would be to cause an entrenchment of the rivers below the levels of their fans at their apices. Whether or not a stream will erode its bed in any place depends, however, upon the gradient of the stream in that place; other factors, such as load carried, merely determine the rate of erosion. In accounting for these terraces it is therefore necessary to explain how a change in gradient occurred by which the river was enabled to erode its own deposits.

In tracing the history of a fan formed by any stream the following two stages may be noted:—

(1.) In its earliest youth the stream, passing down a steep bed, with consequent high velocity and carrying a heavy load, builds up a fan upon reaching the edge of the immature country; as this process goes on the fan is built up until its apex is level with the point where it leaves its rocky bed.

(2.) As erosion continues the fan is extended to a less convex shape, with lower angle of declivity, at the same time the bed of the stream in the alpine area reduces its grade. The point where the stream leaves the rocky bed to pass over the fan will therefore also be lowered. The river must then begin to cut into the top of its own fan-deposits. As this process continues, residual terraces will form at the top of the fan, which will possess the feature of barrier terraces—that the remnant on either side of the river will not correspond in level but will form an alternating series. These terraces, moreover, will decrease in height above the river-level when traced towards the fringe of the fan.

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Fig. 4.—Section along left bank of gorge (line CD in fig. 1). 1, Martai beds; 2, rhyolites; 3, pitchstone; 4, andesites; 5, coal-measures; 6, teschenite; 7, Pleistocene deposits; F, fault.

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Fig. 5.—Section along right bank of gorge (line EG in fig. 1).

Applying this hypothesis to the terraces of the Rakaia River below the gorge, the observed facts appear to be explained—(1) The terraces are discordant in level on either side of the river: (2) they decrease in height above the river-level when traced towards the sea. In the Rakaia River the terraces have completely disappeared about twelve miles inland from the coast.

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5. Conclusion.

The following is an outline of the geological history of the Rakaia Gorge district:—


Permian or Trias: Deposition of greywacke, shales, &c.


Lower Cretaceous: Orogenic activity, with contortion of Maitai beds and emergence of land-surface.


Middle Cretaceous: Period of erosion when land-surface reduced to peneplain; eruption of rhyolites and pitchstones followed by andesites and andesitic breccias.


Upper Senonian: Marine transgression with deposition of coal-measures, and probably followed by more or less continuous deposition until Pliocene times.


Tertiary: Intrusion of teschenite sill into coal-measures.


Pliocene: Orogenic movements characterized by block-faulting and causing emergence of the land-surface.


Pleistocene: Period of erosion giving rise to present topography, and including two periods of extension of the Rakaia Valley glacier as far as this district.

Literature Cited.

Cotton, C. A. 1922. Geomorphology of New Zealand.

Cox, S. H. 1884. Selwyn and Ashburton Counties. Rep. Geol. Expl. N.Z.

Haast, J. 1871. The Malvern Hills District. Rep. Geol. Expl. N.Z.

Iddings, J. P. 1909. Igneous Rocks, vol. 1.

Jaworski, E. 1915. Die systematische und stratigraphische Stellung von Torlessia. Centralblatt fur Min. Geol. Pal.

Judd, J. W. 1881. Volcanoes.

Morgan, P. G. 1908. Mikonui Subdivision. N.Z. Geol. Surv. Bull. No. 6.

Speight, R. 1907. Some Aspects of the Terrace-development in the Valleys of the Canterbury Rivers. Trans. N.Z. Inst., vol. 40.

— 1921. Modification of Spur-ends by Glaciation. Trans. N.Z. Inst., vol. 53.

Speight, R. 1922. The Rhyolites of Banks Peninsula. Rec. Cant. Mus., vol. 2.

Speight, R., and Dobson, A. D. 1924. The so-called “Railroad” at Rakaia Gorge. Trans. N.Z. Inst., vol. 55.

Trechmann, C. T. 1917. Age of the Maitai Series of New Zealand. Geol. Mag., February.

Tyrrell, G. W. 1923. The Analcime Rocks of Scotland. Geol. Mag., June.

Walker, F. 1923. Scottish and Moravian Teschenites. Geol. Mag., June.

Wilckens, O. 1922. Upper Cretaceous Gastropods of New Zealand. Geol Surv. Pal. Bull. No. 9.

Woods, H. 1917. Cretaceous Fauna of the North-east of the South Island. Geol. Surv. Pal. Bull. No. 4.

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Considerations relative to the Age of the Earth's Crust.

[Read before the Otago Institute, 9th December, 1924; received by Editor, 19th December, 1924; issued separately, 6th March, 1926.]

In considering such questions as this we may roughly divide our knowledge of relevant facts into two classes—namely, (1) things which are demonstrated, and (2) things which are proved by the balance of evidence. They are not really two classes, as the inferior value of the second is only a question of the strength of the evidence.


Under the first head we have—

(1.) Life began on this planet: if it also began elsewhere, that does not concern us. There could be no actual connection between the life that began here and the life we may guess began elsewhere. Of such life we have no proof, and if we could import living beings from elsewhere it would not alter the problem.

(2.) Life began at some date: that is to say, there was a time in the earth's history when it was too hot to allow the beginning of life, as we understand the term, to be possible.

(3.) Changes on the hot surface of the planet resulting in a cool surface and condensed waters made the advent of life possible. No form of life that we can conceive of, and consequently no form of life as we understand the expression, can exist on the sun or on the glowing stars, and it is improbable that it exists on Mercury or Saturn. Ascertainable conditions suggest this improbability.

(4.) It began in a sense spontaneously—that is to say, under some natural impulse governed by antecedent conditions affecting the matter which became a living thing. We cannot scientifically postulate any other mode of beginning. This fact may belong to the first or to the second class. We accept it as a fact because reason teaches us to regard it as proved by substantially cogent evidence negativing any other hypothesis.

(5.) Save that we must shut off the early stages of the earth's existence when its surface was unfit to support life, we cannot do more than conjecture when it began. We can only collect such evidence as is available denoting periods when life existed. We must now recognize a tendency to push this further and ever further back in the history of the planet as our actual knowledge increases.


Of the facts made out by strong if not cogent evidence we have—

(6.) Life began by a concourse of matter under chemical, physical, and perhaps electrical conditions favourable to certain aggregations of matter attaining a quality which we call life. Personally I do not see how this can be effectively disputed. It cannot be denied save by substituting something more probable for this hypothesis. The further we go back in the history of speculations as to the origin of life—or, rather, of living beings—

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the more crude do we find these speculations. Milton describes the process from his point of view, veiled as to the origin but visible as to the result:—

The grassy clods now calv'd; now half appear'd
The tawny lion, pawing to get free
His hinder parts, then springs as broke from bonds,
And rampant shakes his brinded mane; the ounce,
The libbard, and the tiger, as the mole
Rising, the crumbled earth above them threw
In hillocks: the swift stag from underground
Bore up his branching head: scarce from his mould
Behemoth biggest born of earth upheav'd
His vastness: fleec'd the flocks and bleating rose,
As plants: …—Paradise Lost, Bk. 7. 463.

If seriously produced in our time, this description of the creation would provoke a smile; but I feel inclined to regard it with respect. It throws into poetic form the view of creation which for a century and a half after Milton wrote was the only known view.

(7.) It seems to me more reasonable to conclude that the aggregations of matter so conditioned as to be ready to receive life were very small rather than large. This conclusion I regard as of the utmost importance in connection with the question of geological time. We now know of the existence of living things so small that it is far under the mark to say, as was recently said of typhoid germs, that a thousand millions of living beings only aggregate to the size of a small pin's head. Why should we assume that they have degenerated to this size rather than that they were brought into being on this or on a much smaller scale? The presumption appears to me to be in favour of my proposition. Under this head we look to presumption until we can have demonstration.

If we venture without the support of any evidence to assume that life began in creatures or aggregations of greater size, how far are we to go? Are we to support the Miltonic creation? Where are we to make our assumed starting-point? I submit that in any case we cannot intelligently postulate anything larger than something still only perceptible with the aid of a microscope. Against a contrary assumption is the fact that certain diseases appear to be caused by poisons emitted by micro-organisms so small that our most vigilant observers have not yet seen them. Smallpox is one of these diseases. It begins and runs a course so like that of other fevers that, using our reasoning-powers, we conclude that it begins as other fevers begin, under the influence of a minute living organism capable of creating or acquiring and exuding a poison. Other familiar fevers, such as scarlet fever and measles, have a similar origin and run parallel courses. One of the most remarkable is trench fever, produced by organisms which are parasites in lice. This organism, like the others, has escaped visual observation. It has, however, been trapped and utilized to render patients immune from its own attacks. It is a reasonable conclusion from what is known of it that it is a highly specialized creature of ultra-microscopic dimensions.

These experimental observations were made during the late war, but more recent experiments have produced definite visual results. I need not go into the subject of ultra-microscopic observation and photography beyond mentioning that this latest triumph in scientific methods results in our obtaining visual cognizance of things that are too small to be seen under the most powerful microscope used in the ordinary manner. By this means it has been found that the microbe of foot-and-mouth disease, which has again appeared among cattle in Europe, belongs to the little-known group of “filter-passers,” organisms so minute as to be inseparable

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by a Pasteur filter from a liquid containing them. This discovery is the work of two Dutch scientists, Paul Frosch and H. Ahmen, who pronounce the causative organism of this disease to be a yeast-like organism propagated by budding. Grown on a solid medium it gave rise to colonies from 7 to 8 microns in diameter. A micron is a millionth of a metre. The investigators report: “With high magnifications and the use of light of the shortest possible wave-length it was found possible to get photographs of individuals of the organism which proved to be bacteria with an estimated length of about one-tenth of a micron.” (English Mechanics, 11th July, 1924.)

It is not essential for my purpose to go further into this subject, as it cannot be determined to what extent the existing inhabitants of the earth are descended from creatures as small as this; but these investigations render it probable that others of the disease-producing germs are of this minute character, and that until reasons to the contrary are shown it is reasonable to assume that life began in creatures of very minute size.

The organisms which produce this and other diseases which I have mentioned are not likely to be of the original forms into which life was infused; they are all highly specialized poison-producers. In the case of foot-and-mouth disease the organism cultivated to the twenty-sixth generation has been found to retain its special virulence.

It seems to me not unreasonable to conclude that before life thus specialized as we know it began, countless generations in more primitive form had preceded the specialized forms, and that specialization involving increased size may have begun in forms more primitive than any we know.

At one time the habit prevailed of referring to amoeba as a primitive form; it is, however, a relatively large specialized form.

(8.) Is it not then a reasonable interpretation of the evidence to suggest that forms of life exist of the minute dimensions at which such forms, and perhaps all life, began?—that is, assuming that life did arise from organic matter. We need not assume that those very forms do not still from time to time come into existence. All that we can say is that we are wholly ignorant on the subject.

This course of reasoning may without violence even carry us down to those objects of which we now have actual cognizance, which we call ultra-microscopic objects.

It is not necessary, nor is it reasonable, to adopt any positive view on the actual question of size, but I think I have shown reasons for regarding it as probable that some of the germs of disease and other germs are of ultra-microscopic dimensions or of dimensions of a similar order when compared with visible living beings. I am content, however, to start with the assumption that life began where we reasonably conclude that it exists, because the qualities of beings that produce certain results are so like those of beings that we can see as to convince us that they are living beings. If that involves too great a draft on the imagination, then I am content to go down no further than to the size of beings which we can actually see—perhaps far below the size of the typhoid bacillus.

(9.) On these grounds I draw the conclusion that life began on a minute scale in that partly explored world of which we gain more knowledge every day. I speak of a conclusion, but I admit that the subject is so far speculative that any conclusion is liable to be upset. It would, however, be most difficult to explain away the case of the germ of foot-and-mouth disease, or to assert on reasonable grounds that it is a solitary case. In this connection it must be borne in mind that I am not writing on the beginnings

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of life, but am attempting to discuss a different subject—namely, a line of inquiry as to the age of the earth's crust. Any one who found it impossible to believe that life originated as I have suggested, and held that the balance of evidence led to an entirely different conclusion, would not find that to discard this would affect the main theme of this paper. Darwin never found it necessary to discuss the subject of the origin of life. He dealt with the history of organized creatures. I am—I hope, with becoming modesty—discussing a parallel topic. It is therefore quite incidentally that I pause to point out that great chemists and biologists—I need only refer to such names as John Tyndall and Henry Chorlton Bastian—may be said to have advanced a long way apparently in the direction of showing how life began. It is, however, sufficient to say that their brilliant experimental efforts, while producing what must be regarded as organic from inorganic substances, and going far to break down the barrier between the inorganic and the organic, have not resulted in the production of living things. Speaking with the greatest respect of these and other great men of science, I find myself driven to say that while throwing down one barrier they have perforce left the other standing. The synthetic production of organic substances by Tyndall has been immensely advanced and widened in the hands of his disciples, but we still await the Frankenstein who can make them live and move and reproduce their kind. We do not know the origin of life; we can only speculate as to whether we ever shall know it. For the present, however, on mere arithmetical grounds based on the ordinary law of probability, and opposed by no more rational conclusion, I have provisionally formed this conclusion:—

(10.) When we attempt to speculate on the age of this planet as the home of living things we are forced to consider the line of reasoning with which I started. Any one may reject it, but he ought to do so either by showing some fallacy appearing ex facie or by producing evidence displacing it. That is the course of reasoning by which any scientific proposition other than a mere dogma must be destroyed or displaced. Incidentally, recent observations in the field of palaeontology may be referred to.

It has long been the habit of geologists to divide the stratified rocks of the earth into two categories. The oldest was called Azoic, when and because, so far as observation went, it was devoid of indications of past life. The second embraced all the rocks in which fossils and other evidence of life were found, from the earliest to those at present in course of formation.

Some geologists have attempted to give the relative thickness of the rocks in these two categories. More than one published scale shows the Azoic rocks as equal to one-half of the whole known series of stratified rocks.

The pre-Cambrian rocks of Scotland are 8,000 ft. to 10,000 ft. thick, resting on a further series of great thickness. A thick series of slates and phyllites lies below the oldest Palaeozoic rocks in central Europe, with coarse gneisses below. In America, about the great lakes, a vast succession of rocks of pre-Cambrian age has been classed as forming six successive layers of great thickness. These have received names, the lowest being the Laurentian. Some of them have, however, been found to contain traces of living organisms. Since 1910–11, the date of the eleventh edition of the Encyclopaedia Britannica, further explorations have yielded remarkable results. Some of these are summarized in the recently added Supplement of 1922. In this summary an attempt is made to apply a time-scale to the whole period covered by the growth of stratified rocks.

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The term “Azoic” is not used, but, allowing sixty millions of “time units” for the building-up of all the periods, one-half is allotted to the pre-Cambrin ages. Here below the Laurentian is a vast assemblage of rocks awaiting further examination and exposition. When this stupendous series of old-world rocks is contemplated it is not surprising that geologists stubbornly stood out against the suggestions that limited the age of the earth's crust to a few tens of millions of years. In discussing the question of age the biologist necessarily goes hand-in-hand with the geologist.

It is when attempts are made to measure time by years that the greatest difference arises between the authorities. Here the physicist claims a voice, and in his present mood entirely reverses the contentions made a few decades back, when he relied on mathematical calculations which appeared to give but a relatively short life, both in the past and in the future, to the heat of the sun. Thus “taking the Lead as all produced by Uranium at the rate [elsewhere] above given we get an age of 925 million years. Some minerals from other Archaean rocks in Norway give a rather longer age … the upshot is that radio-active methods of research indicate a moderate multiple of 1,000 million years as the duration of the earth's crust as suitable for the habitation of living beings, and that no other considerations the side of pure physics or astronomy afford any definite presumption against this estimate.” (Lord Rayleigh, 1921.)

This method, it is said, carries back the appearance of Eohippus, the oldest form of horse, some thirty million years. Obviously this stretches the Tertiary ages in an enormous degree, making them cover seven or eight times as long a period as was not long since regarded as necessary. I have often wondered why the geologists and palaeontologists spoke of the Tertiary ages as extending over four million years. Why not forty million? I had to be silent in the presence of such unanimity—for unanimity there was. I am afraid that I am growing less modest as my years advance. Charles Dolittle Walcott, one of the most successful geological explorers of modern times, appears (1893) to speak of three million years for the age of mammals, while Joseph Barrell (1917) calls for fifty-five or sixty-five million years. The date (1893) given for Walcott's deduction was long prior to his remarkable discoveries. The modern tendency undoubtedly is to recognize immensely longer periods for the building-up of the rocks than were admitted a couple of decades back. Opinion on this comparatively new subject is, however, necessarily in a state of flux; we may not unreasonably hope for something more like demonstration as data accumulate and methods improve. Recent investigations by Walcott have revealed evidence of primitive life in the pre-Cambrian (Proterozoic) rocks of North America. In Montana, at a depth of nearly 10,000 ft. below the earliest Palaeozoic rocks (Cambrian), he found evidence of ancient reef deposits of calcareous algae which ranged upwards through 2,000 ft. of strata. The general barrenness of pre-Cambrian rocks is no doubt attributable to their extensive metamorphosis, presumably under the influence of heat-producing upheaval and other movements. This would tend to destroy everything but the hard casings of calcareous plants and of animals similarly protected. We need not, however, despair of the discovery of what may be termed the portraits of some of the even earlier denizens of the earth. Corals and sponges represent advanced structures—for Walcott has added to our knowledge of such life the discovery even of a monad or bacterium allied to Micrococcus living in pre-Cambrian times. Bastian had in 1910 referred to the fact that B. Renaut had found traces

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of Micrococci and Bacilli “even as low as the upper Devonian strata.” This discovery has been immeasurably surpassed by those of Walcott. If any value can be attached to the presumptions I have ventured to put forward, even a Micrococcus may from its structure be regarded as a creature which has already advanced in point of size at least far beyond the most primitive forms of life.

These considerations encourage further investigation, which may carry the discovery of evidences of life not only down throughout the Laurentian age but into and perhaps through a period equally long which lies below it. It is impossible now to speak conclusively of an Azoic age so long as we are dealing with stratified rocks.

Incidentally it must be allowed that, apart from the suggestions of the physicists, we can have no actual data for fixing the age in point of time of the earliest rocks until we have means of determining the rate of the deposit of strata. This may, but need not necessarily, have been more rapid in earlier than in later times. There may have been, and probably was, a vastly greater rainfall, causing a more rapid destruction and redeposit of the surface soils. We may also have to reckon on tidal action on a far more active scale if the history of the rocks carries us back to a period when the moon was appreciably nearer to the earth than at present. Before this can be touched we must come to an understanding with the astronomer.

I may now briefly refer to certain forms of which we have definite knowledge, and concerning which we may fairly speculate as to their having in the course of ages grown up in successive generations from some of the smallest to some of the greatest living creatures.

The recent expedition to the Ross Dependency found it necessary, for economic purposes, to weigh their captures. The great blue whale was found to weigh 150 tons. If the assumption with which I have started has any foundation, this animal by a long process of evolution may have progressed from an ancestor a thousand million of which would build up a speck the size of a pin's head. This is not proved, but it is not disproved, and it lies within the limits of reasonable argument. There are, of course, difficulties. A dormouse weighing an ounce has presumably an analogous history. In its strcture it has an obvious affinity with its great congener. They are both mammals, and each has seven cervical vertebre in common with almost all mammals. In the course of its evolution it has been found to be to the advantage of the race to which it and its ancestors belonged to remain small, while it has been to the advantage of the whale branch to attain and retain an immense size. Such differences for protective and generally benefit-conferring purposes are familiar throughout animal life. Trees great and small afford a similar illustration. The difference between the kiwi, living in dense forests in New Zealand, and the moa, formerly frequenting open plains, is another familiar instance, the relation between these two birds being much closer than in the first-mentioned instance—so close, indeed, as to leave open no question as to their common origin.

As to the rate of growth in the course of evolution, of which we know so little, we are not confined to one group in our search for animals of immense size. Great developments are scattered through the ages immediately preceding our own. Our largest surviving land mammal is the elephant, and in no other order is there now a near approach to its size. Baluchitherium, the vertebrae and limb-bones of which were found

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in the Upper Oligocene of Baluchistan, was probably much larger. In Pleistocene times the ground-sloths of South America rival the elephant and rhinoceros in bulk. The Megatherium was 18 ft. long, and its skeleton was of a most ponderous character. But perhaps the most remarkable of the mammals of the pampas is the great armadillo, Glyptodon claviceps, with the enormous weight of armour in separate plates over its body, head, and immense tail. It was 9 ft. long. The ponderous carapace indicates the prolonged operation of some cause or impulse, not solely connected with its feeding-habits, which had secured the preferential persistence (in survival of the retention) and progressive expansion of this monstrous development. Changes in the climate of this region probably produced a scarcity of food which led to the extinction of this great and varied fauna. The geological record as disclosed to us gives us at best but a hint in the shape of evidence. The pictures of past life, consisting not merely of skeletons, but of the undigested contents of the stomachs of the animals of the past, and even the fossilized eggs of the Dinosaurus recently (1924) discovered in northern China, are continually illuminating our means of study; but what lies buried beneath whole continents of land surface is only here and there disclosed by almost accidental breaches of that surface. What lies beneath vast stretches of ocean like that which separates southern Asia from Africa must remain for ever undisclosed so far as the human race as we know it is concerned. Thus far I have drawn my examples from the Tertiary period, with its specialized mammalian fauna.

We may, however, leave this and leap backward over the immense and unmeasured interval of time which separates the age of mammals from the age when the great reptiles were dominant. This interval is still unbridged. The prodigious development of the saurian fauna is the wonder of the geology of that great era. The largest surviving saurians, the crocodiles, are simply trivial when compared with their mighty predecessors—ancestors in any direct sense we can hardly call them.

The Brontosaurus excelsus of the upper Jurassic of Wyoming must have been fully 70 ft. long and 25 ft. high. Though its bones were lighter in structure than those of the great land mammals, it was built to walk on land, carrying its enormous weight on sufficiently stout legs. In the same region and of the same period is the Diplodocus carnegii, 80 ft. in length. Its skeleton shows that it must have had an immense muscular development to enable it to carry with comfort its enormous neck, projecting far beyond its body.

These two examples show size and weight, competing with that of our modern whales, carried by animals walking on land, even though living generally in the more sustaining medium of shallow waters. It is enough to mention these two examples of great vertebrates of the far past as showing that in that remote age great and wonderfully perfect development was achieved. *

To grasp the significance of these illustrations of the age in which the great saurians dominated the waters, the earth, and the air, we have to

[Footnote] * Since this paper was written further information has come to hand respecting the extensive discoveries made in recent times in the Tanganyika area of South Africa Amongst other important results these finds make it clear that the Gigatosaurus, with a humerus more than 7 ft. long, must have far surpassed in size any saurian or other form of animal theretofore revealed to the geologist. At least one writer claims for it a size double that of Diplodocus carnegii.

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try our best to appreciate the immense lapse of time which has occurred since the Jurassic rocks were laid down to the enormous depth at which we find them.

I can only briefly—and, I must admit, in a most unsatisfactory way—refer to what I may term the arithmetic of the case. It is a subject which opens up a field for endless speculation and inquiry, but, so far as I know, it has not up to the present been very comprehensively attacked. How many generations are required under given conditions of temperature, environment, &c., to raise from an invisible microbe a Brontosaurus or a whale? The problem is a stupendous one, but it is one that cannot be passed over, and is one which will, I think, receive more attention in the future than it has received in the past. If a creature the size of a typhoid bacillus bred another which doubled its weight in the first generation, and again added its original weight in each succeeding generation, its progeny would by successful development reach the size of a pin's head in, say, for example, a thousand million generations. If this went on unswervingly, and there were ten generations in each year, the result would be achieved in something like a hundred million years. It is, of course, mere guesswork to talk of ten generations in a year; there might be a hundred. On the other hand, it is a bold step to suggest that life has run so smoothly that in each generation the descendant of any particular microbe added its original weight to its progeny. Probabilities are against this having occurred over the whole series of generations. If the ancestry of the whale could be traced to a creature of the earliest date of the age of mammals, or even to some creature approaching the mammalian type as small as those we are told are found as we approach the Eocene, and that ancestor were found to be no more than an ounce in weight, then it has in a given period, not stretching back into the great age of saurians, so increased its bulk that its descendants of to-day are more than five million times the weight of their ancestor. There is, moreover, here in the later stages no question of ten generations in a year. Some of the great mammals do not produce at the rate of one a year, after some years have been spent in reaching maturity. Such figures relate to a comparison in weight between two creatures of the relative sizes, say, of a mouse weighing an ounce and a whale weighing 150 tons. But without going into actual arithmetic it may be pointed out that the disparity in weight between a creature the size of a typhoid bacillus and one the size of a mouse is vastly greater; while between the size and weight of the invisible microbe of foot-and-mouth disease and the comparatively gross typhoid bacillus there may be a disparity approaching either of the above. It is difficult to grasp comparisons of sizes in dealing with such matters.

The growing importance of the subject of examining the minutest forms of life is, however, attested by the discoveries of Dr. F. d'Herelle, of the Pasteur Institute, which came under my notice while I was revising this paper. It is found that certain bacteria which propagate diseases—perhaps hitherto regarded as not curable—so small themselves that they are only revealed by means of a microscope of 3,000 magnifying-power, are preyed upon and destroyed by smaller parasitic organisms of the “filter-passer” type. So minute are these that they are described as “just as small in relation to ordinary bacteria as a flea is to a calf.” These have now been styled “bacteriophages.” They have already been utilized for the destruction of the bacteria of persistent dysentery, and their discovery, isolation, and culture give promise of a revolution in the treatment of various

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diseases. (English Mechanics, 19th September, 1924.) What occurs to me in reading of such organisms is this: Are we to assume that they have retrograded from predecessors of larger size, or is it not more probable that they are rather in the vicinity of the size at which most lines of organized beings have received life ? Geological history, and analogies derived from it, give us the only clue we have at present to the answer. I will leave it to others, curious of arithmetical details, to calculate the relative weights or sizes of the various creatures involved in this discussion from a “filter-passer” to a Brontosaurus or a whale.

As we go down the geological ladder we find, as a general but not universal rule, that the creatures are getting smaller and smaller. It is true that examples of the smallest are always with us, while, as in the case of the giant saurians, the largest are sometimes left by the way; but the rule remains good until we find ourselves groping with Walcott among the immeasurably ancient rocks of Montana in the hope of here and there finding a legible stain or similar sign that life was once there. If our giant vertebrates are not descended from minute ancestors of those ages, we are left without means of even guessing where they came from. If they are so descended, have we not then to face the time-problem on some such lines as I have indicated ? The evidence points to the mammals having originated as small rat-sized creatures in the Jurassic and possibly, but doubtfully, in the Triassic period. That is to say, some of the saurians of the earlier period present some mammalian affinities; but to give this fact a definite value in this connection it would be necessary to show direct ancestral descent. It is considerations such as these which have led me to wonder whether our great geologists are right in assigning a period of only a few million years to the whole Tertiary age. I again most respectfully urge that it is probable that too much importance has been attached to evidence which appears to enforce this limitation.

I have presented certain problems of simple arithmetic. The least that can be said of them is that they deserve serious consideration. In doing so I have ignored evolution per saltum as I have ignored the Miltonic creation. I am quite prepared to admit that there may be an occasional sudden advance in size. We have, however, no knowledge of such a phenomenon. I am also quite alive to the observations of geologists who in certain periods of the earth's history find evidence of rapid expansion of groups of animals, as if new genera and species were appearing at an accelerated rate: Deprét has spoken of an “explosion” of forms in the ammonite genus Neumayria. Moreover, we can no more insist on a universal law of uniformity in the matter of speed of variation than on any other supposed fact of which we have no actual knowledge. What we do know as a general rule to be deduced from the evidence is that evolution as a whole has been slow and gradual, especially perhaps when dealing with large bony structures. When, therefore, we find in a single group what we take to be evidence of a more rapid rate of change we note it as something exceptional, accentuating, without contradicting, the general rule.

My modest efforts to illustrate what we require in the shape of vast measures of time are merely tentative, but again I claim that they call for discussion. My own impression is that when the matters to which I have addressed myself receive from competent investigators the consideration they deserve the result will emerge in the shape of conclusions as to the age of the crust of the earth of a totally different order from those heretofore admitted. My own opinion as to how such inquiries and discussions will probably eventuate is sufficiently disclosed in the above.

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Cretaceous Plants from Kaipara, N.Z.

[Read before the Wellington Philosophical Society, 27th August, 1924; received by Editor. 28th August, 1924; issued separately, 6th March, 1926.]

The following notes are based on some specimens collected by Dr. P. Marshall, who kindly handed them to me for examination. They all come from Bull's Point and Batley, Kaipara Harbour, New Zealand, and the geology of the district has been fully dealt with by Dr. Marshall himself.

Though the plants do not give any indication of their exact horizon, they are associated with ammonites which point fairly definitely to an Upper Senonian (Campanian) age. * Most of the plants so far discovered are either fragmentary or belong to form-genera which are of little use for correlation purposes; they are, however, of considerable interest botanically, more especially as there are some petrifactions as well as impressions.

They occur mainly in hard nodules with an irregular fracture, from which it is difficult to obtain complete specimens; and, moreover, the plants were evidently in a fragmentary condition before fossilization. The impressions are partially petrified; thus, in the case of the fern Taeniopteris batleyensis, sections reveal to some extent the structure of the midrib, while collodion imprints of some of the leaf-fragments show the outlines of the epidermal cells. The araucarian leaves show the rows of stomatal pits very clearly, and can also sometimes be sectioned fairly successfully, while some of the araucarian wood is very well preserved.

One of the nodules is of great interest, for it consists largely of petrified vegetable debris in a sandstone matrix, and at once suggests comparison with plants containing nodules from other places. In its general structure this nodule closely resembles those from Upper Cretaceous beds of Japan described by Dr. M. C. Stopes (1909), from which a large and interesting flora was obtained (Stopes and Fujii, 1910). The matrix of the Kaipara nodule (kindly examined for me by Mr. W. Campbell Smith) consists of very angular quartz-grains in a calcareous cement together with a few crystals of feldspar, traces of mica, some green flakes of chlorite, and a very few grains which might be glauconitic. All these constituents are probably of detrital origin. Scattered through the matrix are abundant petrified plant-fragments and a few Foraminifera and other shells. Sections of other nodules showed a very similar mineral structure (sometimes more finely grained), but with very little plant-debris except for highly comminuted tissue-fragments. The Japanese nodules also vary in the amount of the plant contents, some containing only shells and no plants, while only a few have the fragments thickly massed. The Kaipara nodule differs from the Japanese in being slightly coarser-grained, with rather more quartz, while the plant-remains are not so well preserved. Dr. Stopes gives a detailed

[Footnote] * The collections of ammonites include the well-known form Pseudophyllites indra and the local representative of Gaudryceras kayei; also species of Puzosia, a form close to Acanthoceras rotomagense, and a form close to Phylloceras velledae. It appears, therefore, that if a single horizon is represented it must be somewhat lower than the Campanian. The question is discussed in a paper on “Upper Cretaceous Ammonites” in this volume.—P. Marshall.

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Fig. 1.—Taeniopteris batleyensis n. sp.
Fig. 2.—Taeniopteris batleyensis n. sp. The sori are very worn, and their structure is doubtful.
Fig. 3.—Fern-petiole from stem of Dadoxylon. Diagrammatic view of transverse section.
Fig. 4.—Sphenopteris sp.
Fig. 5.—Araucarites marshalli n. sp. Leaves attached to axis.
Figs. 6, 7.—Araucarites marshalli n. sp. Single leaf.

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Fig. 8.—Araucarites marshalli n. sp. Small portion enlarged to show rows of stomatal pits.
Fig. 9.—? Dammarites sp.
Fig. 10.—Carpolithus zeelandica n. sp.
Fig. 11.—Dadoxylon kaiparaense n. sp. Part of transverse section magnified, showing a fairly well marked annual ring, which, however, was not visible to the naked eye.
Fig. 12.—Dadoxylon kaiparaense n. sp. Radial section.
Fig. 13.—Dadoxylon kaiparaense n. sp. Tangential section.

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comparison of her nodules with the coal-balls and roof-nodules of the English Carboniferous beds. The resemblance is greater to the latter—which, however, are much finer grained, and contain very little quartz, while the plant-fragments are never closely packed. The differences are due to differences in deposition: the plants in the roof-nodules had drifted and been sorted out, but “the numerous minute fragments of the Japanese nodules … could neither have drifted far nor long before they were covered and preserved in that potent preservative and petrifying solution, sea-water” (Stopes, 1909, p. 203). The granular Kaipara nodules with their marine animals were also obviously formed fairly near the shore where detrital matter was accumulating, but the plants had perhaps drifted a little farther, for they had decomposed more than the Japanese plants, so that only pieces of wood and the thicker and more resistant leaves have been at all well preserved. There are at Kaipara a few fairly large stem-fragments (see below), but in the nodule which has been particularly noticed here there are only small scraps of coniferous and dicotyledonous wood, fragments of bark and leaves, and some rather poorly preserved seeds, none of which can be identified definitely. It is possible, of course, that other nodules may be obtainable which, though of unpromising appearance, may contain recognizable petrified plant-remains.

The English roof-nodules are found in the beds immediately above coal-seams, while the Japanese nodules occur at least 100 ft. below the seams in their neighbourhood; but the association with coal-seams is probably-more or less accidental, and the floras of the coal and of the nodules are not necessarily identical. They might, indeed, as Dr. Stopes believes is the case with the roof-nodules, belong to different plant associations, a fact which must be borne in mind when questions of correlation arise.

The recognizable plants from Kaipara are few in number. As in the Japanese nodules, and as in Upper Cretaceous floras generally, there is a mixture of ferns, gymnosperms and dicotyledons, but the material is insufficient for comparison with any other fossil flora. In the case of New Zealand no Upper Cretaceous flora has yet been adequately described, for the early work of Hector and Ettingshausen consists largely of nomina nuda, and Arber's monograph extended only to the Lower Cretaceous.

The following plants are recorded here: Ferns—Taeniopteris batleyensis n. sp., Sphenopteris sp. Gymnosperms—Araucarites marshalli n. sp., Dadoxylon kaiparaense n. sp., Carpolithus zeelandica n. sp., ? Dammarites sp. Dicotyledons—Phyllites sp., dicotyledonous wood.

Descriptions of Specimens.

Taeniopteris batleyensis n. sp. (Figs. 1, 2.)

Frond with the habit and venation of Taeniopteris, with secondary veins at right angles to midrib, but having circular sori with few sporangia between lateral veins, forming a row on each side of and close to midrib.

Dimensions: Maximum breadth, 1 cm.; length (incomplete), 5 cm.; diameter of sori, 1 mm. or slightly less.

Locality: Batley. One specimen only, with portion of counterpart.

The reference of this fertile frond to the form-genus Taeniopteris is purely provisional. The sporangial characters are not very well shown, and though the fern does not agree closely with any known genus of ferns Recent or fossil, it seems inadvisable at present to create a new genus for

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its reception, and additional material may throw further light on its affinities.

The midrib is stout and is partly petrified; the vascular bundle is apparently U-shaped in section, but the details are not clear. The secondary veins are at right angles, or nearly so, to the midrib, usually forking almost immediately, but sometimes running half-way to the margin before forking. In one case one of the branches can be seen to fork again. The sori are always close to the midrib and between the secondary veins. They are round papillae with a depression in the centre, and sometimes seem to contain 5 to 10 sporangia. The structure of the sori and sporangia cannot, however, be made out clearly, and it is impossible to see any trace of an annulus or of an indusium. In general appearance the sori resemble those of Laccopteris, which, however, are not situated definitely between the secondary veins, while the venation is distinctly reticulate. The genus Nathorstia is almost identical with Laccopteris in its vegetative characters, but the sporangia form a synangium, and Nathorstia can be recognized only in extremely-well-preserved material. In any case, the venation characters preclude a reference of our fossil to either of these genera, and it may further be noted that neither of them is known later than the Cenomanian.

Among Recent ferns the same type of venation occurs in Oleandra, but here the sori, which have a reniform indusium and numerous small sporangia, are situated immediately on the lateral veins, and usually at a short distance from the midrib. The genus is not known fossil, though sterile taeniopterid fronds have occasionally been referred to it on entirely inadequate grounds. Dawson described an inconclusive sterile fragment from Upper Cretaceous beds of Canada as Pteris (Oleandra) glossopteroides, but fossil fronds of the Taeniopteris type are mostly found in Jurassic and Lower Cretaceous beds. Some of these are now known to be cycadean, though the provisional generic name doubtless also includes some ferns. Several members of the Marattiales with a similar venation have been found fossil in a fertile condition, but the Kaipara fern does not agree with any of them in soral characters, nor with any of the living ferns of the taeniopterid type with which I am acquainted.

Fern-petiole (fig. 3).—In the hollow pith of the araucarian wood described below a fairly-well-preserved fern-petiole is seen in transverse section. The C-or U-shaped vascular bundle, with the incurved xylem strand, is so characteristic of ferns in general that without further evidence reference even to a family is difficult. Its occurrence is of interest as showing the possibilities of the material, and it seems quite probable that it belongs to the same species as Taeniopteris batleyensis.

Sphenopteris sp. (Fig. 4.)

Locality: Bull's Point.

The only specimen is a small fragment of a fern-frond about 3 cm. long. The pinnules are narrow, cuneate-lanceolate, alternate, decurrent and uninerved, with an entire margin. It is difficult to place such a sterile fragment in its exact systematic position. Similar fronds occur frequently in Mesozoic rocks, and have often been referred to the Cyatheaceae, and especially to the genus Thyrsopteris, on insufficient evidence. Others of a similar type have been named Asplenium or Asplenites, and the present specimen also resembles some species of Onychiopsis, which is a widespread Lower Cretaceous genus. It does not seem possible to identify the fragment with any degree of accuracy, and it is therefore referred to simply as Sphenopteris sp.

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Araucarites marshalli n. sp. (Figs. 5–8.)

Leaves thickly set on axis (usually occurring isolated), coriaceous, ovate or lanceolate, subacute, sessile with unguiculate base, finely striate longitudinally with numerous rows of stomatal pits. Internal structure: Numerous bundles with well-developed centrifugal wood radially arranged, centripetal wood (transfusion tracheides) present but not clearly preserved, resin-canals frequently but not always associated with bundles, numerous strands of hypodermal fibres, thick-walled idioblasts scattered through mesophyll.

The commonest fossils in the collection are leaves which I believe to belong very probably to the genus Araucaria, though, as there is always some uncertainty about detached organs, the usual termination ites is employed. They are referred to a new species, which I have much pleasure in naming after Dr. P. Marshall. Among Recent species there is considerable resemblance to the section Colymbea, and Araucaria brasiliensis in particular shows the rows of stomatal pits clearly, though they are not so distinct and well-marked a feature as in the fossil. This is a character which is not present in living specimens of Agathis as far as I have observed in herbarium specimens. Stomata were apparently present on both surfaces, but in the sections examined their structure was not preserved. Though the detached leaves from Kaipara exhibit considerable variation in size and shape, it seems probable that they all belong to the same species, in view of the similar variation among living araucarians.

Among comparable fossil species the nearest is Araucarites macrophylla (Bozzi, 1892, p. 375, pl. 16, figs. 1, 2) from the Lower Senonian of Italy, which externally is very similar indeed, though Bozzi's species showed no internal structure, and he does not mention the rows of stomata. Another similar type is Araucarites ovatus Hollick (1898, p. 128, pl. xii, figs. 3a, 4) from the Magothy Formation (Cenomanian), in which, however, the leaves are acuminate, while in Araucarites hatcheri Wieland (1910, p. 80, pl. 1, fig. 2) they are narrower and even more acuminate. This species seems to show traces of the stomata, for the figures indicate dots on the surface, and the stomata are more clearly seen in rows in some specimens of Araucarites bladenensis (Berry, 1908, pl. 14, fig. 3).

Kraeusel (1922, p. 7) unites A. bladensis and A. toucasi Sap. with A. crassifoha corda, and also includes the specimen from Lesina figured by von Kerner as Pachyphyllum rigidum. He records the species from the Lower Senonian of Swalmen, Holland, and says that the American specimens differ solely in being slightly larger. All these forms, however, are much smaller and more acuminate than Araucarites marshalli.

The fossil araucarians of New Zealand have scarcely been adequately studied. The foliage described by Ettingshasuen (1887, p. 154) from Shag Point and Malvern Hills as Araucaria haastii seems to be similar to the present species, though the leaves are rather more lanceolate and acute and have a definite median rib. Ettingshausen also states that isolated leaves are not found. The specimens figured by Hector (1886) are without descriptions and are too poorly drawn for exact comparison, but they do not seem to be identical with our fossil. An examination of the originals, if available, might show an agreement between Araucarites carinaria (Hector, 1886, fig. 24a, No. 13: the name is misspelt) and Araucaria haastii, &c. Tertiary araucarians evidently occur in New Zealand, as in Australia, but it is usually very difficult to say whether they should be referred to Agathis or to Araucaria. Among similar Australian species we may note Araucarites imbricatiformis (Johnston) from the leaf-beds of Macquarie Harbour, Tasmania (Johnston, 1888, pl. 36, fig. 1, p. 294).

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Some account of the internal structure has been included in the diagnosis, though the anatomical characters are not in general specific. Attempts to distinguish living species of Araucaria by their anatomy have not been very successful, and have been criticized by Seward and Ford (1906). On the whole, the anatomy of the present specimen resembles that of Araucaria rather more than that of Agathis, but the extent of the transfusion-tissue is not clear. In both Agathis and Araucaria sec. Colymbea the resin-canals alternate with the veins, whereas in the fossil they are usually below them, but also occur above or in the mesophyll more or less between the veins. Seward and Ford state that the canals are below each vein in Araucaria rulei, which is the flattest-leaved of the Eutacta group. The preservation of the fossil is scarcely good enough, however, for detailed comparison with living species.

? Dammarites sp. (Fig. 9.) Batley.

A portion of a leaf 2 cm. long and 1.5 cm. wide, with parallel venation, is obviously distinct from Araucarites marshalli. The veins, about 1 mm. apart, are more prominent, and occasionally bifurcate; there are no signs of stomatal pits, and the texture seems to be thinner. The leaf resembles some living species of Agathis, such as Agathis vitiensis, and some leaves of Podocarpus sec. Nageia are also rather similar. It is recorded here as evidence of another genus of plants, probably gymnospermous, in the Cretaceous rocks of Kaipara.

Dadoxylon kaiparaense n. sp. (Figs. 11–13.)

Annual rings of araucarian type, fairly well marked in places; bordered pits on radial walls in one or two (rarely three) rows in contact and sometimes slightly compressed; no pits on tangential walls; resin parenchyma absent; tracheids bordering rays sometimes containing resin, which is occasionally seen as “spools” in longitudinal section; medullary rays 1–8 cells high (usually 2 or 3), 1–10 pits in the field, and no abietinean pitting.

Locality: Bull's Point.

The above description is based on a stem 4 cm. in diameter, the centre of which is filled with the sandstone matrix containing the fern-petiole described above. A second (smaller) specimen, not so well preserved, doubtless belongs to the same species, the only observed difference being in the slightly greater height of the rays (up to 14 cells), which are sometimes biseriate.

I follow Professor Seward in referring fossil araucarian wood to Dadoxylon, though the present specimens, like many others of Upper Cretaceous and Tertiary age, closely resemble the wood of Recent members of the family. In fact, it seems to me quite probable that this is the wood of a species of Araucaria itself, and that it bore the foliage described above as Araucarites marshalli. There is, however, no evidence of connection, and it is unfortunate that the fragments of axis with leaves attached were not sufficiently well preserved to show the pitting. It must further be remembered that the wood of Araucaria and Agathis is extremely similar, and, indeed, indistinguishable in the fossil state.

The best-preserved fossil araucarian wood from New Zealand is Dadoxylon (Araucarioxylon) novae-zeelandiae (Stopes), from the Mid-Cretaceous (?) of Amuri Bluff. It differs from the present species in having better-marked annual rings, slightly larger tracheids, and a much greater development of resin in tracheids adjoining the rays (Stopes, 1914). The slight differences in the number of pits in the field and the height of the rays are probably unimportant. Dadoxylon ettingshauseni (Stopes) is poorly

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preserved, but has larger tracheids and separated bordered pits. The wood from Amuri Bluff referred by Ettingshausen (1887, p. 156) to Dammara oweni is inadequately described and figured, but the drawing of the radial section (pl. 6, fig. 14) suggests that abietinean pitting of the ray-cells was present, and it seems quite probable that this wood may belong to Protocedroxylon hectori (Stopes). There is, therefore no fossil araucarian wood from New Zealand closely resembling that from Kaipara, and it is consequently described as a new species.

Carpolithus zeelandica n. sp. (Fig. 10.)

Seed oval, winged, 8 mm. × 5 mm.; nucule 5 mm. × 3.5 mm. Wing or margin slightly contracted at base, broadly obtuse at apex. Nucule with median rib.

Locality: Bull's Point. One specimen, with counterpart; also one isolated nucule, probably of the same species.

Among the many figures of Cretaceous species of Carpolithus which I have examined there is none closely resembling the present species, though winged seeds of a somewhat similar type have been recorded from earlier Mesozoic rocks in various parts of the world. It is difficult to be certain of the true nature of the apparent wing, which might be the impression of a sarcotesta, and it is impossible to express any opinion as to the affinites of this seed except that it is probably gymnospermous.

Dicotyledonous Wood.—In addition to the fragments in the nodule mentioned above, a small stem from Bull's Point, about 1 cm. in diameter, was sectioned, but the preservation was too poor for description or identification.

Phyllites sp.

There are several small fragments of dicotyledonous leaves; one of these occurs on the same block as the Sphenopteris. A collodion imprint was obtained which showed only the upper surface of a leaf without stomata. A larger fragment, from Bull's Point, shows venation details very clearly, but in the absence of base, apex, and margin it is impossible to attempt identification. Collodion imprints again showed no stomatal details.


Berry, E. W., 1908. Some Araucarian Remains from the Atlantic Coastal Plain. Bull. Torrey Bot. Club, vol. 35, pp. 249–60, pl. 11–16.

Bozzi, L., 1892. La Flora Cretacea di Vernasso nel Friuli. Boll. Soc. Geol. Ital., vol. 10, pp. 371–82, pl. 15–16.

Ettingshausen, C., 1887. Beitraege zur Kenntniss der fossilen Flora Neuseelands. Denk. k. Ak. Wiss. Vienna, vol. 53, pp. 143–92, pl. 1–9.

Hector, J., 1886. Detailed Catalogue and Guide to the Geological Exhibits, New Zealand Court, Indian and Colonial Exhibition.

Hollick, A., 1898. The Cretaceous Clay-Marl Exposure at Cliffwood, N.J. Trans. N.Y. Ac. Sci., vol. 16, 1898, pp. 124–36, pl. 11– 4.

Johnston, R. M., 1888. Geology of Tasmania.

Kraeusel, R., 1922. Beitraege zur Kenntniss der Kreideflora: 1, Ueber einige Kreidepflanzen von Swalmen (Niederlande). Med. van's Rijks Geol. Dienst., ser. A, No.2; 40 pp., 5 pl.

Seward, A. C, and Ford, S. O., 1906. The Araucariae, Recent and Extinct. Phil. Trans. Roy. Soc., vol. 198, B, pp. 305–411, pl. 23, 24.

Stopes, M. C., 1909. Plant containing Nodules from Japan. Quart. Journ. Geol. Soc., vol. 65, pp. 195–205.

—, M. C., 1914. A New Araucarioxylon from New Zealand. Ann. Bot.. vol. 38, pp. 341–45.

Stopes, M. C., and Fujii, K., 1910. Studies on the Structure and Affinities of Cretaceous Plants. Phil. Trans. Roy. Soc., vol. 201, B, pp. 1–90, pl. 1–9.

Wieland, G. R., 1910. Two New Araucarians from the western Cretaceous. Bull. S. Dakota Geol. Surv. for 1908, pp. 77–81.

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The Upper Cretaceous Ammonites of New Zealand.

[Read before the Wellington Philosophical Society, 7th October, 1924; received by Editor, 13th October, 1924; issued separately, 13th March 1926.]

Plates 1947.


[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]

Discovery 129
Geological Occurrence 130
Descriptions of Species—
Phylloceras nera Forbes 134
Phylloceras radiatum n. sp. 135
Phylloceras forbesianum d'Orb. 136
Phylloceras minimum n. sp. 137
Phylloceras bistriatum n. sp. 138
Vertebrites murdochi n. sp. 139
Gaudryceras propemite n. sp. 142
Gaudryceras particostatum n. sp. 143
Gaudryceras subsacya n. sp. 144
Gaudryceras politissimum Koss. 145
Gaudryceras crenatum n. sp. 146
Zelandites kaiparaensis n. sp. 147
Tetragonites epigonus Koss. 149
Tetragonites latus n. sp. 149
Tetragonites simplex n. sp. 150
Tetragonites margaritatus n. sp. 151
Pseudophyllites indra Forbes 152
Pseudophyllites whangaroaensis n. sp. 153
Baculites rectus n. sp. 154
Diplomoceras wakanene n. sp. 155
Oxybeloceras sp. 156
Ptychoceras zelandicum n. sp. 157
Acanthoceras ultimum n. sp. 158
Gunnarites inflatus K. & R. 160
Gunnarites zelandicus Marshall 161
Gunnarites nordenskjoldi K. & R. 162
Gunnarites antarcticus Stuart Weller 162
Madrasites sulcatus n. sp. 164
Madrasites multicostatus n. sp. 164
Madrasites regularis n. sp. 165
Madrasites fortior n. sp. 166
Jacobites anderssoni K. & R. 168
Jacobites angularis n. sp. 169
Jacobites minimus n. sp. 169
Jacobites whangaroaensis n. sp. 170
Jacobites waitapuensis n. sp. 170
Neomadrasites nodulosus n. sp. 171
Brahmaites rotundus n. sp. 173
Maorites tenuicostatus Marshall 177
Maorites densicostatus n. sp. 178
Maorites suturalis n. sp. 179
Puzosia angusta n. sp. 182
Parapuzosia brevicostata n. sp. 183
Parapuzosia ordinaria n. sp. 184
Tainuia aucklandica n. sp. 186
Parapachydiscus rogeri n. sp. 188
Nowakites denticulatus n. sp. 189
Hauericeras ngapuhi n. sp. 190
Schluteria rarawa n. sp. 192
Importance of Suture-line 192
Geographical Relationship 194
Geological Age of Circum-Pacific Cretaceous Ammonite Horizons 198
Stratigraphical Features of the Batley Series 201
Geological Age of the Batley Series 203
Comparison with other Cretaceous Circum-Pacific Districts—
India 204
Japan 205
California 206
British Columbia 206
Patagonia 206
Chile 207
Seymour Island 207
Fossils found with the Ammonites 207
Literature 208

In the year 1917 (49, p. 443) a brief statement was made of the occurrence of ammonites at Batley, on the Otamatea arm of the Kaipara Harbour, North Auckland Peninsula. Since that time several visits have been made to Batley, and to Bull's Point, a neighbouring locality on the Arapaoa arm of the harbour where similar beds occur and where many of the same species of fossils are found. The first party, in 1916, included Mr. J. A. Bartrum, Lecturer on Geology at the Auckland University College, and Mr. R. Browne. In 1919 and in 1922 further visits were made to Bull's Point, and in 1923 collections were again made at Bull's Point and at Batley. In addition, a few specimens were found by Professor Benson and Mr. Bartrum in 1920, and by Mr. Ferrar, of the Geological Survey, in 1923.

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The result of all these collections has been the revealing of a fairly extensive fauna. It consists of specimens of several species belonging to all the important groups of mollusca, a few brachiopods, and a variety of fish-scales. There are also the remains of five species of plants. Probably the most interesting portion of the collection is that consisting of ammonites. Up to the present time very few species of ammonites have been recorded from the Cretaceous of New Zealand, and, with the exception of three species recorded in the previous paper on Batley, all have been collected in the South Island. They were first mentioned by Hector, and have been classified in papers written by Woods (57, p. 34), Trechmann (56, p. 338), and Spath (54). The species hitherto recorded are as follows: Madrasites haumuriensis Hector, Madrasites sp., Gaudryceras jukesi Sharpe (Whit-eaves), Gaudryceras sacya Forbes, Grossouvrites gemmatus Huppe, Baculites cf. vagina Forbes, Kossmaticeras zelandicum Marshall, Kossmaticeras tenui-costatum Marshall, Madrasites bhavani Forbes, Lytoceras sp.

The latest reference to the Upper Cretaceous ammonites of New Zealand is that of Spath (53, p. 299):—

“The writer has lately recognized the presence in the Upper Senonian of New Zealand of Kossmaticeras (Madrasites) bhavani Stol. sp., and K. (M.) cumshewaense (? Whiteaves) Kilian and Reboul1, as well as of K. (Gunnarites) aff. bhavaniforme Kilian and Reboul, K. (Grossouvrites) gemmatum Huppe sp., and of Pseudophyllites (Tetragonites) sp. juv.2

“Footnote (1): Possibly close to the incompletely known K. (Madrasites) mcKayi Hector sp. (Catal. N.Z. Court, 1886, p. 57, text-fig. 19a, No. 4) (misspelt maeCoyi in Haug. Traite, 2, p. 1345), that Steinmann (loc. cit., 1895, p. 28) considered to belong probably to the group of K. (M.) aemilianum Stol. sp.”

“Footnote (2): In collections kindly sent by Mr. Henry Woods, F.R.S., and by Dr. Trechmann (see Geol. Mag., n.s., dec. 6, vol. 4, 1917, p. 338).”

In the following pages sixty species are recorded, and all are described. This is done even when the specimens are identified as species already described from other countries, a practice adopted for the following reasons: (1) The literature in which the descriptions of these Upper Cretaceous ammonites have been published is very scattered and most difficult to obtain in New Zealand; (2) the distinction of the different species is carried to such a degree of refinement that the identifications made in this paper may be questioned by workers elsewhere, and in such cases it is well to have a description and a figure to which reference can be made; (3) the great distances between some of the countries where a few of the species have been recorded, and the absence of actual specimens for comparison, make identification difficult and even hazardous.

Geological Occurrence.

The formation in which the fossils occur is an unctuous mudstone which becomes arenaceous in places and even conglomeratic, and it contains many concretions varying in size from 5 ft. in diameter downwards. The great majority of the fossils have been found in the concretions, which are sometimes composed of spherical aggregates of calcite crystals—“pseudopisolitic”—and often show on the exterior an extremely well developed layer of cone-in-cone structure. The development of these structures has sometimes obscured certain features of the fossils, and in particular the suture-lines are occasionally hard to follow. The matrix of the specimens is also extremely hard, and the fossils are often fractured while they are being extracted. On the other hand, much of the shell matter remains, and in the great majority of instances the suture-lines are remarkably clear and easy to follow.

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The thickness of the formation in which the fossils are found is difficult to estimate. On the foreshore of the harbour where the strata occur the whole outcrop is covered with debris, for the unctuous nature of the rock allows of earth-sliding to an unusual degree, and it is rarely that any portion of the cliff-face can be seen. Above the immediate shore-line the whole surface of the hill is covered with vegetation, which effectually con ceals the rock from observation. Here and there, where a small exposure is visible, the strata stand at a high angle. The average direction of the strike at Bull's Point is 285°, and at Batley 295°. In both localities the ammonite-bearing strata are succeeded by a fine-grained chalky limestone, composed for the most part of tests of Globigerina—the so-called “hydraulic limestone” (49, p. 440).

At Bull's Point there are occasional nodules of barite, and frequently boulders of the lower part of the hydraulic limestone are thickly impregnated with black manganese oxides. Concretions of pyrite, also, are of frequent occurrence; and in the interior of many of the concretionary boulders of the ammonite beds, which are often septarian in nature, it is not unusual to find well-developed crystals of calcite. It is often found that a portion of the hydraulic limestone is siliceous and even flinty. This is probably due to the presence of siliceous organisms such as diatoms, radiolaria, and sponge-spicules, the occurrence of which has been reported by Marshall in similar material obtained from neighbouring localities (49, p. 434). Manganese dioxide, barite, and pyrite are found in far greater abundance on the northern shore of the Pahi arm, near Mr. Blackwell's house. Mr. Blackwell was good enough to point this out. A similar association is found on the eastern shore of the inlet behind the Batley Peninsula. In all these instances the occurrence of the minerals seems to be in the material near the base of the hydraulic limestone.

At Whangaroa the strata that contain the fossils are very similar to those at Bull's Point; but they appear to lie directly beneath the Wairakau breccia of Bell and Clarke (47, p. 65). No actual contact of the two formations could, however, be observed. The geology of this district was first described by McKay in 1892, but he did not collect any Upper Cretaceous fossils on the Whangaroa, except specimens of the genus Inoceramus (18, p. 68). He does, however, record ammonites as occurring in concretions in the south branch of the Kaeo River, In 1875 Mr. R. Bell, who then lived on the north side of the Whangaroa Harbour, near Totara North, sent several specimens to the Auckland Museum. Among these is a poor specimen of Gaudryceras kayei Forbes, and an excellent specimen of Tainuia multispinosa n. sp., which has also been found at Bull's Point. There is also a specimen of Phylloceras minimum n. sp., numbers of which have been found in the Kaipara localities. Bell's specimens probably came from the south side of the harbour, for in his letter to the curator of the Museum he says that the specimens in “sandstone” came from the south side of the river. Though “sandstone” is not a correct term to apply to the matrix of the specimens, it is certainly more nearly correct than “shale”—the only other material mentioned by Bell. The matrix is very similar to that of the fossils since obtained on the south side of the harbour. On both sides of the harbour concretions have been much used for roadmaking, and at the present time few of them can be found except at Nedler's Point, near the head of Waitapu Bay, about 200 yards distant from the road.

Clarke in 1909 (47, p. 56) stated that the concretions at Nedler's Point contained numbers of fossils, though the specimens he collected were few

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änd in bad preservation. However, he definitely stated that ammonites could be found in the concretions at Nedler's Point and made it possible for subsequent collectors to obtain specimens easily. In April, 1923, a considerable number of fossils, which are described in the following pages, were found in the concretions at Nedler's. Their nature and geographical occurrence will be discussed after the species in the whole collections have been recorded and described. The outcrop of rock is small, and perhaps rather less unctuous than at the Kaipara localities, but otherwise there is little difference. There are not many concretions to be found, but a larger proportion of them is fossiliferous than at Batley and Bull's Point. On the north side of the harbour there are now no concretions remaining, and no ammonites were found by me, but a few large masses of limestone contained numerous fragments of Inoceramus and specimens of another large species of pelecypod.

In addition to the localities mentioned above, Mr. Ferrar, of the Geological Survey, has recently collected specimens of ammonites at Te Opu, which is quite close to Bull's Point. None of the species found there are different from those found at the latter locality. He also found a few species at Matakohe, about six miles to the north-west. There are also a number of specimens of Vertebrites murdochi in the Auckland Museum. These were obtained at Hokianga, though the exact locality is not recorded. In the same place there is also a good specimen of Baculites rectus and one of Gaudryceras semileve, said to have been found at Morant's Island, in the Kawhia Harbour. This is probably an error, for Morant's Island is composed entirely of a glauconitic limestone, containing a great number of Tertiary fossils, whereas the matrix of the ammonite specimens is a shale. It is possible that “Kawhia” is a lapsus calami for “Kaipara.”

In the collection of the Geological Survey there are two ammonites collected by McKay at Awanui, near the East Cape; but subsequent collectors have failed to find any additional specimens. The two species are similar to some found at Bull's Point. One is close to Zelandites kaiparaensis, and the other is probably Schluteria rarawa. Ongley has recently found a small ammonite at the Pourere Beach, about forty miles south of Napier, but unfortunately it is not in a condition that admits of identification. He has also found a few impressions of ammonites near Whatatutu, inland from Gisborne.

All these localities are shown on the accompanying map, and it is evident that we now have definite information of the occurrence of a considerable extent of Upper Cretaceous strata in which ammonites have been found. There is reason to hope that a large fauna will ultimately be discovered. A map of the Kaipara will be found in 49, p. 435, and a map of Whangaroa in 47, map 3.

I wish to record my grateful thanks to the following institutions and friends: To the authorities of the South Kensington Natural History Museum, for giving me the use of a room and for allowing me to use their vast collections for comparisons; to the authorities of Victoria University College, Wellington, for giving me the use of a room in which to do my research; to the New Zealand Institute, for allowing me a grant of £40 from the Hutton Fund, and one of £50 from the Research Grant, to defray some of the expenses incurred. The following gentlemen have materially assisted me in collecting material: Mr. J. A. Bartrum, Lecturer on Geology, Auckland University College; Professor Cotton, Victoria College; both

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of these gentlemen kindly allowed me to use the resources of their laboratories. Mr. R. Brown and Mr. W. Blackwell also assisted me most materially in making the collections, and R. Marshall also added important specimens. Dr. L. Spath and Professor W. Kilian kindly examined many of my specimens and gave me most valuable advice. Dr. J. Marwick kindly copied drawings of the cross-sections for publication. The Australian Museum generously granted me a long loan of Kossmat's work; this is out of print and there are no copies in New Zealand.

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Sketch-Map of New Zealand, showing Outcrops of fossiliferous rocks of upper cretaceous age.

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Descriptions of Species.

Dimensions.—In all cases the first column under each species gives the actual measurement; the second column gives the proportional measurement, assuming the diameter to be 100 mm.

All dimensions are given in millimetres, and all types, except four—Maorites tenuicostatus, Gunnarites zelandicus, Oxybeloceras sp., Tainuia aucklandica—are at present in my collection.

Phylloceras Suess.

Phylloceras nera Forbes. (Plate 19, fig. 4; Plate 26, figs. 1, 2.)



Phylloceras nera Forbes (1, p. 106, pl. 8, fig. 7).


Phylloceras velledae Mich., in Stol. (11, p. 116).


Phylloceras nera Forbes, in Koss. (28, p. 109, Taf. 16, fig. 2 a—d).


Phylloceras sp. Woods (57, p. 331, pl. 41, fig. 4).


Phylloceras nera Forbes, in Paulcke (42a, p. 3, Taf. 14, fig. 5 a, b, c).


Phylloceras nera Forbes, in Spath (59, p. 40).


Phylloceras woodsi van Hoepen (52, p. 3, pl. 2, figs. 1–6, text-fig. 1).


Phylloceras woodsi van Hoepen, in Spath (54, p. 117).


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A. B. C. D. E.
Diameter 42 100 13 100 24.5 100 46 100 20 100
Height of last whorl 23 54 8 61 14.6 60 28 61 12 60
Width of last whorl 11 26 4 30 7.4 30 13.2 29 6 30
Umbilicus 6 12 2.3 17 2.0 8 2.8 6 1.5 8

A and B, Phylloceras nera Forbes, Bull's Point, Kaipara Harbour, N.Z.; C and D, Phylloceras woodsi van Hoepen (52, p. 4); E, Phylloceras nera Forbes, in Koss. (28, p. 160, Taf. 16. fig. 2, a–d).

Shell discoidal, slightly umbilicated. Whorls strongly involute, high and narrow with greatest thickness about one-third of height measured from umbilicus. Umbilical wall steep and passing gradually into flank, curve gradually increases, and finally periphery is almost a circular arc Umbilicus cannot be properly cleared but is evidently extremely narrow, and a very small portion of any of inner whorls is exposed.

Ornamentation: Numerous thin rounded ribs, twenty in number, on a length of 10 mm. of periphery when height of whorl is 12 mm. They are hardly visible near umbilicus, and have a slight bend forward, but soon straighten up and pass over periphery without any change. On lower part of flank are six conspicuous folds in a half-revolution. These are strongly marked at umbilicus and curve forward in the same way as ribs, but soon weaken, and disappear completely half-way up flank.

Suture-line does not materially differ from that of P. nera figured by Kossmat; but in the largest specimen only five saddles can be distinguished, though in a smaller specimen, in which the details are less clear, eight saddles can be distinguished. The excellent preservation of the large specimen allows all the details to be seen where the whorl height is 19 mm. Owing to the shortness of the external lobe and the large development of the external branch of the first lateral lobe, the overlapping and inclined nature of the first lateral saddle is well shown. This feature was recorded by van Hoepen in the suture-line of Phylloceras woodsi.

The umbilicus of this species is wider than that of P. nera from Patagonia and of P. woodsi of Pondoland, and in addition there are twice as many folds as in the last species. The umbilicus is smaller than in P. decipiens Koss. This species differs from P. velledae in its more compressed form, and the folds disappear more quickly than in P. surya. The suture-line, however, is more like that of P. surya than that of P. nera.

Four specimens have been found, but all are incomplete. It has been found at Bull's Point, Batley, and at Whangaroa.

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Phylloceras radiatum n. sp. (Plate 19, fig. 7; Plate 26, figs. 3, 4.)



Ammonites velledae Mich., Mag. de Zool., pl. 35.


Ammonites velledae Mich., in d'Orb., Terr. Cret. T. I. Ceph. pl. 82.


Ammonites (Scaphites) ramosus Meek, Trans. Alb. Inst., vol. 4, p. 45.


Ammonites velledae Mich., in Pictet et Camp., Terr. Cret. Ste Croix, vol. 1, p. 268, pl. 36, fig. 8.


Ammonites ramosus Meek, in Gabb, Geol. Surv. Canada, Pal., vol. 1, p. 65, pl. 11, fig. 4; pl. 12, fig. 12.


Ammonites velledae Mich., in Stol. (11, p. 116, pl. 59, figs. 1, 4).


Ammonites ramosus Meek, in Gabb (3, p. 65, pl. 11, fig. 4: pl. 12, fig. 12).


Ammonites velledae Mich., in Schmidt, Kreidepetrefacten von Sachalin (Mem. Acad. St. Petersburg), p. 10, Taf. 1, figs. 3, 4.


Phylloceras ramosum Meek, in White. Bull. U.S. Geol. Surv., vol. 4, p. 372, pl. 5, fig. 1.


Ammonites velledae Mich., in Whiteaves (8, p. 103).


Phylloceras velledae, Mich., in Yokoyama (17, p. 177, Taf. 19, fig. 1).


Phylloceras ramosum Meek, in Steinmann (26, p. 80, Taf. 5, fig. 4 a, b).


Phylloceras ramosum Meek, in Anderson (33, p. 34).


Phylloceras velledae Mich., in Boule, Lemoine, and Thevenin, Ceph. Diego Suarez, p. 7, pl. 1, figs. 6, 10, 11.


Phylloceras (Schluteria) ramosum Meek, in Kilian and Reboul (46, p. 9, pl. 1, fig. 3).


Phylloceras umzambiense van Hoepen (51, p. 142, pl. 24, figs. 1, 2, 3).


Phylloceras umzambiense van Hoepen, in Spath (54, p. 117).


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A. B. C. D. E.
Diameter 43 100 32 100 150 100 44 100 38 100
Height of last whorl 25 58 56 57 27 62 16 42
Width of last whorl 18 42 26 58 17 40 13 34
Umbilicus 2 5 9 6 2 5

A. Phylloceras radiatum n. sp., Kaipara Harbour, N.Z.; B, Phylloceras surya Forbes, in Stol. (11, p. 115, pl. 58, fig. 5); C, Phylloceras velledae Mich., in Stol. (11, p. 116, pl. 59, figs. 1–4); D, Phylloceras umzambiense van Hoepen (51, p. 142); E, Phylloceras velledae Mich., in Yokoyama (17, p. 177, pl. 19, fig. 1 a, b).

The dimensions show that this species is distinct, though it approaches D.

Shell discoidal, strongly involute, and higher than wide. Greatest thickness rather nearer to umbilicus than to external border. Flank has a uniform curve from umbilicus almost to median line. Umbilicus deep and extremely narrow.

Ornamentation: Numerous narrow ribs steeper in front than behind. They commence as extremely fine lines at bottom of umbilicus, and gradually increase in size towards external margin, when at a height of 12 mm. there are eighteen ribs on a length of 10 mm. of periphery. Within umbilicus they are curved strongly backwards, but on leaving it curve a little forward, but straighten half-way between umbilicus and periphery, which they cross without any curve. As far as can be seen, there are no interstitial ribs. A series of wide folds, twelve in a half-whorl, extend from umbilicus across periphery, but they lose somewhat in height as they approach periphery. There are four or five ribs on each of the folds, which do not curve like the ribs, but are strictly radial in their direction. The folds seem to be more pronounced on surface of body-chamber, but the specimens are too imperfect to be certain of this.

The dimensions given above show that the species is much wider than Phylloceras surya and a good deal narrower than P. velledae. In dimensions and form of whorl it approaches more closely to P. umzambiense than to any other species of which details are available. Umbilical wall not so steep as that of P. ramosum Meek in Steinmann, and the whorl appears rather wider than that of P. ramosum Meek in Kilian and Reboul.

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Suture-line a good deal less divided than in P. ramosum Meek in Steinmann, and less than that species in Kilian and Reboul as far as can be seen from their figure. It is, however, of a typical Phylloceratid type, rather more complex than that of P. velledae, and decidedly more so than the suture-line of Phylloceras ellipticum Koss., a species that it closely resembles in the details of ornamentation. Phylloceras radiatum clearly belongs to the velledae group of the genus, which persisted through all Upper Cretaceous times. The marked radial folding, combined with the form of the whorl and the nature of the costation, seem to me to distinguish it from all others. In the costation it comes nearer to P. ellipticum Kossmat; in the form of the whorl and the folding the closest species is P. umzambiense; while in the development of the suture-line it lies between P. ramosum and P. velledae.

The internal portion of the suture shows only three saddles. The only figures available for comparison are those given in Zittel's Handbuch, vol. 2, p. 436. Here the line of a Dogger species is shown, but it has six saddles. It is interesting to note that Sowerbyceras has three saddles.

One good specimen and two fragments from Bull's Point, and one fragment from Whangaroa.

Phylloceras forbesianum d'Orbigny. (Plate 19, fig. 6; Plate 27, figs. 3, 4.)



Ammonites rouyanus d'Orb., in Forbes, Geol. Soc. Lond., ser. 2, vol. 7, pl. 8, fig. 6, p. 108.


Ammonites forbesianus d'Orb., Prodrome, vol. 2, p. 13.


Ammonites rouyanus d'Orb., in Stol. (11, p. 117, pl. 59, figs. 5–7).


Phylloceras ezoense Yokoyama (17, p. 178, pl. 19, fig. 1 a–c).


Phylloceras forbesianum d'Orb. and P. whiteavesi Koss., in Kossmat (28, p. 109, pl. 15, fig. 1; also p. [189]).


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A. B. C. D. E.
Diameter 25 100 18 100 13 100 14.5 100 28 100
Height of last whorl 15 60 12 66 8.2 63 8.5 58 18 64
Width of last whorl 15 60 12 66 8.2 63 9 62 17 61
Umbilicus 1 04

A, Phylloceras forbesianum d'Orb., in Koss. (28, p. 109, pl. 15, fig. 1 a–d); B, C, D, E, Phylloceras forbesianum, Bull's Point, Kaipara Harbour, N.Z.

Whorls rapidly increasing and extremely involute, so that umbilicus is practically obscured. Height and width almost equal, and cross-section almost circular. This is the same as in dimensions of P. forbesianum given by Kossmat, though the figure shows a far greater height than width. The whorl rises at first steeply from umbilicus, but curves uniformly in circular arc and has its widest part about half-way between umbilicus and periphery.

Ornamentation: Comparatively little of the material of the shell remains, but this shows a fine striation beginning on umbilical wall, at first bending backward strongly, but at half-way along flank it is succeeded by a bend forward; thereafter the striae become straight and cross periphery without a bend. After every seventh or eighth rib is a decided groove, most noticeable near umbilicus but becoming obscure on periphery.

Suture-line very similar to that given by Kossmat for Phylloceras forbesianum, though it is rather more complex and there is a far greater number of saddles, of which as many as twelve can be distinguished. These, of course, are very small in umbilical region, and they are much higher than wide. The external lobe is unusually deep. The external saddle is

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largest, and there is a uniform decrease in size as umbilicus is approached. This species differs from P. forbesianum in dimensions of whorl if Kossmat's figure is correct, though the measurements that he gives agree closely with those of this species. Suture-line of this species also is rather more developed than that of the Indian one. Yokoyama compares his species P. ezoense with P. forbesianum, but states rightly that the whorl is too high. His figure also suggests that the costation is too coarse.

Kossmat, in an appendix to his work on the Indian ammonites, separates the specimens which in the earlier part of his work he had classed together under P. forbesianum into two species, P. forbesianum and P. whiteavesi. Some of these specimens had come from the Utatur formation, of Cenomanian age, and others from the Valudayur beds, of Upper Senonian age. In the appendix he retains the former under the name of P. forbesianum, and establishes the species P. whiteavesi for the latter. The former species is somewhat more inflated and does not show the funnel form of the umbilicus so distinctly as the latter. In addition the saddles of the suture-line are larger, and the median saddle is simple and lancet-shaped, while in the two other occurrences (Valudayur beds of Pondicherry and the Nanaimo beds of Vancouver) it is strongly indented. He says that these are truly small distinctions, but they accord with different geological occurrences, so that great importance must be attached to them. He attaches great importance to the relation between the ultimate and penultimate whorls. In the specimens of P. forbesianum from the Utatur formation the relation in different specimens is 1: 2.75, 1: 2.5, 1: 2.6, while in a specimen of ezoense which is similar to P. whiteavesi it is 1: 3.2 (11, p. 124).

In this respect the New Zealand specimens are similar to P. forbesianum, for the relation is 1: 2.5 and 1: 2.25. In other words, the increase of the size of the whorls of the New Zealand species is less rapid than even that of the Utatur species. On the other hand, in the details of the suture-line there is a greater similarity to P. whiteavesi, for the median saddle is sharply indented and at the same time the saddles are relatively longer. It seems, then, that this species has some of the determining characters of each of the related Indian ones.

Several specimens, most of which come from Bull's Point, though others come from Batley and from Whangaroa.

Phylloceras minimum n. sp. (Plate 19, fig. 8; Plate 26, figs. 5, 6.)


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A. B.
Diameter 13 100 8.5 100
Height of last whorl 7.5 57 5 58
Width of last whorl 7 54 4.5 53

A, B, Phylloceras minimum, Kaipara Harbour, N.Z.

Shell small, highly involute, and, as dimensions show, height and width almost equal. Umbilicus obscured. There is a steep slope from umbilicus, and the greatest thickness is reached at a distance of about one-third of height from it, whence the curve is that of a circular arc over periphery. A series of fine hair-like striations commences at umbilicus, and at first are directed backwards; then they bend forwards and then straighten towards periphery, which they cross without any bend. They remain fine, but are much more distant on periphery.

The suture-line is highly developed when the size of the species is considered. The saddles are relatively long and well separated from one

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another. There is also a large number of auxiliary saddles. In this species, again, the internal portion of the suture-line has three saddles only. The internal saddle, as before, has the curious form characteristic of the genus.

This species is very similar to the last, but the ornamentation is far less distinct, and the whorl is wider. The suture-line also is more developed.

A number of specimens of this small species from Bull's Point and from Batley.

Phylloceras bistriatum n. sp. (Plate 19, fig. 5; Plate 27, figs. 1, 2.)



Ammonites subalpinus d'Orb., in Stol. (11, p. 114, pl. 58, fig. 3).


Phylloceras ellipticum Koss. (28, p. 107, pl. 15, fig. 2).


Phylloceras ezoense Yokoyama (17, p. 19, pl. 2, fig. 178).


Phylloceras ezoense Yabe (61, p. 54, pl. 8, fig. 2).


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A. B.
Diameter 43 100 25 100
Height of last whorl 27 63 15 60
Width of last whorl 22 51 15 60
Umbilicus 1 4

A, Phylloceras bistriatum n. sp., Bull's Point, Kaipara Horbour, N.Z.: B, Phylloceras forbesianum d'Orb., in Koss. (28, p. 109, Taf. 15, fig. 1).

A discoid highly involute form, with umbilicus reduced to smallest dimensions. Cross-section nearly circular, but there is a distinct angle at top of umbilical wall. The shell matter is poorly preserved, but it can be seen that interior whorls have an extremely fine, somewhat distant striation. On body-chamber there are in addition a number of large rounded ribs, which are narrow at umbilicus and gradually widen as they approach periphery, which they cross without change. The fine striations can be seen between and on the slopes of these large ribs.

Suture-line: Unfortunately only five of the saddles can be exposed. They are less finely divided than those of the other species, and the terminations are rounded and coarse. The secondary saddles in the lobes are but little divided.

So far as the suture-line is concerned, this species approaches most closely to the improvisum-ellipticum group in the Indian Lower Cretaceous fauna. It comes very close to P. ezoense Yokoyama (in Yabe, 1921).

A single specimen, in fair order, at Bull's Point.

Vertebrites n. gen.

It is considered necessary to establish this new genus because, though numerously represented by many specimens in splendid condition, it cannot be included in the genus Gaudryceras as emended by Kossmat (28, p. 113). The particular distinction is found in the internal portion of the suture-line, which shows six distinct saddles gradually decreasing in size from the antisiphonal line to the umbilicus. This condition is wholly different from that described by Kossmat as characteristic of Gaudryceras, which has a high single and narrow saddle in the internal portion of the suture-line. It may, of course, be maintained, as is suggested by Spath (53, p. 239), that such a development “is a result of the adaptation of a suture-line to wider sides.” This, however, does not seem to be a sufficient explanation in this instance, for the various species of the closely related

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genera Tetragonites and Pseudophyllites, in which the reduction of the sides is even more extreme than in the species of Gaudryceras, have an internal suture-line in which there are two well-separated saddles.

The species placed in this genus is clearly the New Zealand representative of the well-known and widely-distributed Indo-Pacific species Gaudryceras kayei Forbes. It seems, however, that the internal suture-line of that species has not yet been observed; and if, as seems probable, it should prove to be similar to that of Vertebrites murdochi it will have to be included in this new genus. It is not the suture-line alone that distinguishes these forms from the typical species of Gaudryceras: there is in addition the curious low and wide shape of the whorl, as well as the conspicuous difference in the ornamentation on the flanks as compared with that on the periphery.

The name Vertebrites is given to the genus because of the resemblance of a cross-section to that of a fish-vertebra.

Vertebrites murdochi n. sp. (Plate 20, figs. 9, 9a; Plate 30, figs. 1, 2; Plate 40, fig. 3.)



Ammonites kayei Forbes (1, p. 101, pl. 8, fig. 3).


Ammonites kayei Forbes, in Stol. (2, p. 156, pl. 77, fig. 3).


Ammonites kayei Forbes, in Griesbach, Q.J.G.S., vol. 27, p. 63.


Ammonites jukesi Sharpe, in Whiteaves (8, p. 111, pl. 13, figs. 3 a, b, c).


Gaudryceras planorbiforme Bohm, in de Grossouvre (19, p. 231, pl. 27, fig. 2).


Lytoceras kayei Forbes, in Steinmann (26, p. 86, pl. 5, fig. 5).


Lytoceras (Gaudryceras) kayei Forbes, in Kossmat (28, pp. 124, 162, pl. 16, fig. 5; pl. 17, fig. 2).


Lytoceras (Gaudryceras) kayei Forbes, in Anderson (33, p. 83).


Lytoceras (Gaudryceras) kayei Forbes, in Kilian and Reboul (46, p. 12).


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A. B. C. D. E. F. G.
Diameter 25 100 27 100 27 100 42 100 36 100 29 100 26 100
Height of last whorl 7 28 7 26 6.5 24 10.25 24 9 25 7.75 27 6.75 26
Width of last whorl 10.5 42 11 41 11 41 16 39 14 39 13 44 12.2 47
Umbilicus 14.5 58 14 53 15 55 25 60 21 58 16 55 14.8 57
H. J. K. L. M. N. P.
Diameter 29.8 100 30 100 31 100 28.8 100 30 100 26.2 100 10 100
Height of last whorl 7.0 24 7.4 25 7.2 23 7.2 24 6.9 23 5.9 23 2.2 22
Width of last whorl 11 37 11 37 13.2 43 12.6 43 13 43 11.8 45 5 50
Umbilicus 18 60 16 53 15.4 53 16.0 55 17 57 14.8 56 6.2 62
Q. R. S. T. V. W. X.
Diameter 22.5 100 58.5 100 12.6 100 30 100 57 100 46 100 24 100
Height of last whorl 6.5 29 18.5 32 3 24 8 27 19 33 15 33 8 33
Width of last whorl 9.5 33 18 31 5 40 10 33 18 32 15 33 9 37
Umbilicus 12.0 53 27.5 47 7 56 16 53 28 50 23 50 13 55

A, B, C, Vertebrites murdochi, Batley, Kaipara, N.Z.; D, E, F, G, H, J, K, L, M, P, Vertebrites murdochi, Whangaroa, N.Z.; N, Vertebrites murdochi, Hokianga, N.Z.; Q, Ammonites kayei Forbes, in Steinmann, l.c.; R, S, Gaudryceras kayei Forbes, in Koss., l.c.; T, V, W, X, Gaudryceras kayei Forbes: specimens in Kaye's collection in British Museum, Natural History Branch, South Kensington, London; measured by P. Marshall.

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These dimensions clearly show that the New Zealand specimens from three localities belong to the same species, and they differ from the typical Indian specimens in having a distinctly lower and also wider whorl and a slightly wider umbilicus. It might appear as though the dimensions of S contradicted this statement; but small specimens are all wider, and S should be compared with P.

Shell discoidal, with an involution of about one-third. Whorls much wider than high, with a surface rising steeply from umbilicus and passing with a gradually decreasing curve to periphery, where it is much flattened. The deep umbilicus has a steep slope almost uniform except for upper part of each whorl, which breaks the regularity with rather sharp curve. Whorls do not increase rapidly, and seven can be counted when shell has a radius of 10 mm. Strong sharp wire-like ribs start at bottom of umbilicus and are at first radial, directed slightly backwards; almost at once they curve strongly forwards, but become almost radial again on upper part of flank at point where covered by succeeding whorl. A little below this an interstitial rib generally appears between each pair of wire ribs, and at this line each rib usually breaks up into three fine striations. Other striations also take their origin at this level, though the original rib often continues straight on, but with the dimensions of one of the striations only. These striations at once bend forward and cross periphery with a gentle broad forward curve. On body-chamber the coarse wire ribs become much less numerous and are irregularly spaced, but the fine striations maintain their number and appearance.

The external portion of the suture-line is similar to that shown by Stoliczka for Gaudryceras kayei; Kossmat does not give a drawing of it. Six saddles can be distinguished, the external by far the largest. Auxiliary saddles much inclined, and with auxiliary lobes form a deep umbilical lobe. External lobe a little deeper than first lateral lobe. Internal portion of suture-line at once distinguishes the genus. Instead of the single lengthened saddle which is found in Gaudryceras, six distinct saddles can be seen. Of these the first, which is close to the antisiphonal line, is the largest and is bifid, though the division is not deep. The next saddle is shorter, and the size afterwards rapidly decreases to umbilicus. Second and third saddles are bifid. Antisiphonal lobe a little deeper than lobe next to it, but not nearly so deep as the decline of the whole suture-line on umbilical margin. Lobes all simply bifid.

The specimens that have been described show a close resemblance to the specimens of Gaudryceras kayei in the collection at the British Museum, Natural History Branch, at South Kensington. These are in the collection made by Kaye and described by Forbes. In the New Zealand specimens the periphery is somewhat flatter, sculpture coarser on umbilical surface, and finer on peripheral region, where also striae have a stronger forward bend. The width which Stoliczka mentions as so remarkable is still more marked than in the Indian specimens of the Kaye collection, though the measurements of the drawings given by Stoliczka show proportions almost exactly the same as in the New Zealand specimens. Yabe (37, p. 25) suggests that various specimens of G. kayei from South India, as well as Steinmann's specimens from Quiriquina, and the Griesbach specimen from South Africa, should be included in his species Gaudryceras tenuiliratum var, ornatum. The specimen of this species figured by him is, however, clearly a close relative of G. sacya, and quite different from the true

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G. kayei in form and sculpture. The specimen in the Griesbach collection has now rightly been recognized as a distinct species by van Hoepen, and he has named it G. varicostatum, which, as shown in this paper, is quite close to the New Zealand species G. semileve. G. jukesi Sharpe in Whiteaves (8) is certainly this species, and is quite different from the species that he refers to under G. jukesi later (27). There is no description of the forms identified by Anderson in California and by Kilian and Reboul in Seymour Island as G. kayei, but there is little doubt that they are typical forms of this species.

This species has been found at Batley and Bull's Point in the Kaipara, and plentifully at Whangaroa, where it seems to be the species called Desmoceras sp. by Clarke (47, p. 56, pl. 12, fig. 3 a, b).

Three specimens in the Auckland Museum come from Hokianga.

It is stated by Kossmat, and the remark is quoted by Kilian and Reboul, that G. kayei Forbes is closely related to G. planorbiforme Bohm from the Upper Cretaceous of Bavière. I have no specimens of this species for comparison, but, basing my opinion upon the description and figures given by de Grossouvre, I am unable to agree that the relationship is close. The specific name is given in memory of the late R. Murdoch, a well-known New Zealand conchologist.

Gaudryceras de Grossouvre (19, p. 225).

The following is the diagnosis of the genus given by de Grossouvre: “Ce genre est caracterisé par l'allure des striés de la coquille partant de l'ombili infléchies en avant, et par une ligne suturale formée de lobes et de selle assez nombreux, les lobes étant tous à terminaison paire.” He lays stress on the striae being directed back from the edge of the umbilicus. The sutural line is not so much reduced in auxiliaries as in Lytoceras. In the greater number of species the whorls increase slowly, and are but slightly involute in the young, but afterwards increase more rapidly, especially in height.

Kossmat later emended the description (28, p. 113). Slightly involute forms, with periodic constrictions forming bolsters on the shell or furrows on the cast. These, like the fine wire-like ribs, start at right angles to the umbilicus, and in the under half of the flank bend forward and in the upper half backward, forming a shallow forward bend on the periphery. The suture-line is of the greatest importance for the whole group. The median saddle is small, like a spear; the external lobe long and small, as deep as the first lateral lobe. The bifid external saddle is conspicuous. There are always several auxiliary lobes, which form a distinct umbilical lobe. In the internal portion there is a small and deep antisiphonal lobe, which goes as far back, or farther, than the umbilical lobe. A long tree-like saddle separates the two lobes, and on its side it is deeply dissected by sharp points. The bottom of the deep highly characteristic umbilical lobe is two-pointed, like the antisiphonal lobe. (Translation of extract.)

The nature of this internal suture-line separates Gaudryceras from Tetragonites. All species of Gaudryceras have at the base of the partition wall a single deep groove which corresponds in position to the antisiphonal lobe, while Tetragonites, with a greater number of internal lobes, has more than one on each side of the antisiphonal line.

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Gaudryceras propemite n. sp. (Plate 20, fig. 4; Plate 28, figs. 3, 4.)



Ammonites sacya var. multiplexus Stol. (11, p. 155, pl. 76, fig. 1).


Lytoceras mite von Hauer, Neue Cephalopoden der Gosaugebilde, Sitz. Akad. der Wiss. Wien, Bd. 53, p. 7, Taf. 2, figs. 3, 4.


Ammonites sacya var. sachalinensis Schmidt (4, p. 15, Taf. 2, figs. 1, 2, 6.


Gaudryceras mite Hauer, in de Grossouvre (19, p. 227, pl. 26, fig. 4; pl. 39).


Gaudryceras jukesi Sharpe, in Whiteaves (27, p. 129, pl. 2, figs. 1, 2).


Lytoceras (Gaudryceras) multiplexum Koss. (28, p. 121, pl. 15, fig. 6).


Lytoceras varagurense var. patagonicum Paulcke (42a, p. 6, Taf. 17, figs. 1, 2).


Gaudryceras aff. jukesi (Whiteaves) Sharpe, in Woods (57, p. 35, pl. 20, fig. 2).


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A. B. C. D.
Diameter 64 100 78 100 101 100 64 100
Height of last whorl 24 37 23 29 40 40 25 40
Width of last whorl 28 44 24 30 38 38 26 41
Umbilicus 19 30 38 49 35 35 24.5 39

A, Gaudryceras propemite, Bull's Point, Kaipara Harbour, N.Z.; B, Lytoceras (Gaudryceras) multiplexum Koss. (28, p. 121, Taf. 15, fig. 6); C, Lytoceras (Gaudryceras) varagurense (28, p. 122, Taf. 18, fig. 2, pl. 17, fig. 9): D, Lytoceras (Gaudryceras) vertebratum Koss. (28, p. 126, Taf. 15, figs. 4, 5).

The dimensions given above show that the breadth and height of the whorl are almost equal, and that they come nearest to G. vertebratum of the Indian species.

Form discoidal, with an involution of one-third. Seven whorls when diameter is 64 mm., with its greatest thickness just above slope to umbilicus; from this point there is a gradual slope to periphery, which is evenly rounded. Whorls slightly wider than high; whorls do not increase rapidly in height.

Ornamentation: A large number of rounded ribs of moderate size, which on body-chamber begin rather indistinctly near base of umbilical wall. They rapidly increase in distinctness, and at top of umbilical slope many divide into two, and other interstitial ribs arise, their number being increased about twofold. Ribs at first almost radial, but soon bend strongly forward, then distinctly backward, and at periphery again slightly forward so that they cross it with a slight forward curve. Ornamentation of inner whorls the same up to the point where they are covered by the next succeeding whorl; here they break up into striations, and similar additional striations arise interstitially and cross periphery with a slight forward bend. There are periodic furrows, sometimes very distinct, about four in a whorl.

Suture-line does not differ to any important extent from that of the different coarse-ribbed forms of Gaudryceras from South India which have been mentioned above. On the whole, it is rather more like that of G. vertebratum than the others. The most distinctive character is in the development of the median secondary saddle in the first lateral lobe.

The species is evidently very close to the G. vertebratum-varagurense group in all respects, but in none of them is there any mention of the curious change in the nature of the sculpture in all of the inner whorls. Gaudryceras cinctum Spath, from Pondoland, is also close, as is G. mite Hauer, from the Santonian of Europe. Again, G. varagurense var. patagonica Paulcke, from Patagonia, is closely related, but the details of the ornamentation seem to be distinct. G. varagurense and G. mite have been recorded by Kilian and Reboul from Seymour Island, but in both cases

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it appears that the state of preservation does not allow any details of the ornamentation to be seen.

A single complete specimen has been found at Bull's Point, as well as several fragments.

G. multiplexum and G. vertebratum come from the Utatur formation, and G. varagurense from the Upper Trichinopoly (= Lower Senonian) of Europe.

Gaudryceras particostatum n. sp. (Plate 20, fig. 7; Plate 30, figs. 3, 4.)



Gaudryceras kayei Forbes, in Woods (40, p. 335, pl. 41, fig. 8).


Gaudryceras varicostatum van Hoepen (52, p. 7, pl. 2, figs. 10–12, text-figs. 3, 4).


Gaudryceras varicostatum van Hoepen, in Spath (54, p. 117).


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A. B. C. D.
Diameter 27 100 24 100 33 100 39.6 100
Height of last whorl 10 37 9 38 13 39 13.7 35
Width of last whorl 10 37 9 38 11 34 14.7 37
Umbilicus 10 37 11 45 13 39 17.2 43

A, B, C, Gaudryceras particostatum, Bull's Point, Kaipara Harbour, N.Z.: D, Gaudryceras varicostatum van Hoepen (52, p. 7).

The dimensions show at once that height and width of whorl are almost equal, and umbilicus extremely wide. Shell small, with whorls that show little involution and do not increase rapidly in size. Umbilicus wide, with a nearly uniform slope. When the radius is 14 mm. six whorls are developed. The umbilical slope is steep, but passes by a rapid curve into the flank, which slopes steeply to the well-rounded periphery.

Ornamentation: Flanks of inner whorls covered with thin wire-like ribs, which begin at umbilicus and at once bend sharply forward. At edge of periphery, where whorl is covered by the succeeding one, the wires change into a number of minute striations, which require a high-powered lens to show them up. These have a slight bend forward where they cross periphery. On body-chamber the wire-like ornamentation cannot be seen, and its place is taken by fine striations; these are not easily seen, and without a careful inspection the surface appears to be smooth. Four conspicuous constrictions in a revolution.

Suture-line in both external and internal portions very similar to that of Gaudryceras varicostatum. Antisiphonal lobe differs from typical form in its greater breadth. External portion of suture-line is quite typical of the genus, though, on the whole, it is perhaps rather less deeply dissected. Umbilical lobe unusually deep and extremely steep. The width of the single saddle in the internal portion of the suture-line is noticeable, and in this respect this species resembles G. propemite, G. varicostatum, and G. varagurense, though it is less marked in the last species.

This species certainly comes nearer to Gaudryceras varicostatum than to any other that could be found in the available literature. It is, however, distinguished by the finer ribbing and the somewhat higher whorl. There seems to be no Indian species to which it shows a close resemblance, for it has a higher whorl and smaller umbilicus than G. multiplexum; finer ribbing and a wider umbilicus than G. varagurense; a narrower and lower whorl than G. vertebratum, and finer ribbing as well; and in none of these species has it been recorded that the ribbing changes in its character on the upper part of the flank.

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Several good specimens have been obtained at Bull's Point, and one in poor condition from Whangaroa. In the Auckland Museum there is a specimen in good condition labelled as coming from Morant's Island, in the Kawhia Harbour. This, however, I regard as a mistake, as the rocks on that minute island are entirely Tertiary in their age. It is possible that this and Baculites rectus n. sp., which have the same matrix, come from Hokianga or from Kaipara.

Gaudryceras subsacya n. sp., (Plate 20, figs. 8, 8a; Plate 29, figs. 1, 2.)


  • Ammonites sacya Forbes (1, p. 113, pl. 14, fig. 10).

  • Ammonites buddha Forbes (1, p. 112, pl. 14, fig. 9).

  • Ammonites sacya Forbes, in Stol. (11, p. 154, pl. 75, figs. 5–7; pl. 76, figs. 1, 2, 3).

  • Ammonites sacya var. sachalinensis Schmidt (4, p. 15, pl. 2, figs. 3, 4).

  • Ammonites filicinctus Whiteaves (7, p. 43, pl. 2, figs. 2, 3).

  • Lytoceras sacya Forbes, in Whiteaves (7, p. 203, pl. 25).

  • Lytoceras sacya Forbes, in Yokoyama (17, p. 178, pl. 18, fig. 12).

  • Lytoceras sacya Forbes, in Jimbo (22, p. 34, pl. 6, fig. 1).

  • Lytoceras sacya Forbes, in Stanton (20, p. 445).

  • Lytoceras (Gaudryceras) sacya Forbes, in Koss. (28, p. 119).

  • Lytoceras (Gaudryceras) sacya Forbes, in Anderson (33, p. 82).

  • Gaudryceras tenuihratum Yabe (48, p. 19, pl. 3, figs. 3, 4; also var. ornata, pl. 3, fig. 2).

  • Gaudryceras striatum var. pictum Yabe (48, p. 33, pl. 4, fig. 3).

  • Gaudryceras sacya Forbes, in Woods (57, p. 11, pl. 5, figs. 4 a, b).

  • Lytoceras sp. Marshall (49, p. 445, pl. 33, fig. 3, text-fig. 4).

  • Gaudryceras sacya Forbes, in Spath (59, p. 54).


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A. B. C. D. E. F. G.
Diameter 28 100 22.75 100 36 100 29 100 24 100 16 100 45 100
Height of last whorl 10.5 37 9 39 12.5 35 11 38 9 37 6.2 39 33
Width of last whorl 10.75 42 10.25 45 14 39 12 41 10.25 42 8 50 34
Umbilicus 11.5 41 10 44 13.5 38 11.25 39 9 37 7 44 44
H. J. K. L. M. N. P.
Diameter 95 100 40 100 50 100 120 100 113 100 300 100 35 100
Height of last whorl 43 30 34 40 50 44 160 53 12 34
Width of last whorl 36 35 30 38 45 40 165 53 13 38
Umbilicus 32 50 46 36 36 32 55 18 15 42

A, B, C, D, E, F, Gaudryceras subsacya, Bull's Point and Batley, Kaipara Harbour, N.Z.; G, H, J, K, L, Gaudryceras sacya Forbes, in Stolicka (11, p. 154, pl. 75, figs. 5–7; pl. 76, fig. 23); M, Gaudryceras sacya Forbes, in Kossmat (28, p. 119); N, P, Gaudryceras tenuiliratum Yabe (37, p. 19).

The six specimens of G. subsacya that were measured show at once that the species always has a whorl that is wider than high, and that the larger specimens are both lower and narrower than smaller ones. The Indian specimens that were measured by Stoliczka are in most cases higher than wide, and this is also the case in the Indian specimen quoted by Kossmat. Yabe's measurements of Gaudryceras tenuiliratum are much higher and wider in the large specimens than in the smaller. The measurements show that species is characterized by its uniformly wide whorl.

Shell discoidal with wide umbilicus, downward slope of which is modified by rounded contour of inner whorls. Involution about one-third. Slope from umbilicus steep, and gradually rounds off into flank; periphery flattened. Well-preserved specimens show five distinct constrictions marked

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on shell by prominent rounded ribs, which have a large furrow behind and a smaller one in front. Regular sculpture consists of extremely numerous ribs, so fine that when radius is 7 mm. there are 45 on an arc of periphery 5 mm. in length. Large ribs that mark constrictions are covered with these fine ribs, which arise at bottom of umbilical slope and at first have a slight forward inclination. This soon becomes pronounced, but decreases again at about two-thirds of distance from umbilicus. Ribs cross periphery with the very slightest forward bend. Near top of umbilical slope an interstitial rib arises between each pair, and all continue without interruption across periphery. Ribs and sculpture on body-chamber differ in no respect from those on younger whorls, except in size, which increases proportionately to diameter.

Suture-line is typical of the smaller species of Gaudryceras, and by itself can hardly be distinguished except by the slightly displaced position of the small secondary saddle in the first lateral lobe. Internal portion of suture-line also is quite typical of these smaller species, and shows the long and narrow single saddle which Kossmat considers an essential feature of the genus.

The position of this species is very close to G. tenuiliratum var. ornatum of Yabe; but comparison of the measurements shows that the whorl is both higher and wider than in the Japanese species, and at the same time the umbilicus is narrower. The resemblance to G. striatum var. pictum of Yabe is also close, but the ribs appear to be much more flexuous in that species. The specimens considered to be G. sacya by Yokoyama and Jimbo are differently classed by Yabe. Crick's specimen of G. pulchrum in the South Kensington Museum was also compared with those from New Zealand, and it was found that its ribs were distinctly sharper and farther apart than in the present species. Forbes's type of G. sacya is also in the Museum, and though it is in poor condition it is clear that the ribs are a little larger and farther apart than in the New Zealand examples. The species appears to be very similar to G. filicinctus of Whiteaves from the Queen Charlotte Islands. The examples here described from the north of New Zealand are quite close to the specimen from the Clarence Valley of the South Island of New Zealand, which was classified by Woods as G. sacya, for the only observable difference in that the ribs in the South Island specimen are rather coarser than those of all but one of the specimens from the North Island.

This species is common at Batley and at Bull's Point in the Kaipara Harbour, but no specimens were found at Whangaroa.

Gaudryceras sacya comes from the Lower Utatur group. No specimen yet found in New Zealand shows the corrugations which characterize the mature forms of G. sacya as figured by Whiteaves and of G. limatum figured by Yabe.

Gaudryceras politissimum Kossmat. (Plate 20, fig. 3; Plate 28, figs. 1, 2.)



Gaudryceras politissimum Koss. (28, p. 128, Taf. 15, figs. 7 a-c).


Lytoceras (Gaudryceras) politissimum Koss., in Kilian and Reboul (46, p. 14, pl. 1, figs. 7, 8).


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A. B.
Diameter 48 100 89 100
Height of last whorl 20 49 33 37
Width of last whorl 14 29 28 31
Umbilicus 16 33 34 38

A, Gaudryceras politissimum Koss., Bull's Point, Kaipara Harbour, N.Z.; B, Gaudryceras politissimum Koss. (28, p. 128).

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These dimensions show that the only specimen that is in good enough condition to be measured has a whorl a little higher than that of Kossmat's type, while the umbilicus is distinctly narrower. Comparisons with the dimensions of G. subsacya show that the whorl in that species is not so high as, but is much wider than, that of G. politissimum. In the specimen that has been measured the greater part of outer whorl is somewhat compressed. The inner whorls increase a good deal in their relative width, the penultimate whorl measuring—height, 4.5 mm.; width, 5.5 mm.; and the whorl next to that—height, 2.5 mm.; width, 3.5 mm. The broadest part of outer whorl is at top of umbilical wall, where there is a sharp curve to the gently sloping flank; but at periphery the curve is much sharper.

Ornamentation: Five constrictions can be distinguished in the last whorl, which appear as rounded ribs on the shell. They are low where they leave umbilicus. They at once bend forward and increase in size, but the forward curve is soon reduced, and they cross periphery with a slight forward bend. At first sight the surface of the shell appears to be polished, but when examined with a lens a number of regular but extremely fine hair-like ribs can be distinguished. About every tenth or fifteenth of these is larger. They start almost straight from the umbilicus, or with a slight backward trend. They soon bend forward on the edge of umbilicus; then they straighten up, but pass over periphery with a broad forward curve.

Suture-line agrees fairly well with that of G. politissimum figured by Kossmat. The saddles, however, are less divided, the slope to umbilicus begins sooner, and there are fewer auxiliary saddles; umbilical lobe is also deeper.

The specimen agrees quite well with description that has been given by Kossmat, though the relative height and width of inner whorls are not referred to by him. The ornamentation and also the form of G. politissimum, G. subtilineatum Koss. (28, p. 123), and G. valudayurense Koss. (28, p. 127) are very similar, but it appears that the ornamentation of the surface of G. valudayurense is rather stronger and the suture-line is rather more divided. It would be extremely difficult to distinguish the inner whorls of the New Zealand specimen of G. politissimum from those of G. subtilineatum described by Kossmat.

One specimen, in moderately good condition, from Bull's Point, and a second, of the inner whorls only, from Batley.

Gaudryceras crenatum n. sp. (Plate 20, fig. 10; Plate 31, figs. 3, 3a.)


Diameter 27 100
Height of last whorl 9 33
Width of last whorl 10 37
Umbilicus 12 44

A, Gaudryceras c [ unclear: ] enatum, Bull's Point, Kaipara Harbour, N.Z.

Shell discoidal, with wide umbilicus which has very even slope. Involution about one-third. A steep even slope of umbilical wall, which curves rather quickly into flank, and therafter a uniform curve which passes without interruption over rounded periphery.

Ornamentation: Four shallow periodic furrows in last whorl, with occasionally a low rounded rib behind. The surface appears bright and shining, but examination shows that it is closely covered with fine thread-like ribs. These arise at umbilicus and are at first straight, but soon bend

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forward and maintain this direction until they reach beyond umbilical edge; they then bend slightly backwards, but soon again bend forward and pass with a broad curve across periphery. Ribs become less distinct at periphery, but there are no interstitial ribs. Ribs have a sharp crest, but are distinctly crenate on backward side. A good specimen and a high-power lens are necessary to reveal this structure.

Suture-line not well seen. Saddles not so highly divided as in the other species of Gaudryceras. External lobe is perhaps a little deeper than usual, and three auxiliary lobes are present.

Three specimens only, two of which come from Bull's Point and one from Batley.

The crenation of the small ribs is the most distinct characteristic of this species. So far as Cretaceous species of this family are concerned, it has been recorded only in Lytoceras mahadeva Stol., from the Lower Utatur of India; L. batesi, from the lower beds of Queen Charlotte Island of British Columbia; and L. ezoense, from the Lower Ammonite beds of Japan. These are included in the Lytoceras fimbriatum group of Kossmat, which is mainly distinguished by the absence of auxiliary lobes and saddles in suture-line (28, p. 112). The present species has three auxiliary lobes and saddles and at the same time its small size and fine ribbing ally it rather to the Gaudryceras group. It is perhaps permissible to regard the crenation as a vestigial remnant that shows clearly the lytoceratid ancestry.

Zelandites n. gen.

This genus is suggested for some ammonites that could not be placed in any of the genera of which detailed descriptions and literature were available. The ornamentation is so similar to that of several species of Gaudryceras that it seems necessary to place it close to that genus. The narrow and high whorl has the general form of that of Gaudryceras varuna, which has been recorded from Quiriquina by Steinmann as well as from its original locality. On the other hand, the suture-line, so far as its general appearance is concerned, and especially in the nature of the first lateral lobe, is so peculiar that it becomes doubtful if the species should be included in the Lytoceratidae. However, in the absence of specimens of allied species for comparison, and of more extensive literature, it has been placed here provisionally.

Zelandites kaiparaensis n. sp. (Plate 20, figs. 9, 9a; Plate 31, figs. 1, 2.)



Ammonites varuna Forbes (1, p. 107, pl. 8, fig. 5).


Ammonites varuna Forbes, in Stol. (11, p. 111, pl. 58, fig. 1).


Desmoceras kawanoi Jimbo (22, p. 28, pl. 1, fig. 7 a, b).


Lytoceras (Gaudryceras) odiense Koss. (28, p. 129, pl. 18, fig. 1 a-c).


Lytoceras (Gaudryceras) varuna Forbes, in Koss. (28, p. 130, pl. 16, fig. 4 a-b: pl. 17, fig. 8).


Lytoceras varuna Forbes, in Steinmann (26, p. 84, Taf. 5, fig. 2 a, b; text-fig. 7).


Gaudryceras kawanoi Jimbo, in Yabe (37, p. 41).


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A. B. C. D.
Diameter 25 100 18 100 23 100 26 100
Height of last whorl 12 48 8 44 12 53 13.5 52
Width of last whorl 7.45 30 6.5 35 7.5 33 7 27
Umbilicus 6.5 26 5 28 4.5 20 5.2 20

A, B, Zelandites kaiparaensis n. sp., Bull's Point, Kaipara Harbour, N.Z.; C, Lytoceras (Gaudryceras) varuna Forbes, in Koss. (28, p. 161); D, Lytoceras varuna Forbes, in Steinmann (26, p. 84).

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The dimensions show a fairly close resemblance to Gaudryceras varuna, though the whorl of the New Zealand species is not so high and the umbilicus is a good deal wider.

Shell small, discoidal, with an involution of about two-thirds. The greatest width is just above umbilicus, to which the slope is steep, though it rounds off gradually at top of wall; thence the flanks slope gradually to periphery, which is narrow and sharply rounded. Body-chamber has the length of two-thirds of a revolution. Surface of inner whorls covered with fine hair-like ribs, at first strongly inclined forwards, but at top of umbilical slope they become straight and pass straight over periphery. Body-chamber has the same ornamentation as inner whorls. Nine deep constrictions in a whorl. They are at first radial in direction, but soon take a strong bend forward, and near periphery become radial again, passing over it with a barely noticeable backward curve. These constrictions are visible on the shell and have no ribs before or behind.

Suture-line peculiar, and the absence of much literature makes it hard to classify the species. The median saddle is sharp and has a few jags on side. External saddle high and bifid, with a much larger development on external than on internal side. The two lateral saddles are also bifid, but are much smaller, and seven auxiliary saddles can be distinguished. All divisions of the saddles have deep and sharp jags, which give rather a distinct appearance. External lobe about as deep as first lateral lobe, and rather wide. First lateral lobe wide, and the small secondary saddle in the middle is much less conspicuous and important than in Gaudryceras. There are larger projections on either side of it, and of these the internal one is by far the largest element in the whole. This is similar to the same lobe in Diplomoceras cylindraceum Defr. (47A, fig. 1259; 45, Taf. 45, fig. 47) and to that in D. wakanene, figured in this paper (Plate 19, fig. 3), but it is quite different from that in the four Indian species of Anisoceras Pictet = Diplomoceras Hyatt figured by Kossmat (28, Taf. 19). The other lobes present no unusual features. Auxiliary lobes and saddles highly inclined, and form a deep umbilical lobe. Internal portion of suture-line not much different from that of G. semileve or G. propemite, or of G. varicostatum van Hoepen, and approaches that of G. varagurense as drawn by Kossmat. In the three first-mentioned at least it may be said that there is a single wide and rapidly thinning saddle, which is high, while smaller saddles are represented by projections on its outer side. This is certainly distinct from the single thin and high saddle of G. subsacya and G. vertebratum as drawn by Kossmat.

The form of this species shows affinities with that of G. varuna and G. kawanoi, while the ornamentation is similar to the general characters of that of Gaudryceras.

Four specimens, all of them from Bull's Point, on the Kaipara Harbour. A specimen of a closely allied species is in the collection of the New Zealand Geological Survey. It was found by McKay at Awanui, near East Cape, in Cook County. It differs from the Kaipara form in ornamentation, as it is almost destitute of ribs, but in form and suture-line it differs but little.

Tetragonites Kossmat, 1895.

Whorls strongly involute and rapidly increasing. Cross-section trapezoidal, the broadest part near umbilicus: flanks and periphery flattened. Fine growth-lines only, and usually periodic growth forms bolsters on the shell and deep constrictions on the cast. Siphuncular saddle lance-shaped; external lobe same depth as first lateral lobe, which has a median secondary saddle. Several auxiliary saddles. Internal portion of suture-line may have

– 149 –

as many as four saddles, which are bifid like saddles of external portion. The genus begins with T. duvalianum in the Neocomian, and extends in a single Indian species into the Senonian.

The above is a brief summary of the generic diagnosis given by Kossmat (28, p. 131).

Tetragonites epigonus Kossmat. (Plate 21, fig, 10; Plate 29, figs, 6, 7.)



Ammonites timotheanum Mayor, in Stol. (11, p. 146, pl. 73, fig. 5).


Lytoceras (Tetragonites) epigonum Koss. (28, p. 135, Taf. 17, figs. 4 a, b, c, 5 a, b).


Lytoceras (Tetragonites) epigonum Koss., in Paulcke (42A, p. 8).


Lytoceras (Tetragonites) epigonum Koss., in Kilian and Reboul (46, p. 14).


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A. B. C. D.
Diameter 38 100 59 100 23 100 28 100
Height of last whorl 20 53 26 44 10 43 11 39
Width of last whorl 22 58 27 46 11 44 12 43
Umbilicus 8 21 15 25 7 30

A, Tetragonites epigonus Koss., Bull's Point, Kaipara Harbour, N.Z.; B, C, Lytoceras (Tetragonites) epigonum Koss. (28, p. 135); D, Lytoceras (Tetragonites) epigonum Koss., in Paulcke (42A, p. 8).

Shell discodial but whorls increasing rapidly in size. Involution about one-half, and umbilicus small but deep. Whorls wider than high. Umbilical wall almost vertical but soon curving rapidly into flank, which is gently rounded and passes without any sudden angle into periphery. Thickest part of whorl just above umbilicus. Surface appears smooth and polished, but examination shows it traversed by fine lines which arise radially on the steep slope of umbilicus, then bend forward, but soon change again and cross periphery with a slight curve backward. There are deep constrictions, most pronounced on middle of flanks but soon fade away towards periphery and in umbilicus; there are five in a revolution, and they follow the same general course as the striations.

Suture-line very similar to that of Pseudophyllites indra, and I fail to find any important differences in the figures given by Kossmat for the two species and also in the specimens found in New Zealand. The internal portion of suture-line shows a similarity to the figure of Kossmat, but the saddles are less divided and the saddle nearest to umbilical surface is relatively shorter.

One specimen at Bull's Point, and one at Batley.

Tetragonites latus n. sp. (Plate 20, figs. 6, 6a; Plate 32, figs. 1, 2.)



Ammonites timotheanus Pictet and Roux; also the same species in. Stoliczka, Schmidt, Whiteaves. Anderson, and Kossmat.


Tetragonites glabrus Jimbo, in Yabe (37, p. 43, pl. 7, figs. 2, 5).


Tetragonites kingianum var. involutior Paulcke (42A, p. 9, Taf. 17, figs. 3 a, b, 4 a, b).


Tetragonites superstes van Hoepen (52, p. 11, pl. 2, figs. 17–20, text-fig. 6).


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A. B. C. D. E. F.
Diameter 19 100 13 100 13.5 100 80 100 32.4 100 35 100
Height of last whorl 9.5 50 7 54 7.3 54 41 50 14.2 44 17 48
Width of last whorl 11 58 7.7 59 8 59 45 56 15.5 48 19 54
Umbilicus 3 16 2 16 2 15 16 20 8.3 26 8 21

A, B, Tetragonites latus, Bull's Point, Kaipara Harbour, N.Z.; C, Tetragonites latus, Batley, Kaipara Harbour, N.Z.; D, Tetragonites timotheanus Pictet, in Koss. (28, p. 133, Taf. 17, figs. 11, 13 a, b); E, Tetragonites superstes van Hoepen (52, p. 11, pl. 2, figs. 17–20, text-fig. 6); F, Tetragonites glabrus Jimbo, in Yabe (37, p. 43, pl. 7, figs. 2, 5).

– 150 –

Shell small, discoidal, somewhat inflated and involute. Four whorls only in a radius of 10 mm. Whorls rise very steeply from umbilicus, where there is a distinct rounded angle, and afterwards a uniform curve to periphery, which is slightly flattened. Whorls always distinctly wider than high, and increasing rapidly, thus forming a deep and narrow umbilicus.

Ornamentation: The surface is practically smooth, but longitudinal grooves can be seen distinctly. There is one on each side of the median line, about 1 ½ mm. distant from it in the largest specimen. There is another, less distinct, where the whorl curves from the flat periphery to the flank. These spiral lines are not distinct in all specimens.

Surface of shell appears to be bright and shining, but is actually marked by very fine indistinct and irregular striations, between which low rounded ribs can be distinguished. These bend forward slightly at first, then cross periphery without change. Spiral lines of a similar nature to those seen in this species have been recorded in T. superstes by van Hoepen, and in the related genus of Pseudophyllites in the species garuda by Stoliczka which is included in the species indra by Kossmat. No periodic grooves whatever can be seen on any of my specimens. The body-chamber shows for half a revolution in one specimen, but it is not complete. Suture-line shows that the first lateral lobe is unusually wide. Divisions of saddles and lobes very simple. Second lateral saddle is already on slope of umbilicus. Three auxiliary lobes and saddles. Internal portion of suture-line has a deep antisiphonal lobe and two saddles nearly equal in height, but projections on outer side of second saddle may represent partially suppressed saddles.

This species differs from the widely-distributed T. timotheanus in the entire absence of constrictions, and from this and apparently from all other closely related species in the great width of whorl and the narrowness of umbilicus. It is perhaps nearer to T. superstes van Hoepen than to any other. T. kingianum var. involutior Paulcke is quite close to this species, but has a wider umbilicus.

Several specimens have been obtained from Batley, Bull's Point, Whangaroa, and perhaps Tetragonites sp. juv. Spath (53, p. 300) from Selwyn Rapids is the same species.

Tetragonites simplex n. sp. (Plate 20, figs. 11, 11a; Plate 32, figs. 3, 4.)



Ammonites cala Forbes (1, p. 104, pl. 8, fig. 4).


Ammonites cala Forbes, in Stol. (11, p. 153, pl. 75, fig. 4).


Lytoceras (Tetragonites) cala Forbes, in Koss. (28, p. 136, 163, Taf. 17, figs. 12 a-d).


Tetragonites nuperus van Hoepen (52, p. 13, pl. 3, figs. 3, 4, text-fig. 8).


Tetragonites nuperus van Hoepen, in Spath (54, p. 119).


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A. B. C. D. E.
Diameter 19 100 21 100 20 100 80.5 100 37.6 100
Height of last whorl 7 31 7.5 35 8 40 25 31 14 37
Width of last whorl 7.2 37 8 42 8.4 42 29 37 15.4 41
Umbilicus 7.6 40 8 42 7 35 38 47 15.5 41

A, B, C, Tetragonites simplex n. sp., Batley, Kaipara Harbour, N.Z.; D, Tetragonites cala Forbes, in Koss. (28, p. 163); E, Tetragonites nuperus van Hoepen (52, p. 13).

Shell small, discoidal, slightly involute, about one-quarter and therefore with a wide umbilicus. Five whorls in a 10 mm. radius. Whorls rise steeply from umbilicus and curve sharply, almost with an angle, to the evenly-rounded flanks, and pass with another sharp curve or angle to the much-flattened periphery. The section of a whorl is not truly

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angular, though the curves are rapid. Surface of shell quite smooth and polished. There are three conspicuous constrictions on the cast in each whorl, but they cannot be distinguished on the shelly matter. They commence on edge of umbilicus and bend sharply forward, becoming much less distinct towards periphery, which they cross with a low forward curve.

Suture-line very simple. Median saddle rounded. External saddle with three simply rounded projections which are less distinct on first lateral saddle and just indicated on second. The five auxiliary saddles are quite simple. First lateral lobe has one simple secondary saddle.

This species is close to Tetragonites nuperus van Hoepen, but the suture-line is a good deal simpler, and in this respect it resembles G. rouvillei de Grossouvre (19, p. 288, pl. 37, fig. 10), though it seems to be quite different from the larger form, fig. 7. It differs from T. cala Forbes in the less angular whorl-section and in the simpler suture-line. Tetragonites kingianum var. involutior is much less inflated, and has a more complex suture-line. In suture-line and in general aspect this species shows a great resemblance to G. bucculentum Pervinquière (Mem. Geol. Soc. de France, 17, Fasc. 2, 3, 1910).

This is not an uncommon species at Bull's Point and Batley; a specimen has also been found at Whangaroa.

Tetragonites margaritatus n. sp. (Plate 20, fig. 5; Plate 30, figs. 5, 6.)



Tetragonites popetensis Yabe (37, p. 48, pl. 7, figs. 4 a, b, 6).


Lytoceras (Tetragonites) kingianum var. involutior Paulcke (42A, p. 8, pl. 17, figs. 3 a, b, 4 a, b).


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A. B. C. D. E.
Diameter 15.5 100 12 100 11.5 100 34 100 20.5 100
Height of last whorl 7 45 4.7 39 5 43 14.5 43 9 43
Width of last whorl 8 51 5 42 5.5 49 15 44 9 43
Umbilicus 5 32 4 35 4 36 10.5 30 5.5 26

A, B, Tetragonites margaritatus, Batley, Kaipara Harbour, N.Z.; C, Tetragonites margaritatus, Bull's Point, Kaipara Harbour, N.Z.; D, E, Tetragonites popetensis Yabe (37, p. 48, pl. 7, figs. 3 a, b, 4 a, b).

A very small species, of which there are five examples. Shell is discoid, with an involution of about one-half. Wall of umbilicus steep, but passes quickly into circular curve of flanks and periphery, though the latter is distinctly flattened. Whorls increasing rather rapidly. Umbilicus deep, its slope interrupted by curved surface of whorls, of which there are five in a radius of 8 mm. In one specimen body-chamber is three-quarters of a revolution, but it is not complete. Surface of shell perfectly smooth, and in the type specimen pearly, though a few very indistinct growth-lines can be distinguished. There are occasional deep constrictions which bend strongly forward from bottom of umbilicus until they reach halfway across the flanks, from which point they extend straight to periphery and cross it without change. No more than two can be seen in a revolution. Suture-line with six saddles, well divided considering the small size of the shells. Saddles high and narrow, and there are three auxiliary saddles. On the whole, the suture-line is quite characteristic of the smaller species of Tetragonites.

A number of specimens were found at Batley, and two at Bull's Point.

This species closely resembles T. popetensis, but the umbilicus is rather wider than in that species. The smoothness of the surface of the present

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species is another distinction. Umbilicus wider and whorls more projecting than in Paulcke's species T. kingianum var. involutior from South Patagonia.

Pseudophyllites Kossmat, 1895.

Kossmat gives no definite statement of the characteristics of the genus, but from the description of the only species of it that he recognizes the following statement may be taken as embodying the essential features:—

Strong involution, rapid increase especially in the later stages of growth, and very small umbilicus. The whole surface covered with very fine hair-like lines which bend backwards at periphery; there are also broad bands which may be 5 mm. apart, but no constrictions. In the young the suture-lines are phylloid, and the external and first lateral saddles are trifid. Kossmat states that in the larger examples the inner division lobe in the trifid external saddle is the deeper, and the outer division lobe of the first lateral saddle is the deeper. The unsymmetrical nature of the external and first lateral saddles distinguishes Pseudophyllites from Gaudryceras and Tetragonites.

Pseudophyllites indra Forbes. (Plate 20, fig. 1; Plate 29, figs. 3, 4, 5.)


  • Ammonites indra Forbes (1, p. 105, pl. 11, fig. 7).

  • Ammonites garuda Forbes (1, p. 102, pl. 7, fig. 1).

  • Ammnoites indra Forbes, in Stol. (11, p. 112, pl. 58, fig. 2).

  • Ammonites garuda Forbes, in Stol. (11, p. 149, pl. 74, fig. 5).

  • Ammonites indra Forbes, in Whiteaves (8, p. 105, pl. 13, fig. 2).

  • Lytoceras (Pseudophyllites) indra Forbes, in Koss. (28, p. 137, pl. 16, figs. 6, 9; pl. 17, figs. 6, 7; pl. 18, fig. 3).

  • Phylloceras indra Forbes, in Whiteaves (27, p. 129).

  • Pseudophyllites indra Forbes, in Whiteaves (34, p. 331).

  • Lytoceras indra Forbes, in Boule and Thevenin (42, p. 44, pl. 1, fig 1).

  • Pseudophyllites indra Forbes, in Woods (40, p. 334, pl. 41, fig. 6).

  • Pseudophyllites indra Forbes, in Kilian and Reboul (46, p. 14).

  • ? Tetragonites teres van Hoepen (51, p. 144, pl. 25, figs. 1, 2).

  • Tetragonites (?) sp. indef. Spath. (59, p. 42, pl. 7, fig. 3).

  • ? Tetragonites virgulatus van Hoepen (52, p. 11, pl. 3, fig. 1, 2; text-fig. 7).


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A. B. C. D.
Diameter 34 100 28 100 24 100 21 100
Height of last whorl 16 47 14 50 12 50 10 48
Width of last whorl 17.5 51 16 57 13 54 11 52
Umbilicus 8 24 6 21 5.5 23 5 24
E. F. G. H. K.
Diameter 21 100 113 100 20 100 27 100 33 5 100
Height of last whorl 10 48 61 54 9.5 47 12 44 16.7 50
Width of last whorl 12 57 51 45 9 45 12 44 17.1 51
Umbilicus 5 24 18 15 4.8 24 6 22 7.3 22

A, B, Pseudophyllites indra Forbes, Batley, Kaipara Harbour, N.Z.; C, D, E, Pseudophyllites indra Forbes, Bull's Point, Kaipara Harbour, N.Z.; F, G, H, Pseudophyllites indra Forbes, in Kossmat (28, p. 164); K, Tetragonites virgulatus van Hoepen (52, p. 13).

Shell discoidal, but whorls rapidly increasing with an involution of three-quarters. Whorls slightly wider than high in all measured specimens. In this respect the New Zealand specimens differ slightly from the Indian shells that were measured by Kossmat. Umbilical wall steep, and passing into the slightly flattened flank with a broad curve. There is, however, no suggestion of the angle at top of umbilical wall that is characteristic of Tetragonites epigonus Koss. Umbilicus narrow and deep showing four

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whorls when diameter is 12 mm. Surface of shell apparently smooth and polished, but when viewed with a lens it is seen to be much striated. The striae start nearly radially at bottom of umbilicus, but curve strongly forward, the curve beginning to decrease at top of umbilical wall, and at top of flank it passes into a faint backward curve precisely as shown by Kossmat (28, Taf. 18, fig. 3). The larger rounded bands to which Kossmat refers are also clearly visible, but they are not prominent, and are somewhat variable in importance. There are no constrictions.

Suture-line is much divided—the median saddle is rather long and has several jags on the side. External saddle unsymmetrically bifid, with the interior portion much smaller than exterior. First lateral saddle also bifid, but exterior portion is here the smaller, and the larger interior portion is divided by a secondary lobe almost as deeply as the main division of saddle, thus giving almost the appearance of a trifid form. Second lateral saddle small, and only three auxiliary saddles can be seen (Kossmat shows four in the Indian examples). External lobe as deep as first lateral, which is very wide and has a prominent secondary saddle in the middle. Auxiliary elements are moderately inclined, and umbilical lobe is not so deep as in most species of Gaudryceras. Internal portion of suture-line can be seen in one of the specimens, but it is a little eroded. Two high saddles not much divided, and the one near umbilical surface is considerably shorter than the other. It is therefore not very different from that of Tetragonites simplex, and of T. epigonum as drawn by Kossmat.

The form and ornamentation are so similar to those of the typical Indian examples which I had the opportunity of examining at the British Museum that I do not feel justified in separating it, notwithstanding the slight but constant differences in dimensions. A single example of P. indra was recorded by Kilian and Reboul in the collection from Seymour Island, and it has also been recorded from South Africa (Pondoland), and from the Nanaimo formation of Vancouver. In India it occurs in the Valudayur and Trigonoarca beds of the highest Senonian. It is considered by de Grossouvre that this species is the same as G. colloti from the Upper Campanian of the south-east of France; but Kossmat points out that, though the ornamentation and apparently the suture-line are closely similar, the dimensions of the European form are different, and he therefore retains this species as distinct. Kossmat also points out that it is closely related to G. postremus Redtenbacher from the Gosau. Spath classes van Hoepen's two species Tetragonites teres and T. virgulatus from Pondoland with this species.

Numerous examples have been found, especially at Bull's Point, also at Batley, and one at Whangaroa. Ferrar has found examples at Te Opu, in the Kaipara Harbour, as well as at Matakohe, on the shores of the harbour six miles north of Bull's Point. A specimen was sent to Professor Kilian, and was identified by him as Pseudophyllites indra.

Pseudophyllites whangaroaensis n. sp. (Plate 20, fig. 2; Plate 21, fig. 11; Plate 32, figs. 5, 6.)

Compare with the species quoted under P. indra Forbes.


Diameter 42 100
Height of last whorl 21 50
Width of last whorl 22 53
Umbilicus 11 26

A, Pseudophyllites whangaroaensis n. sp., Whangaroa Harbour, N.Z.

– 154 –

Shell of moderate size, with whorls rapidly increasing in size. Involution almost one-half, with umbilicus narrow and deep. Whorls rise steeply from umbilicus and curve quickly into an inclined but somewhat flattened flank, which in turn leads to a rather flattened periphery. Four whorls at a diameter of 42 mm., but depth of umbilicus makes it almost impossible to expose them. The surface shows fine hair-like ribs, which at the bottom of umbilicus are almost radial; but they bend strongly forward and on flanks become more irregular and discontinuous. They are crossed by a series of low rounded spiral elevations, which are intersected by similar but more irregular radial elevations. These give rise to a check pattern, which is best shown near periphery. With a lens the fine hair-like striae can be seen crossing this checkwork, but they are too fine to show in a photograph. No constrictions can be seen.

The suture-line, which is incompletely shown in the single specimen, is similar to that of P. indra, but is perhaps rather less deeply dissected. The elements of first lateral lobe are somewhat less developed, but the general nature of saddles and lobes is the same.

This species is rendered distinct by its ornamentation, which makes such a striking checkwork pattern on the shell. Kossmat, however, says of Pseudophyllites indra that lightly impressed spiral lines are almost always present, but not so markedly as Stoliczka represents them. I can find no mention of this in Stoliczka. * The statement seems to refer to fine lines rather than the rounded swellings that are so striking in this species. None of the specimens of the species here called Pseudophyllites indra Forbes shows any sign of fine spiral lines or of the sculpture described for this species.

A single specimen has been found at Whangaroa.

Baculites Lamarck.

Straight forms, circular or oval in cross-section, smooth or ornamented.

Baculites rectus n. sp. (Plate 19, fig.]; Plate 32, figs. 9, 10.)



Baculites chicoensis Trask, Proc. Cal. Acad. Nat. Sci., p. 85, p]. 2, fig. 2.


Baculites chicoensis Gabb (3, pl. 14, fig. 27b; pl. 17, figs. 27, 27a).


Baculites cf. anceps Paulcke (42A, p. 10, pl. 16, figs. 6, 6a).


Baculites sp. cf. vagina Forbes, in Woods (57, p. 36, pl. 20, figs. 5 a-d).

The largest fragment measures 135 mm. in length, and the diameters of its cross-section are 30 mm. and 22 mm. at anterior end, and 21 mm. and 17 mm. at posterior end. All except the anterior 38 mm. is septate.

Ornamentation: Surface of shell quite smooth except for some obscure and indefinite growth-lines, and there are no signs of tubercles. Cross-section oval, and the siphuncle is situated at the more pointed end, which is not carinate. Chambers unusually long. First lateral lobe has a prominent projection which makes a distinct bifid division of the lobe. In second lateral lobe are three projections of practically the same size and shape, and the two divisions on either side of median projection are quite similar in development. Saddles deeply bifid. The only species that

[Footnote] * I find this is mentioned under the species Ps. garuda, which Kossmat regards as a synonoym of Ps. indra (11, p. 149, pl. 74, figs. 5, 5a).

– 155 –

approaches closely to the New Zealand species in characters of suture-line is Baculites chicoensis Trask from California. Baculites ovatus Say from North America, where it extends right through the Fort Pierre beds (Senonian) is also somewhat similar. The smooth surface as well as the suture-line distinguishes this species from all recorded from South Africa.

The species that occurs at Quiriquina, identified as B. vagina by Stein-mann (26 p. 89, Taf. 6, figs. a-e, also text-figs. 8–12), is quite distinct from the New Zealand species both in external sculpture and in the form of the suture-line. The species recorded by Paulcke from South Patagonia (42A, p. 11, pl. 16, figs. 6, 6a) has much more sculpture, and lacks the most distinctive features of the suture-line. Kossmat states that a specimen from India is very similar to an unnamed specimen from California figured by Gabb (3, pl. 17, fig. 27), and names it Baculites vagina var. simplex. It, however, is quite different from the present species.

It thus appears that the present species, B. reclus, is more closely allied to B. chicoensis Trask than to any species of which descriptions are available. This species occurs in Queen Charlotte Sound of British Columbia, as well as in California, where in the type locality it is found only in the, upper division of the Chico, though it has been recorded also in the Lower Chico of San Diego and Silverado (33, p. 27).

Baculites cf. vagina Woods (57, p. 36, pl. 20, fig. 5), which was found at the Amuri Point, in the South Island of New Zealand, almost certainly belongs to this species.

A large number of specimens has been found, though most of them are small fragments. They are most frequent at Bull's Point, though not uncommon at Batley and Whangaroa.

Diplomoceras Hyatt.

The Indian species Diplomoceras rugatum Forbes, to which the following New Zealand species is closely allied, was originally included by Forbes (1, p. 117) in the genus Hamites, but Stoliczka in 1885 used the generic name Amsoceras for this and a large number of related species. Kossmat in 1895 employed the generic title Hamites (Anisoceras). Hyatt in 1900 gave the generic name Diplomoceras to a number of the Upper Cretaceous uncoiled Ammonites, using as the genotype D. cylindraceum Defr. I am unable to obtain a full description of this genotype, but figures are given in Zittel (30A, p. 571, fig. 1187), and the same appear in the next edition, where the Ammonite section is edited by J. P. Smith (47A, p. 654, fig. 1259). The genus Diplomoceras is provisionally adopted for the single New Zealand species, though the suture-line has a very deep external lobe.

Diplomoceras wakanene n. sp. (Plate 19, fig. 3; Plate 31, figs. 8, 9.)


  • Diplomoceras indicum Forbes, in Koss. (28, p. 145, Taf. 19, figs. 4 a-c).

  • Diplomoceras subcompressum Forbes, in Koss. (28, p. 145, Taf. 19, figs. 10 a-b, 11 a,b, 12).

  • Diplomoceras rugatum Forbes, in Koss. (28, p. 146, pl. 19, figs 7 a, b, 8 a, b, 9): on the pages quoted full references will be found to the work of Forbes and Stoliczka on these species.

Dimensions: Height of whorl, 11.2 mm.; width, 9 mm.: also, height, 7 mm.; width, 5.5 mm.

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Shell nearly elliptical in cross-section; found in widely-curved fragments without any straight portions, which are apparently derived from an irregular spiral shell. Numerous rounded ribs of moderate size extend right round whorl, but they are most prominent on middle of flank and decrease greatly in size as they approach ventral and dorsal surface. No interstitial ribs and no unequal spacing to suggest that constrictions occur. No tubercles. Suture-line generally resembles that of D. rugatum more closely than that of any other Indian species, which are mainly distinguished by the size of second lateral saddles, whose greatest development is in D. rugatum. In the present species second lateral saddles are still larger, and are separated by a lobe as deep as first lateral lobe. External lobe also of much the same depth. First lateral lobe has the peculiarity of showing two prominent projections, of which the internal is the more pronounced. This feature is seen in D. cylindraceum Defr. (47A, fig. 1259; 45, Taf. 45, fig. 47), but not in any of the Indian species, D. indicum D. rugatum, D. subcompressum, as figured by Kossmat (28, Taf, 19). In the present species, however, both the external and the antisiphonal lobes are of unusual width and depth. First lateral lobe has a very similar structure in Zelandites kaiparaensis (Plate 31, fig. 2).

As indicated, this species seems to be closest to D. rugatum Forbes from the Valudayur beds of Pondicherry, but is distinguished by the rounded form of the ribs, absence of constrictions, as well as the form of suture-line. Kossmat states that D. rugatum is very similar to D. cylindraceum Defr. from the highest Senonian of Europe.

A single specimen has been found on the south-west side of Bull's Point, on the Kaipara Harbour, N.Z.

The specific name is that of a well-known Maori chief.

Oxybeloceras Hyatt.

Oxybeloceras sp. (Plate 15.)

Shell nearly straight, with a maximum width of 35 mm., thickness of 16 mm., and length of 292 mm. Surface marked with numerous ribs 4 mm. apart and strongly inclined. On each rib is a small bead-like tubercle on the side of the specimen that is preserved, and there is an indication of another small tubercle near periphery, but the condition of the specimen does not allow this to be seen distinctly. Unfortunately no indication of any suture-line is to be seen, and the only specimen seems to have been a body-chamber only. Costation quite distinct from Aniso-ceras sp. in Woods (57, p. 35, pl. 20, figs. 3, 4). The species of Anisoceras notabile and A. obstrictum figured by Killian and Reboul (46, p. 15, 16) have not inclined costation, and apparently no tubercles.

The present specimen is unfortunately far too imperfect for any specific description. There is a single specimen from the Mangamuka River, Hokianga, in the Auckland Museum. I am indebted to Dr. F. L. Spath for kindly informing me that this species probably belongs to the genus Oxybeloceras of Hyatt. I have not been able to find any statement of the characteristics of this genus, but Spath refers a Pondoland species to it (59, p. 50.)

Ptychoceras d'Orbigny.

Uncoiled forms, for the most part straight, but at intervals bent through 180°

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Ptychoceras zelandicum n. sp. (Plate 19, fig. 2; Plate 32, figs. 11, 12.)



Ptychoceras sipho Forbes (1, p. 118, pl. 11, fig. 5).


Ptychoceras sipho Stol. (11, p. 194, pl. 90, figs. 5–9).


Ptychoceras sipho Koss. (28, p. 150).

Straight fragments only have been found, but, as the matrix is extremely hard and the specimens fracture easily, it is possible that the species has actually a hooked form, as it usual in this genus. Nearly all the fragments are small, 1–5 mm. in diameter, but there is one piece of a living-chamber 72 mm. in length and 16 mm. in diameter. Cross-section is almost circular, the taper is hardly noticeable, and thus the specimens have a cylindrical appearance. There are deep constrictions at intervals, strongly inclined to the axis. Anterior apex of constriction is at siphuncular side, and in quite small specimens is 4 mm. in advance of posterior portion. Surface of shell quite smooth, or feebly marked with indistinct lines following the general direction of constrictions. Suture-lines simple and symmetrical. Six prominent saddles and six lobes. Siphuncle situated at top of a small saddle in a deep lobe and the antisiphonal line is in a similar deep lobe, which divides the dorsal saddle into two divisions each as large as the other saddles. The six saddles are almost equal in size, and are broader than the lobes. Each saddle is bifid, and the two halves are rounded, and have a slight indentation in each half. The suture on the whole is similar to that of Ptychoceras sipho figured by Forbes (which figure is in the opinion of Kossmat much superior to Stoliczka's). Little distinction can be made between the different saddles in size or form, and the lobes also are almost indistinguishable from one another. The only distinction that can be made is that the external lobe is slightly wider than any of the other, and the median saddle is rather wider than the small saddles at the base of the other lobes.

This species is distinguished from Ptychoceras sipho by its simpler suture-line and by the apparently unhooked form. Ptychoceras sipho has not yet, so far as can be seen in the literature consulted, been compared with any other form. That species occurs in the Valudayur beds of Pondicherry (Upper Senonian).

A large number of fragments of this species has been obtained. It is very frequently found at Whangaroa, and specimens have also been found at Bull's Point and Batley.

Acanthoceras Neumayr.

Kossmat, following Zittel, describes this genus as follows (28, p. 108): “Ribs simple or dichotomously split, straight, thickened in the outer portion; lateral and umbilical tubercles. External portion broad, with a median row of tubercles.“

Group of A. rotomagense: Whorls broad, the ribs usually alternately long and short, with a straight radial arrangement, and seven rows of tubercles—siphonal, umbilical, and two marginal on each side, though either or both of the last may be obliterated. The suture-line has only two large saddles. The bifid external saddle is usually club-shaped. The second lateral saddle is on the umbilical slope and is not much larger than the auxiliaries, of which there are always a few present. The first lateral lobe is usually forked and is about as long as the external lobe. (A summarized translation).

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Acanthoceras ultimum n. sp. (Plate 25, figs. 1, 1a; Plate 34, figs. 1, 2.)



Acanthoceras newboldi Koss., typical form (28, p. 5, pl. 12, fgs. 2, 3) = A. rotomagense var. asiatica Jimbo (22, p. 177, Taf. 4, fig. 1).


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A. B. C. D.
Diameter 94 100 99 100 220 100 149 100
Height of last whorl 37 39 42 42 81 37 60 40
Width of last whorl 40 42 53 53 90 41 80 54
Umbilicus 27 29 32 32 76 35 50 34

A, Acanthoceras ultimum n. sp., Bull's Point, Kaipara Harbour, N.Z.; B, Acanthoceras newboldi var. spinosa Koss. (28, p. 7); C, Acanthoceras newboldi form. typ. Koss. (28, p. 5); D, Acanthoceras rotomagense var. asiatica Jimbo (22, p. 31) = A. newboldi, form. typ. Koss. (28, p. 5).

The table shows that in form this species is quite close to the typical A. newboldi Koss.

Shell of moderate size, discoid, but rather inflated; involution slightly less than one-half. Whorls wider than high. Umbilical wall steep, with widest part of shell just above top, and afterwards the slope towards periphery is gentle until the rounded angle on its edge. Periphery nearly flat. Whorls increasing rapidly in size. Ornamentation striking, but the only specimens are imperfect and fail to display it fully. Four constrictions on last whorl; they are narrow and extend right over periphery without a curve from bottom of umbilicus. Ribs on the two sides of constrictions have small tubercles. Nine tubercles on edge of umbilicus in last whorl. Most of these are large, vertical, and rounded, but a few are almost lamellar, following direction of ribs. Ribs large and conspicuous, rounded when shell is present but sharp and irregular when there is a cast. There are about nineteen on a half-revolution. Some start close to base of a tubercle, but there are also a few interstitial ribs. Half-way between umbilical edge and angle of periphery they rise into a low projection, and at the angle itself they form large projecting sharp tubercles which are much steeper behind than before. On siphuncular line are other lower tubercles which appear to be steeper before than behind. Ribs much flattened between edge of periphery and siphuncle. In the small inner whorls umbilical tubercles alone are present.

Suture-line is relatively simple, and the various arms of the saddles are not deeply intersected. Median saddle only slightly ornamented. Second lateral saddle rather distorted by development of the large umbilical tubercle. Auxiliary saddles are three in number, rapidly decreasing in size, but are not inclined so as to form a deep umbilical lobe. Internal portion of suture-line closely approaches that of A. newboldi, illustrated by Kossmat (28, pl. 3, fig. 2), though the smaller third saddle is less distinct. First lateral lobe indistinctly forked and rather shorter than external lobe. The suture-line generally much more finely divided than that of A. newbold and of the other Indian species figured by Kossmat.

This species is rather more tuberculate than Jimbo's form, which comes from the Ikushumbets. The dimensions show that it comes fairly close to the Indian species.

Only two specimens have been found, one of which is in extremely bad condition. Both of them were found at Bull's Point.

Kossmaticeras (de Grossouvre).

A number of species of ammonites that had been classed in the group Ligati by Stoliczka in 1885 (11) were in 1895 placed in the genus Holcodiscus

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by Kossmat (28). However, de Grossouvre pointed out in 1901 that these Upper Cretaceous forms were not cophyletic with the species of Holcodiscus from the Lower Cretaceous, and he then proposed the name Kossmaticeras for them. This name has subsequently been adopted by other authors.

Kilian and Reboul pointed out in 1909 (46) that great differences were displayed in the ornamentation of the species that were referred to the genus Kossmaticeras, and for the groups of these that appeared from their ornamentation to be related he suggested the following subgeneric names: Gunnarites, Madrasites, Grossouvrites, Jacobites, Seymourites, and Grahamites. This subgeneric classification has been adopted by other authors, and has been followed here, though the subgenera are treated as genera. It is also suggested that Madrasites bhavani var. densicostata Kilian and Reboul should be placed in a new genus Maorites, closely allied to the genus Papapuzosia.

Gunnarites Kilian and Reboul, 1909.

Genotype: Gunnarites antarcticus Stuart Weller.

The group of Gunnarites was established by Kilian and Reboul (46, p. 26) in the following words: “Dans le groupe de Kossmaticeras antarcticum Stuart Weller les caractères se modifient par l'apparition de crénelures de plus en plus accentuées sur les côtes, et il finit par en rester, dans l'adulte, une ornamentation qui rapelle vraiment celle de certains Douvilleiceras. Ce sous-groupe, qui contient plusieurs espèces notamment K. kalika Stol., K. antarcticum Stuart Weller, peut-être désigné sous le nom de Gunnarites.” In such descriptions from other countries as are available to me I can find no mention of other species that can be referred to this genus. It is therefore of some interest, and it must be regarded, so far as our present knowledge is concerned, as having its headquarters at Seymour Island. Its representation in New Zealand is, however, considerable, and one species is recorded from India. Possibly there is a species in Madagascar identified by Boule, Thevenin, and Lemoine as K. theobaldianum Stol. Stoliczka did not draw the suture-line of G. kalika, and Kossmat apparently did not see any specimens of that species. Kilian and Reboul (46) merely say of G. antarcticus that the suture-lines are imperfectly preserved, and of G. nordenskjoldi that it closely resembles that of K. karapadense figured by Kossmat (28, p. 41, Taf. 8, fig. 4c). It thus happens that the suturelines in Plate 22 appear to be the first that have been drawn of species of this genus. It is well to make a comparison with those of Madrasites, of which there are abundant drawings by Kossmat, van Hoepen, Spath, and others, and as far as possible with those of other genera originally established by Kilian and Reboul as subgenera of Kossmaticeras. The specimen of G. inflatus from which the suture-line was taken has a whorlheight of 38 mm., that of G. antarcticus is 35 mm., and that of G. zelandicus 24 mm.

The median saddle has a considerable amount of frilling, especially near the top, but generally the suture-line is not finely divided. The external lobe has a strong projection in it which reaches nearly as high as the top of the median saddle. The external saddle is deeply bifid, with three well-developed arms on each side. The exterior half of the saddle is rather larger than the interior half. The first lateral lobe is a little deeper than the external lobe, and it is quite symmetrical. The first lateral saddle is not so deeply divided as the external saddle. The development of the second lateral saddle is much affected by the umbilical tubercle, over and around which its arms pass. In large specimens there may be

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six auxiliary saddles, gradually decreasing in size and strongly inclined. The second lateral lobe is not nearly so deep as the external or the first lateral lobe.

The internal portion of the suture-line shows first a large, broad, and much-divided saddle, which is separated by a highly-divided lobe from a second high but narrower saddle, which is succeeded by a deep antisiphonal lobe. It is thus very different from the internal portion of the suture-line of Madrasites theobaldianus Forbes, which is figured by Kossmat (28, pl. 7, fig. 5). This is the only drawing of the internal portion of a suture-line of any Kossmaticeras that I have been able to find. In this drawing there are three well-developed saddles and four auxiliary saddles in the internal suture-line. In general features the whole of the suture-line is not very different from that of Madrasites, though it is somewhat more frilled.

The suture-line of Grossouvrites—taking G. gemmatus as the genotype, as is done by Kilian and Reboul (46, p. 26)—is much more highly divided (46, p. 22, text-fig. 15). It has a great similarity to that of Pachydiscus, as remarked by other authors, and I am inclined to think that it should be placed in that genus. This species has been recorded from New Zealand by Trechmann (56).

One species of Gunnarites has already been recorded from New Zealand under the name Kossmaticeras zelandicum by Marshall (49, p. 444, pl. 33, fig. 2, text-fig. 2).

Gunnarites inflatus Kilian and Reboul. (Plate 22, figs. 1, 1a, 1b; Plate 40, figs. 1, 2.)



Kossmaticeras (Gunnarites) antarcticum var. inflatum Kilian and Reboul (46, p. 33, pl. 10, fg. 1; pl. 11, fg. 1; pl. 16, fg. 1).


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A. B.
Diameter 130 100 108 100
Height of last whorl 54 45 50 46
Width of last whorl 52 40 46 42
Diameter of umbilicus 38 29 31 29

In the specimen B the succeeding whorls measure H. 20–25, W. 21–25: H. 13, W. 15; H. 7.5, W. 9, respectively. The height is thus seen to increase more rapidly than the width.

A, Gunnarites inflatus, Batley, Kaipara Harbour, N.Z.; B, Gunnarites inflatus, Bull's Point, Kaipara Harbour, N.Z.

The shell is of moderate size, discoid in shape, with a deep umbilicus. Involution nearly one-half. Width of whorls slightly less than height. Wall of umbilicus steep but not perpendicular. A sharp curve at top of umbilical wall. Flanks almost flat and a low curve over periphery. Ornamentation distinct. There are twenty-five strong ribs in a half-revolution. High tubercles situated at top of umbilical wall; they are sharp, slightly twisted, and bent a little backward. From each of the tubercles two ribs arise. An interstitial rib generally arises between each pair of ribs a little distance from umbilical angle. No additional ribs arise higher on flanks. Five constrictions can be distinguished in a revolution. All the ribs are distinctly denticulated. They have a slight backward bend at top of flank, and afterwards bend a little forward at periphery. The first two and a half whorls have tubercles but no ribs.

Suture-line not well preserved so far as the external saddle is concerned, but in general is not highly divided. Median saddle well ornamented at

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top, and is rather more than half the height of external saddle. There is a strong process in external lobe. External saddle strongly bifid, and divisions are again nearly symmetrically bifid. First lateral lobe rather deeper than external lobe and quite symmetrical. First lateral saddle large and bifid. Second lateral lobe and second lateral saddle rather irregular because of umbilical tubercles situated there. Auxiliary saddles six in number, rapidly declining backwards, forming a deep umbilical lobe. In internal portion of suture-line is a deep antisiphuncular lobe. Two large much-divided saddles of nearly equal size between it and umbilical angle. These saddles are separated by a deep lobe which has several sharply-pointed processes at base. Outer side of exterior of the two saddles forms with auxiliary saddles a deep umbilical lobe. This internal portion of suture-line differs in a marked manner from that of Madrasites theobaldianus Stol. (28, pp. 36–143, pl. 7, fig. 5). This is the only figurer of the internal portion of a suture-line of Madrasites or Gunnarites that is to be found in accessible literature. This difference alone would justify the separation of Gunnarites from Madrasites, though there is at present no certainty that the same feature occurs in the other species of Gunnarites.

This species was originally placed by Kilian and Reboul as a variety of G. antarcticum Stuart Weller (46, p. 33). Professor Kilian kindly identified a specimen as identical with those obtained from Seymour Island. The specimen is in good condition, and was found by Miss Linley at Batley. It is embedded in a hard sandstone, and part of the suture-line cannot be seen distinctly. Another specimen has since been found at Bull's Point, and fragments probably of this species at Whangaroa. Kilian and Reboul record it from the Snow Hill beds, correlating them with the Trichinopoly beds of India, which they consider the equivalent of the Santonian.

Gunnarites zelandicus Marshall. (Plate 22, fig. 2; Plate 39, figs. 1, 2.)



Kossmaticeras zelandicum Marshall, Trans. N.Z. Inst., vol. 49, p. 444.

This species has already been described and figured (49, p. 444, pl. 33, fig. 2, text-fig. 2), but the figures now given are more complete. The specimen there described was sent to Professor Kilian, and I am much indebted to him for the note that the species is distinct, but very close to G. antarcticum Stuart Weller, so abundant at Seymour Island. The following dimensions were not given in the original description, and must be added:—

Diameter 75 100
Height of last whorl 33 44
Width of last whorl 28 37
Diameter of umbilicus 24 32

The species is distinguished from G. antarcticum Stuart Weller by the following characters: Ribs much more numerous, as many as thirty-six in a half-revolution. The constrictions generally interfere with the development of a single ordinary rib. Ribs more strongly bent forward than in G. antarcticum. Umbilical knots much more numerous. Suture-line generally similar to that of G. inflatum, but the external saddle is rather less symmetrically divided than in that species owing to the greater development of the external portion. The less depth of second lateral lobe is also a more pronounced feature. The nature of the auxiliary lobes unfortunately cannot be seen, nor does any specimen show the internal portion of the suture-line.

No complete specimen has yet been found. The only specimen comes from Batley. The type is in the Otago Museum.

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Gunnarites nordenskjoldi Kilian and Reboul. (Plate 21, fig. 8; Plate 36, fig. 6.)



Kossmaticeras antarcticum var. nordenskjoldi (46, p. 33, pl. 12, figs. 4, 5; pl. 14, fig. 2; pl. 15, fig. 3).


Diameter 12 100
Height of last whorl 5 42
Width of last whorl 5.5 46
Umbilicus 3.75 31

The specimen measured is the only one in a good state of preservation, and is quite juvenile.

Shell discoid and involute; about one-third of whorl exposed in umbilicus. Whorls almost circular in form. Nine large-tubercles in a revolution on the edge of umbilicus. From each of these three or four ribs take their origin, and usually two interstitial ribs arise between tubercles. All ribs very sharp, but diminish greatly in height towards periphery, and on periphery itself are seen with difficulty. Three constrictions in a revolution are distinct. They commence at the bottom of umbilicus and sweep over flank with a slight forward bend, which is much more pronounced on periphery. They cut off a single rib. Ribs inclined forward, but show no curve until periphery, when there is a strong forward curve. Ribs not denticulate in this young specimen.

Suture-line shows a median saddle about half the length of external saddle, which is relatively slender and not deeply bifid. Both lateral saddles are distinctly bifid, auxiliary saddles small, and the suture-line slopes rapidly backward so as to form a deep umbilical lobe. The external and first lateral lobes are of nearly the same depth, but second lateral lobe much shallower. Both lateral lobes are symmetrically bifid.

Professor Kilian was good enough to identify this specimen for me, and kindly sent me a cast of the Antarctic type for comparison. There appear to me to be some important differences. The ribs in the New Zealand specimen are sharper and are inclined farther forward. The umbilical tubercles also are far less numerous.

A single specimen of this species, found at Batley. Kilian and Reboul compare it to Madrasites karapadensis Koss., and remark that the sutureline is similar to that of M. Karapadensis and M. buddhaicus Koss. Actually, as will be seen from the figures in this paper, there are not many features of distinction between the suture-lines of the various species of Madrasites and Gunnarites—at any rate, in the external portion.

Gunnarites antarcticus Stuart Weller. (Plate 22, fig. 3; Plate 39, figs. 3, 4.)



Holcostephanus antarcticus Stuart Weller (35, vol. 11, p. 4, pl. 2, figs 1, 2).


Kossmaticeras (Gunnarites) antarcticum Kilian and Reboul (46, p. 31).


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A. B. C. D.
Diameter 22 100 19 100 65 100 84 100
Height 10 44 8 42 26 40 33 42
Width 9 41 8 42 25 39 31 39
Umbilicus 7 32 6 32 23 35 29 31

A, B, D, Gunnarites antarcticus Stuart Weller, Bull's Point, Kaipara Harbour, N.Z.; C, Gunnarites antarcticus, a cast of No. 135, given to me by Professor Kilian.

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This species has already been fully described by Kilian and Reboul (46, p 31), but, as the publication is not easily obtainable in New Zealand, a short description based on the New Zealand specimens is given here.

Shell discoidal, and large when mature. Whorls increasing rather rapidly in size; three whorls in a diameter of 8 mm. Whorls well rounded, but rather higher than wide. Umbilicus of moderate size, with a steep but not vertical wall. Ribs strong; some of them begin a little above bottom of umbilical wall, and at the angle they form twisted knots or tubercles. At this point they bifurcate, and there is usually one interstitial rib between each pair of knots. Every second or third rib is at first strongly inclined forward, but it soon becomes straight. At edge of periphery there is again a slight bend forward, and periphery is passed in a slight forward curve. About twenty-six ribs in a half-revolution, all of them crenulated. Six constrictions in a revolution, bordered in front or behind by a stronger rib, which is toothed like the rest, not smooth as in K. kalika.

No drawing of the suture-line of Gunnarites antarcticum has yet been published. Gunnarites kalika (Stol., 11, p. 140, pl. 70, fig. 5) was represented by a single specimen, which did not show a suture-line; and the many specimens from Seymour Island did not provide Kilian and Reboul with satisfactory material.

The suture-line in general closely resembles that of Madrasites, and we have excellent figures of several species of these by Kossmat (28, p. 39, pl. 7, figs, 1d, 5). The suture-line (Plate 22, fig. 3) is not very complicated; the saddles are distinctly bifid, though external half is rather larger than internal. First lateral lobe not quite symmetrical. First lateral saddle also distinctly bifid and nearly symmetrical, and considerably smaller than external saddle. Unfortunately, the whole of suture-line of this species cannot be seen, and the number of the auxiliary saddles is not known.

Gunnarites antarcticum is clearly related to G. kalika from the Aryalur beds of India, which are not lower than Upper Senonian and perhaps of Campanian age. Kilian and Reboul also compare this species to M. carlottense Whiteaves, from the Senonian of Vancouver (7, pl. 6), but the resemblance to this species does not appear to me to be a close one. Gunnarites kalika differs from this species in having four ribs springing from each umbilical tubercle, while the ribs which border the constrictions are not toothed. Kilian and Reboul make another comparison with Madrasites pachystoma Kossmat (28, p. 39, pl. 7, fig. 1 a-d), but the ribs in that species are not crenulated.

I am very much indebted to Professor Kilian for giving me a cast of Gunnarites antarcticum from Seymour Island, and for being good enough to identify my specimens for me.

Only three specimens have been found, all of which were collected at Bull's Point, but recognizable fragments were found at Batley and Whangaroa.

Madrasites Kilian and Reboul.

This was established by Kilian and Reboul in 1909 as a subgenus of Kossmaticeras, and was characterized by them in the following words (46, p. 25): “Un premier groupe comprend des formes, telles que K. karapadense Kossm. sp., K. bhavani Stol. K. theobaldianum Stol., qui conserve jusque dans l'adulte leur ornamentation caraéristique; nous designerons cette section sous le nom de Madrasites. Certaines formes, comme K. aemilianum Stol., prennent aussi, que l'a justment remarqué M. Kossmat, une ornamentation rappelant celle d'Astieria.”

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The New Zealand species that can be referred to Madrasites show no striking differences from these of South India which have been described and figured by Forbes, Stoliczka, and Kossmat. Several of them are clearly seen to be closely allied to Indian species, as will be stated in detail in the specific descriptions. It is interesting to find species that are considered to be closely allied to M. mooraviatoorensis Stol. and M. buddhaicum Kossmat in the same locality. These New Zealand species of Madrasites seem to be quite different from those described from Zululand and Pondoland by Spath (53, 54, 59), by van Hoepen from Pondoland (51, 52), from Japan by Yokoyama (17) and Jimbo (22), and, in my opinion, from those of Vancouver and of California. On the other hand, Madrasites tenuistriatus Paulcke from South Patagonia (36) is closely related to M. sulcatum n. sp., and at Seymour Island Kilian and Reboul recorded M. karapadense (46, p. 30).

Madrasites sulcatus n. sp. (Plate 38, figs. 1, 2.)

Dimensions :—

Diameter 33 100
Height of last whorl 13.5 43
Width of last whorl 12 36
Umbilicus 12 36

Shell discoidal and rather compressed. Whorls distinctly higher than wide. Involution about one-third. Umbilical slope very steep; the surface passes through 80° over a sharp curve. Lower half of flank slopes at first slightly outwards, then is almost flat, but soon slopes gently to periphery, over which it passes in a narrow curve.

There are three deep constrictions in a half-revolution, which always cut off three ribs. Each constriction has a prominent bolster on both sides. Ribs narrow and sharp, with wide interstices. They arise just above umbilical wall and hardly touch tubercles, which are numerous. There are forty-seven ribs in a half-revolution. Some of them begin quite close to an umbilical tubercle, but most of them originate rather higher on the flank and are almost straight. They do not fork, and are not tuberculate. Twelve umbilical tubercles in a half-revolution. Suture-line not sufficiently well shown to allow a satisfactory diagram to be drawn, but it can be seen that it does not differ very much from the other species of Madrasites

This species differs from all other known species of Madrasites. The relatively narrow aperture, close but straight ribs, and numerous tubercles are distinctive.

A single specimen has been found at Bull's Point. It was sent to Professor Kilian for examination, and he kindly sent the following note: “A new variety of Kossmaticeras nordenskjoldi K. & R.“

Madrasites multicostatus n. sp. (Plate 21, fig. 6; Plate 35, figs. 1, 2.)


  • Holcodiscus buddhaicus Kossmat (28, p. 42, pl. 8, figs. 3 a, b, c).

  • Holcodiscus karapadensis Kossmat (28, p. 41, pl. 8, figs. 2, 4).


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A. B. C. D. E.
Diameter 36 100 24.5 100 20 100 36 100 30 100
Height of last whorl 14.5 40 9.5 39 8 40 16 44 10.8 36
Width of last whorl 15.5 44 11 45 9 45 16.5 46 9.2 31
Umbilicus 13.5 37 9 37 8 35 11 31 11.5 38

A, B, C, Madrasites multicostatus from Bull's Point; D, Holcodisens buddhicus Koss; E, Holcodiscus kasapadensis Koss

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Shell moderate size with whorls that do not increase rapidly. Involution about one-third. Whorls rather wider than high, with a steep slope from umbilicus, then gradually curving to periphery, which is gently rounded.

Ornamentation: A row of nineteen or twenty tubercles, small and rounded, but varying considerably in size, situated at top of umbilical wall. From these arise a variable number of small rounded ribs, which follow a rather sinuous curve to periphery. At first they are directed slightly forwards, but after passing top of umbilical slope they bend backwards, and on edge of periphery they again bend forward and so continuing pass over periphery. Most of the ribs become quite indistinct in peripheral region. Interstitial ribs occasionally arise at various points. Here and there a number of fine striations can be seen parallel to the ribs. Sometimes striations occur on or between ribs, and sometimes they entirely displace them; in such cases they pass over periphery without interruption. Seven constrictions in last whorl. They arise from bottom of umbilicus and have a thick bolster on either side. They bend forward rather more than ribs, of which they intersect two or three. Constrictions form a conspicuous curve over siphuncle.

Suture-line quite typical of species of Madrasites so far as the main portion of it is concerned, but there is a sudden change of direction after second lateral saddle. The three auxiliary saddles are directed almost radially, and thus a most pronounced umbilical lobe is formed almost like that figured by Kossmat for H. bhavani (28, p. 38, pl. 8, fig. 5). In other respects, however, there is no resemblance between these two species.

Professor Kilian was good enough to examine the type of this species. In a note of a purely preliminary nature he classed it as Kossmaticeras buddhaicum n. var. multicostatum. I consider, however, that it is quite distinct from M. buddhaicum. The present species has a much higher whorl, and more distinct and fewer tubercles. Its ribs do not bifurcate nearly so frequently, and are more numerous—forty-two in place of thirty-one in a half-whorl. The ribs are more rounded and more often vanish on periphery. Suture-line is, of course, quite distinct from that of M. buddhaicum. Ribs in this species are not inclined forward as in M. karapadensis, and aperture is much wider. The constrictions are more numerous, being seven in number, compared with four in the Indian species. M. buddhaicum, which is probably the most closely allied species, is found in the middle horizon of the Trichinopoly group of the Indian occurrences.

Madrasites regularis n. sp. (Plate 21, fig. 7; Plate 35, figs. 3, 4.)


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A. B.
Diameter 30 100 24 100
Height of last whorl 10.25 34 9 37
Width of last whorl 12 40 10.25 43
Umbilicus 12 40 10 42

A and B, Madrasites regularis, Bull's Point, Kaipara Harbour, N.Z.

These dimensions differ slightly from those of M. multicostatus, immediately preceding.

The most notable features are that the height is small and the umbilicus wide. Shell of moderate size, discoidal. Whorls wider than high. Umbilical wall steep but sloping. Flanks sloping gently to periphery, which is well rounded. Involution about one-half. Whorls not increasing rapidly in height or width.

– 166 –

Ornamentation: Top of umbilical wall crowned with a row of fourteen to seventeen rounded tubercles. These are first noticed on second whorl at a diameter of 7 mm. From each of these tubercles rise two or three moderate rounded ribs, and usually two interstitial ribs begin at the same level in each interval. Ribs much narrower than intervening furrows, but height decreases slightly at periphery, which is crossed with a gentle forward curve. Four large and conspicuous constrictions in a revolution. They begin at bottom of umbilicus and are inclined forward more strongly than ribs, one or two of which they intercept. Constrictions bordered by a large bolster behind and a much smaller one before.

Suture-line in several features is not very different from that of M. multicostatus, but auxiliary saddles are less developed and umbilical lobe is not so conspicuous. Serrations of saddles more finely cut. External lobe wide and process at the bottom more developed. First lateral lobe extremely symmetrical.

This species is distinguished from M. multicostatus by the more regular umbilical tubercles, stronger and more continuous ribs, fewer constrictions, and much less pronounced forward curve of constrictions on periphery. When compared with the Indian species it is found that the shape of the whorl distinguishes it from M. karapadensis Kossm. (28, p. 41, pl. 8, fig. 4). Ribs more continuous, and have not the same strong forward bend at periphery as in the Indian species. Ribs not inclined forward so strongly as in M. bhavani (28, p. 41, pl. 8, fig. 4), which is probably different from Ammonites bhavani Stol. (11, p. 38, pl. 8, fig. 5). They are also less numerous, and the umbilical lobe is less pronounced. M. buddhaicus Koss. (28, p. 42, pl. 8, fig. 3) has less continuous ribs. Of the Indian forms this species resembles M. mooraviatoorensis Stol. most closely (11, p. 158, pl. 77, fig. 4), but the latter species has rather smaller tubercles, while its ribs are closer, the constrictions five to a whorl, and bent more strongly forward. Its suture-line is not dissimilar. This species is placed by Kossmat (28, p. 33) in his first division of Holcodiscus, which he says shows special relationship to the forms of this genus found in the chalk of Europe.

Three specimens have been found at Bull's Point and one at Whangaroa.

Madrasites fortior n. sp. (Plate 21, fig. 12; Plate 41, fig. 3.)


Diameter 13.5 100
Height of last whorl 5.5 41
Width of last whorl 6.4 47
Umbilicus 4 30

A, Madrasites fortior, Whangaroa Harbour, N.Z.

Shell of moderate size. Whorls nearly circular, increasing rapidly. Whorl rises rapidly from umbilicus and curves evenly into the rounded flank, which in its turn maintains the curve over periphery. Large and prominent smooth ribs, fifteen in a half-revolution, arise from tubercles at top of umbilical wall. Usually two arise from each tubercle, and there are rarely any interstitial ribs. They are inclined slightly forwards, and cross periphery with a slight curve and without any diminution in size. Seven constrictions in a revolution, and they have a stronger inclination than the ribs, but intersect only one of them. The bolster behind each rib is not larger than an ordinary rib. Constrictions cross ribs with a strong forward bow.

– 167 –

Suture-line not well seen; it is apparently of the typical Madrasites form, but less dissected than in the other species.

Only three specimens of this species have been found, two of them very small and immature. There is only one (imperfect) specimen of a mature form. Two come from Whangaroa, the third from Bull's Point.

Madrasites sp. (Plate 41, fig. 4.)

A fragment from Whangaroa in some respects resembles M. cumshewensis. No suture-line could be developed.

Jacobites Kilian and Reboul, 1909.

Kilian and Reboul regarded this group of species as constituting a sub-genus of Kossmaticeras, and gave the following diagnosis of it: “Dans la serie des Kossmaticeras anderssoni K. et R. nous assistons à une importante et rapide modification dès le diametre de 38 mm.; les côtes, fines et flexueuses des tours internes grossissent, s'espacent et finissent par se résoudre en tubercles et épines latérales; des tubercles siphonaux et une caréne apparaissent; ce type est assez analogue a celui que se trouve réalisé dans les Noumayria du Jurassique; cette curieuse section pourra porter le nom Jacobites “(46, p. 26). Up to the present time, unless the form known as Acanthoceras rotalinus Neum. be included, as Kilian and Reboul suggest, the group of Jacobites has not been found outside of Seymour Island. The authors did not give any diagram of the suture-line of Jacobites, but merely state that it was not clearly shown.

Professor Kilian was good enough to examine a few specimens from the present collection, and amongst them he recognized some specimens that were practically identical with J. anderssoni (55, pp. 175, 176). Since then I have obtained many additional specimens from Bull's Point, and one from Whangaroa, while Mr. J. A. Bartrum has given me a cast of a specimen from the entrance of the Wairua River, Hokianga.

The criteria that have been used for placing species in this genus are: The whorls have relatively flat flanks; ribs usually arise from umbilical tubercles and often bifurcate; irregularity in the ribs after a diameter of 60 mm. has been attained; the presence of spines on the flanks and the periphery.

I have obtained specimens of other species which I have no hesitation in placing in this genus, though in the one case the wider spacing of the ribs begins at a diameter of 40 mm. and in the other not before the diameter is 115 mm.

The species here called J. angulare is placed in this genus with some hesitation, because there is no sudden increase in the size of the ribs, though the increase in their development within the limits of a single revolution is considerable. There is a prominent angle on the edge of the periphery which almost has the dimensions of a spine, and there is also a small tubercle on the siphuncular line.

In general the suture-line differs but little from that of Gunnarites; in fact, Spath (54, p. 121) regards the genus Jacobites as a late Campanian development of the sparsicosta group of Kossmaticeras. The first lateral lobe is distinctly wanting in symmetry.

The affinity with M. sparsicostatus and M. denisoni mentioned by Kilian and Reboul does not seem to hold closely in these species.

– 168 –

Holcodiscus sp. described by Woods from Pondoland (46, p. 336, pl. 42, fig. 2) was compared by Kilian and Reboul to J. anderssoni (46, p. 35). Spath points out, however, that this species must be regarded as closely related to Madrasites madrasinus (54, p. 135).

Jacobites anderssoni Kilian and Reboul. (Plate 21, figs. 3, 3a; Plate 37, fig. 4.)



Kossmaticeras (Jacobites) anderssoni Kilian and Reboul (46, p. 35).

Several fragments that were sent to Professor Kilian were identified as belonging to this species.


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A. B.
Diameter 33 100 54 100
Height of last whorl 13 39 25 46
Width of last whorl 11 33 19 35
Umbilicus 9 27 16 29

A, Jacobites anderssoni K. & R., Bull's Point, Kaipara Harbour, N.Z.; B, Jacobites anderssoni K. & R. (46, p. 35), from measurement of cast kindly sent by Professor W. Kilian, No. 136.

Shell discoidal, with involution about one-half. Whorls higher than wide. Umbilical wall nearly vertical, with a sharply rounded angle to the flanks, which are nearly flat for about two-thirds of their height and then curve in an almost semicircular arc over periphery.

Ornamentation: Large number of tubercles at crest of umbilical wall. Numerous ribs arise from them at about one-third of the distance along the flank, and then pass over periphery with hardly any bend. Three constrictions in a revolution, bordered with a thick bolster before and behind; they are curved forward more sharply than ribs, two of which they intersect. Suture-line well shown. Median saddle slightly frilled. External saddle high and deeply bifid. First lateral saddle much smaller and second situated where tubercles arise at top of umbilical wall. Auxiliary saddles small and suture-line slopes back in umbilical region. External lobe not so deep as first lateral lobe. The latter is trifid but more developed on external side than on internal. Second lateral lobe about as deep as external lobe. Internal portion of suture-line has a deep antisiphuncular lobe and two high saddles separated by a lobe not so deep as the antisiphuncular lobe. Both saddles are bifid. Second saddle the wider, and on its external side are some projections that probably represent additional saddles. This internal portion of the lobe-line more closely resembles that of Gunnarites than that of Madrasites. It suggests an affinity with Acanthoceras.

From this description it will be seen that there is a considerable difference between this form and the mature form of M. anderssom (46, p. 35). The whorl in the New Zealand form is not so high, and the number of umbilical tubercles is a good deal larger. The specimens are immature, and nothing can be said about costation of mature whorls, which is so striking a feature in Kilian and Reboul's specimens from Seymour Island. Kilian and Reboul do not give a figure of the suture-line of the species, but they state that there is a resemblance between this species and K. pondicherrianum Koss. and K. sparsicostatum Koss. (46. p. 36), and they state that the horizon is that of the Valudayur beds.

– 169 –

Jacobites angularis n. sp. (Plate 21, fig. 2; Plate 36, figs. 4, 5.)


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A. B.
Diameter 55 100 38 100
Height of last whorl 23 42 15 39
Width of last whorl 25 45 17 45
Umbilicus 17 31 12 31

A, B, Jacobites angularis, from Bull's Point, Kaipara Harbour, N.Z.

Shell of small or moderate size. Whorls rather wider than high, with a distinct angle on edge of periphery as well as one much less distinct halfway between edge of periphery and umbilicus. Involution rather less than one-third. Whorls increasing rather rapidly in size.

Ornamentation: A few large ribs arise at bottom of umbilicus, and when they reach top of umbilical slope they develop into prominent elevated tubercles from which arise three and sometimes four ribs. There are also one or two interstitial ribs which commence on edge of umbilical wall between tubercles. Ribs at first bend slightly forward, and this bend is maintained until they reach edge of periphery, where each of them swells into a low rounded tubercle and bends backwards so as to pass over periphery almost at right angles, or with a slight backward curve. Ribs do not decrease in peripheral region, and on siphuncular line swell into small rounded tubercles. None of the ribs are serrated. In young stage the ribs when well preserved are very sharp and are situated quite close together; but when older they become spaced out considerably. Constrictions are not conspicuous, and do not interefere with the regularity of the ribbing.

Suture-line : Median saddle a little frilled and external lobe is not particularly deep. External saddle rather high, narrow, and deeply bifid; its first large external arm is well over top of median saddle. First lateral lobe slightly deeper than external lobe, and, as in the other species of Jacobites, it is not quite regularly trifid, for the external projection at bottom of this lobe is rather larger than internal one. First lateral saddle a good deal smaller than external one, and second lateral is on edge of umbilicus and its development is rather affected by umbilical tubercle. Five auxiliary saddles are small, slope backwards, rapidly diminish, and form a distinct umbilical lobe. This species is distinguished by its relatively wide aperture, strong umbilical tubercles, and the regular rounded tubercles on each of the ribs. In the available literature I cannot find a description of any species that comes close to this one.

Three specimens found at Bull's Point.

Jacobites minimus n. sp. (Plate 21, fig. 5; Plate 38, figs. 3, 4.)

This small shell is probably immature, but the very marked sculpture characterizes it as a different species from any other in this collection.


Diameter 8.75 100
Height of last whorl 4.1 47
Width of last whorl 3.75 43
Umbilicus 2 23

A, Jacobites minimus, Bull's Point, Kaipara Harbour, N.Z.

Whorls slightly compressed and strongly involute. Umbilical wall steep but not vertical. Widest part of whorl just above umbilicus, whence there is a gradual slope on flank towards periphery, on the edge of which is a rounded angle. Periphery gently rounded.

– 170 –

Ornamentation; On the edge of umbilicus are seven large tubercles in a whorl, from each of which two ribs arise, and there are two or four interstitial ribs in each interval. Seventeen ribs in a half-revolution. Ribs are rounded and at first bend slightly backward, but at a higher level on flank they take a gentle bend forward, and at edge of periphery there is a strong rounded tubercle on each rib. Across periphery the ribs run straight, and there is another rounded tubercle on median line. Between the tubercles the ribs are less elevated than on flanks. Suture-line shows five saddles, which gradually decrease in size from ventral surface to umilicus. The external and first lateral saddles are symmetrically bifid, and lobes trifid.

Lobes all of much the same depth, and suture-line bends backward to a considerable extent as it approaches umbilicus. On the whole, the suture-line has a close resemblance to that of a young specimen of Gunnarites (Plate 22, fig. 1b), but it is a good deal simpler, and more even

In the available literature I am unable to find any species that resembles this one closely. It is perhaps closer to A. idoneus Stol. than to any other, but differs from it in having umbilical tubercles and in the symmetrical development of the two sides.

The single specimen from Batley is probably immature, but is in such a good state of preservation and so distinct that it has been described as a new species.

Jacobites whangaroaensis n. sp. (Plate 21, fig. 1; Plate 37, fig. 1.)


147 100
Height of last whorl 63 43
Width of last whorl 56 38
Umbilicus 46 31

Involution about two-fifths. Whorls higher than wide. Umbilical wall steep but not vertical, rounded at angle. Whorl broadest at about one-third of its height above umbilical angle; thence it slopes gently to periphery, which is broad and well rounded.

Ornamentation: About ten large tubercles on edge of umbilicus; they are larger radially than longitudinally. Four ribs rise from each tubercle, and there is usually one interstitial rib between each pair of tubercles. Ribs close together and rounded on inner whorls, and more widely spaced on body-whorl, where at margin of periphery many of the ribs bifurcate and become extremely sharp and high and apparently spiny.

Suture-line only partially visible. Median saddle a little dissected on margin. External saddle high and deeply bifid, external half being slightly larger than internal half. First lateral lobe deeper than external lobe, and unequally trifid, for the external of the two major projections is rather larger than the internal one.

The species differs from J. anderssoni Kilian and Reboul in the sharpness of the ribs in the body-whorl and their regularity. They are also more closely spaced. There is no peripheral angle as in J. angularis n. sp.

A single specimen, found at Whangaroa.

Jacobites waitapuensis n. sp. (Plate 23, fig. 2; Plate 44, fig. 1; Plate 45, fig. 2.)


Diameter 100
Height of last whorl 40
Width of last whorl 31
Umbilicus 33
– 171 –

Aperture widely oval. Umbilical wall steep, with a rounded angle to the flank, which is moderately flat, though it has its. widest part rather nearer umbilicus than middle of flank, which slopes uniformly and gently to the well-rounded periphery.

Ornamentation: Five constrictions in a whorl, each with a prominent bolster before and behind. They are first inclined slightly forward, but in middle of flank they become almost truly radial, soon taking a stronger bend forward and passing over periphery with a distinct forward loop. At top of umbilical wall there are prominent rather rounded tubercles, eighteen in a whorl. From each of these arise three or four ribs, though some of these arise from bottom of umbilicus. Ribs at first bend slightly forward but soon straighten up and pass straight across periphery. Each constriction cuts off three ribs on each side. There are in addition low rounded tubercles on margin of periphery, twenty-three in a revolution, but unfortunately these can hardly be seen in the photograph. Indistinct tubercles also occur on periphery.

Sature-line: Median saddle frilled to a considerable extent, much more than in the two previous species. External saddle deeply bifid, and the two portions are about as unequal as in other species of this genus. First lateral saddle relatively rather larger than in the three previous species. The development of second lateral saddle is affected by umbilical tubercles. Auxiliary saddles, as in the other species, are strongly inclined, forming a deep umbilical lobe. The internal suture-line, which, however, is not figured, is close to that of J. angulare in form.

I am unable to find any described species that approaches closely to this one. The tuberculate periphery and median line, as well as the suture-line, distinguish it at once from all species of Kossmaticeras. The same features, as well as the more rounded form and coarser ribs, enable it to be distinguished at once from Maorites. It certainly comes nearer to Parapachydiscus than the other species of the genus. The whorl, however, is rather narrow, the ornamentation is distinctive, and the first lateral saddle shows lack of symmetry.

A single specimen, in good condition, from Nedler's, at Whangaroa.


Tuberciles on shoulder and on median line. Suture-line resembles that of Madrasites.

Neomadrasites nodulosus n. sp. (Plate 21, figs. 4, 4a; Plate 36, figs. 1–3.)


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A. B.
Diameter 29 100 20 100
Height of last whorl 11 35 7.5 37
Width of last whorl 13 45 9.25 46
Umbilicus 13 45 9 45

A, B, Neomadrasites nodulosus. from Bull's Point, Kaipara Harbour, N.Z.

Shell small, discoid. When the diameter is 29 mm. there are three whorls. Whorls decidedly wider than high, and increasing in width rather rapidly. Involution about one-half. Umbilical wall steep but rounding off into flanks that are gently inclined towards periphery, at edge of which there is a distinct angle, and periphery itself is gently rounded.

Ornamentation: At top of umbilical slope are a number of rounded tubercles, as many as seven in a half-revolution. Three or four sharp

– 172 –

but smooth ribs start from each tubercle, and there are two or three interstitial ribs between each pair of tubercles. Altogether forty ribs in a half-revolution. At edge of periphery ribs unite together in twos or threes into rounded tubercles situated irregularly with respect to umbilical tubercles, and here are also one or two interstitial ribs which do not touch tubercles. Ribs almost straight to edge of periphery, when they bend strongly forward and across it with a pronounced forward curve. At median line each rib has a small rounded projection or tubercle. On inner whorls the umbilical tubercles are well marked but the others are indistinct. In a half-revolution there are two constrictions with a prominent bolster on each side. These have the median projection, and they may join in the tubercles at edge of periphery.

Suture-line shows no striking features as compared with other related species of Kossmaticeratidae. Median saddle nearly simple, and external lobe not important. External saddle high and symmetrically bifid, as are lateral saddles, which rapidly decrease in size. First lateral lobe symmetrically trifid. At edge of umbilical wall suture-line bends sharply backwards and auxiliaries are directed almost internally. A similar sudden declension of suture-line is noticeable in Madrasites multicostatus (Plate 21, fig. 6) and in Brahmaites brahma (28, pl. 8, fig. 9). The internal portion of suture-line shows five saddles, which is quite distinct from that of Madrasites (28, pL. 7, fig. 5), and from that of Jacobites anderssoni (Plate 21, fig. 3a) as well as from Gunnarites.

A specimen sent to Professor Kilian was characterized by him as follows: “II convient de signaler en outre une forme trés curieuse, que je rattache à Jacobites, et dont M. Marshall m'a soumis un fragment, qui présente une modification progressive de l'ornamentation (apparition outre la présence des tubercles ombilicaux, de tubercles réunissant deux côtés du côté externe des flancs, et en outre, indication d'une série des tubercles siphonaux moins accentués, mais trés nets).” (55, p. 176.)

I have not adopted Professor Kilian's suggestion, because the development and drawing of the suture-line appears to me to indicate a closer relationship to Madrasites than to Jacobites. In its details, however, the ornamentation is distinct from both. I therefore suggest a new genus, Neomadrasites.

Several specimens, but none quite complete, have been found at Bull's Point.

Brahmaites Kossmat.

In the original diagnosis of this genus Kossmat says (28, p. 45) that it is very closely related to Holcodiscus (= Kossmaticeras Grossouvre). The most characteristic features are the presence of siphonal knots placed on ribs, the prominent bolsters bordering constrictions, and the highly variable nature of sculpture. In three specimens of middle age the flanks only are sculptured. At a later stage the ribs develop into high ridges which leap over periphery in a manner that is quite different from that of Holcodiscus.

The present species, B. rotundus, has the young form only, but the largest specimen shows a tendency towards the development of tubercles on the shoulder—a fact that definitely distinguishes it from B. brahma and the two other species B. vishnu and B. haugi.

So far as records go, the genus appears to be restricted to India. Some of the New Zealand specimens were sent to Professor Kilian, who was good enough to inform me that he agreed that they belonged to a new species of Brahmaites (55, p. 175).

– 173 –

Brahmaites rotundus n. sp. (Plate 21, fig. 9; Plate 31, figs. 4, 5.)


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A B. C. D. E.
Diameter 19.75 100 16 100 12 100 11 100 88 100
Height of last whorl 7 35 5 31 4 33 3.8 36 25 28
Width of last whorl 8.75 44 7.50 46 7 58 6.5 60 38 43
Umbilicus 7 35 5.75 36 4.75 39 4 39 42 48

A, B, C, D, Brahmaites rotundus, all from Bull's Point, Kaipara Harbour, N.Z.: E, Brahmaites brahma Forbes, in Koss. (28, p. 45).

The whorl is clearly much wider and the umbilicus narrower than in Brahmaites brahma.

Shell small, whorls much wider than high, especially in the younger specimens. Involution about one-quarter, and umbilicus deep. Wall of umbilicus steep but not vertical. Flank sharply rounded; periphery wide, with gentle curve.

Ornamentation: About eleven rounded tubercles in a revolution, situated on edge of flank near but not on edge of umbilicus. Usually three ribs start from each knob, and there are other interstitial ribs which start on side of umbilical wall. Ribs rather sharp, and separated by considerable intervals. At first they bend a little forward; the curve is soon increased, and near periphery it is strong, though the ribs themselves become less distinct. They cross periphery with a strong bow forward. Two or three deep constrictions in a revolution, bordered behind by a thick bolster. They have exactly the same forward inclination as the ribs, with which they do not interfere. Occasionally there is a slight suggestion of the presence of tubercles on edge of periphery, as in Neomadrasites nodulosus, but the ribs are not affected, and there is no indication of median tubercles on periphery, except that the bolster behind each constriction is swollen on the median line, as is also the case in B. brahma.

Suture-line was drawn from a very small specimen, but it is clear in all the specimens that the first lateral lobe is unsymmetrical. The irregular trilobed nature is due to the larger development of external than internal projection in this lobe. Suture-line does not show such a sudden backward curve in the region of auxiliary saddles as is seen in so many of the related species of Madrasites. This asymmetry of the first lateral lobe is not, however, noticeable in B. brahma. Internal portion of suture-line is more like that of B. brahma (28, pl. 8, fig. 9).

This species is not very different from B. brahma (28, p. 45, figs. 7—9), especially in the form of aperture and in ornamentation, though ribs are rather closer together. Its form is rather more rounded and suture-line is less divided.

Several specimens have been obtained from Batley and Bull's Point, but none at Whangaroa. This species is commonest at the first locality.

Maorites n. gen.

Genotype: Maorites tenuicostatus Marshall.

This genus is suggested for species that have the general form of Puzosia or Parapuzosia but possess umbilical tubercles—at any rate, in the young form. The suture-line is typically of the Puzosid type, though usually more highly developed than in the known genera of that group.

The general form has a highly involute whorl with flat flanks, the widest part of which is just above the steep umbilical slope. Ornamented with a number of ribs, usually quite fine. These at first are directed forward,

– 174 –

but on the flanks they become radial and cross periphery at right angles. About five constrictions more or less decidedly marked in a revolution. Many of the ribs arise from well-marked tubercles on edge of umbilicus, but there are always several interstitial ribs as well.

Suture-line: Median saddle a good deal frilled, and external lobe wide and deep. First lateral lobe has a highly important development. It is a good deal deeper than external lobe, and also is very wide. It is unsymmetrically bifid owing to the unequal development of the elements in lobe, those on external side being much larger and stronger than those on internal side. Second lateral lobe also unsymmetrical. Five or six auxiliary lobes are present and strongly inclined outwards. External saddle bifid, but with a much stronger development on external than on internal side. First lateral saddle also bifid, and in this case internal portion of saddle is a good deal larger than external. Second lateral saddle much smaller, and sometimes a little distorted by the tubercles, for it is situated on edge of umbilicus. Five or six auxiliary saddles quite small, and strongly inclined.

It is noticeable that Stoliczka placed his species Ammonites madrasinus, A. Kandi, A. aemilianus, A. bhavam, and A. kaloka in close proximity to Ammonites bhima, A. planulatus, &c., as though he thought that there were affinities between them. The last two of these species were afterwards classed by Kossmat in the genus Puzosia. On the other hand, Ammonites pacificus, A. papillatus, A. mooraviatoorensis were by Stoliczka far removed from those first named, though both groups were placed in the large and comprehensive class of Ligati (11, App. 6, 7). Kossmat, on the other hand, subsequently placed both of these groups (A. madrasinus and A. pacificus) in the genus Holcodiscus, but, of course, excluded A. bhima and A. planulatus. It seems, however that Kossmat did not have the opportunity of examining any of Stoliczka's specimens of the first group, but he saw one specimen of A. aemilianus in the museum at Vienna. Kossmat also had ten specimens that he ascribed to Stoliczka's species A. bhavani in the Warth collection, which was the one that provided most of the specimens that he studied. It is also noticeable that Stoliczka's drawings of the suture-lines of the species in the A. madrasinus group, though always somewhat sketchy, are in some respects distinctly different from those in the second, or A. pacificus, group, now mainly classed by Kilian and Reboul as Madrasites. (Compare 11, pl. 69, fig. 7, and pl. 70, figs. 3, 8, with pl. 77, figs. 5, 8, 9b, &c.). The suture-lines in the former are more divided, and they have dissymmetry in the first lateral lobe. In addition, the first lateral lobe is very deep. In the second, or A. pacificus, group, on the other hand, the suture-line is far less finely divided; there is almost complete symmetry in the first lateral lobe, which is relatively less important and either not deeper or very little deeper than the ventral lobe. The descent to the umbilical lobe is also far less regular in this second group. The shells of the first group also in most cases show a definite tuberculate character in the umbilical region, which is generally absent in the second group—that is, the ribs more decidedly and definitely arise from somewhat lamellar tubercles, which may be absent in the adult stage. The tubercles in the second group are much more rounded in their form.

The genus Maorites agrees with the species of the first group in the majority of their characters, and it is here suggested that the species Ammonites kandi, A. aemilianus, and A. madrasinus of Stoliczka should be placed in this genus, though this point cannot be definitely settled until

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the types of the Indian species which were described by Stoliczka are re-examined and their suture-lines have been carefully drawn.

In regard to Ammonites bhavani there is considerable difficulty. It is a form without any striking specific characters in its ornamentation; and in the absence of well-drawn suture-lines of Stoliczka's specimens it is hard to be certain whether a species in another collection is correctly identified with it. If Kossmat's specimens do belong to the true A. bhavani of Stoliczka there is no doubt that the species is correctly placed in the second group referred to above (the A. pacificus group). This has been done by Kossmat, and Kilian and Reboul have subsequently placed it in their subgenus Madrasites. On the other hand, Stoliczka's classification leads to the idea that his A. bhavani should be placed in the first of the two groups—that is, the group of Ammonites madrasinus. In order to make this point clear, a table is given in which the relative dimensions of Stoliczka's specimens of Madrasites bhavani are given as well as those of Kossmat, and also two of Kilian and Reboul's specimens of Kossmaticeras (Madrasites) bhavani var. densicostata, the last measurements being obtained from the casts that were kindly forwarded to me by Professor Kilian.

[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]

A. B. C. D. E. F.
Diameter 38 100 65 100 36 100 66 100 63 100 125 100
Height of last whorl 37 41 15 42 26 40 30 47 51 40
Width of last whorl 32 31 14.5 40 24 36 18 29 31 25
Umbilicus 24 26 11 30 22 33 13 21 31 25

A, B, Ammonites bhavam Stol. (11, p. 138); C, D, Holcodisens bhavani Koss. (28, p. 145); E, F, Kossmaticeras (Madrasites) bhavani var. densicostatum Kilian and Reboul (46, pl. 15, fig. 4; pl. 18, fig. 1).

These dimensions show that Kossmat's specimens are distinctly wider than the original ones of Stoliczka, and have a wider umbilicus, while Kilian and Reboul's specimens have a slightly narrower whorl.

Ammonites kalika appears to me to belong to a different group. It is the only Indian species of this nature that shows denticulations on the ribs; and it should almost certainly be classed with the species of Gunnarites described in the present collection. Stoliczka gives no drawing of the suture-line of this species, and Kossmat apparently saw no specimen of it. Those obtained in the present collection show that the species of Gunnarites are closely related to Madrasites, though the internal portion of the suture-line shows important differences if Kossmat's drawing of M. theobaldianus is taken as type. The New Zealand species M. regularis, however, shows a similarity to Gunnarites. The large and important collection of Antarctic species from Seymour Island described by Kilian and Reboul were in too bad a condition to allow the suture-lines to be traced.

The affinities of Maorites and its genealogy are a little hard to decide. As stated more fully subsequently, the author, in spite of the great weight of authority in favour of regarding Puzosia and Parapuzosia as derived from the Desmoceratidae, is himself of opinion that they are an independent branch derived from a Hoplitid ancestry. In connection with this it is important to recall the words of Sarasin, who wrote as follows (29A, p. 799): “Puzosia est du reste derivée directement de Desmoceras … Quant a l'origine de Desmoceras … il se rapproche nettement de Hoplites et il est a mon avis incontestablement derivée de Hoplites ou tout au moins de Perisphinctes.”

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Djanélidze in 1922 established Dalmasiceras, a new subgenus of Hoplites, from the Upper Tithonian of France (60, p. 256). The species of Dalmasiceras described by him differ so much in appearance and ornamentation from the typical Hoplites that doubt might well be entertained as to the real position of the genus. However, Djanélidze says, “Remarquons d'abord que Dalmasiceras est sans conteste une branche specialisée de Hoplites.” The forms of Maorites and of Puzosia are extremely similar to that of Dalmasiceras, and the same is equally true of the general features of the suture-line of these genera, both in its external and internal portions. There is, however, a difference in sculpture. Apart from the tuberculate nature of the ornamentation of the typical Hoplites, which is not developed to any extent in these genera, there is the marked break of the costation at the periphery. This feature, however, is not constant even in Hoplites itself, for Hyatt remarks that in the young forms the costation is sometimes continuous across the periphery (47A, p. 668). In addition, it is noticeable that Sonneratia, a genus included by Hyatt in the Hoplitidae, has continuous ribs across the periphery, and it therefore seems that this feature of ornamentation should not be emphasized too much. Maorites appears to be closer to the Hoplitid stock through Dalmasiceras than does Puzosia. In the suture-line the great depth and width of the external lobe, the depth of the first lateral lobe, and its asymmetry are more strongly marked than in Puzosia, and make a close approach to the conditions of the suture-line of Dalmasiceras, though they are more marked than in that genus. Tuberculation on the edge of the umbilicus also is present in Maorites, though absent in Puzosia. It is, however, to be noted that both Jacob and Nowak ascribe asymmetry to the flattening of the flank. This does not appear to hold in the New Zealand species; for, e.g., M. tenuicostatus, which is relatively rounded, has far more marked asymmetry than Puzosia angusta which has an extremely narrow form.

The first specimen of this genus that was found was classed by me under the name Kossmaticeras tenuicostatum (49, p. 445). This specimen was afterwards taken to Europe, and Professor Kilian was good enough to examine it. He classed it with his Madrasites densicostata from Seymour Island (55, p. 175). When the suture-line was developed and drawn, it at once became clear that the specimen could not belong to either Kossmaticeras or Madrasites, and that, on the other hand, it possessed close affinities with Puzosia. Professor Kilian, with the greatest kindness, sent me a cast of two specimens of his Madrasites bhavani var. densicostata, and inspection of these showed that in all probability this species belonged to the same genus as the New Zealand species. The condition of the Seymour Island specimens was, however, so bad that Professor Kilian was not able to develop the suture-lines. I have already stated that in my opinion some of the Indian species described first by Stoliczka, and afterwards placed by Kossmat in the genus Holcodiscus, and later in Kossmaticeras (Madrasites) by Kilian and Reboul, especially K. madrasinum, K. kandi, and K. aemilianum, may also possibly belong to this genus.

The characters of the genus Maorites may be summarized as follows: Form somewhat compressed and resembling that of Puzosia. Ornamentation shows umbilical tubercles a little extended radially. Ribs generally rather fine and sinuous, but continuous across periphery, which they cross without a forward bend. Constrictions deep and with a strong forward inclination. Suture-line in general resembles that of Puzosia, but the first lateral lobe is a little deeper and the external lobe is wide. The unsymmetrical development of the first lateral lobe is marked. The suture-line is greatly dissected.

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Maorites tenuicostatus Marshall. (Plate 23, figs. 1, 1a; Plate 42, figs. 1, 2; Plate 45, fig. 1.)



Kossmaticeras tenuicostatum Marshall (49, p. 445, pl. 33, fig. 1).

Dimensions :—

[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]

A. B. C. D.
Diameter 142 100 143 100 20 100 63 100
Height of last whorl 60 42 60 42 9 45 30 47
Width of last whorl 43 30 43.5 31 7 35 18 29
Umbilicus 35 24 35 24 5 25 13 21

A, B, C, Maorites tenuicostatus Marshall, Batley, Kaipara Harbour, N.Z.; D, Kossmaticeras bhavani var. densicostatum K. & R. (measurements from a cast).

Shell large, discoid. Involution rather more than one-half. A steep rounded slope, nearly vertical in young forms, rises from umbilicus, and greatest width of whorl is attained just above umbilical slope. Flanks nearly flat but with a gently increasing slope towards periphery. Edge of periphery well rounded, and a well-rounded curve over periphery.

Ornamentation: In mature form constrictions not distinct, and in no way interfere with costation, for they have the same slope as ribs. A large number of narrow rounded ribs, which in mature form are almost constant in size, from bottom of umbilicus to periphery. In younger forms ribs do not all extend to bottom of umbilicus, and on its margin about every fifth rib is swollen and almost forms a small tubercle extended in a radial direction. In the youngest forms there are no ribs or tubercules and the whorl is much wider. Some interstitial ribs arise on lower part of flank, but very few above a third of distance from umbilicus. With a radius of 70 mm. there are thirteen on a 10 mm. arc of periphery; at 55 mm. fifteen ribs; at 23 mm. twenty ribs; at 12 mm. twenty-eight ribs. The ribs bend slightly backward at top of umbilical edge; thence to the middle of flank there is a forward inclination, but from that point the ribs pass straight over periphery without any curve.

Suture-line is highly dissected. Median saddle much ornamented. External saddle deeply bifid, but external half much larger than internal half. First lateral saddle also deeply bifid, but internal portion much smaller than external portion. Second lateral saddle small. Six auxiliary saddles. External lobe wide with several frilled projections. First lateral lobe very deep and noticeably wanting in symmetry, for the projections and forks are much more developed on external than on internal side. Second lateral lobe much shallower, but still wanting in symmetry. Auxiliary lobes small and unimportant, but with the saddles form a deep umbilical lobe. Antisiphonal lobe also deep and again wanting in symmetry. The saddle next to it is very high, but the four others rapidly decrease in height, and the lobes are small, and thus the internal side of umbilical lobe is extremely steep. Both internal and external portions of suture-line have a remarkable resemblance to Dalmasiceras, and a rather less marked one to Puzosia.

A specimen of this species was sent to Professor Kilian, who kindly examined it, and returned it with a note: “Kossmaticeras (Madrasites) n. sp., possibly the mature form of M. bhavani var. densicostatus.” I consider, however, that it is to be distinguished from that species by the fineness of the costation, the gradual slope into the umbilicus, the indistinct constrictions, the small size of the umbilical tubercles.

Two adult specimens and several smaller ones from Batley and Bull's Point, and a small one from Whangaroa. The type is in the Otago Museum.

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Maorites densicostatus Kilian and Reboul. (Plate 24, fig. 1; Plate 44, fig. 2; Plate 45, fig. 4.)

Compare the species mentioned below under Maorites suturalis n. sp.

Dimensions of species of Ammonites apparently related to Maorites

densicostatus :—

[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]

A. B. C. D. E. F. G.
Diameter 49 100 55 100 60 100 93 100 63 100 125 100 142 100
Height of whorl 23 47 40 50 42 30 47 51 40 60 42
Width of whorl 12 24 34 36 21 18 29 31 25 43 30
Umbilicus 12 24 33 16 29 13 21 31 25 35 24
H. J. K. L. M. N.
Diameter 100 74 100 36 100 66 100 38 100 65 100
Height of whorl 40 35 47 15 42 26 40 37 41
Width of whorl 31 22 30 14.5 40 24 36 32 31
Umbilicus 33 18 24 11 30 22 33 24 26

A, Maorites densicostatus K. & R., Bull's Point, Kaipara Harbour, N.Z.; B, Ammonites kandi Stol. (11, pl. 70, fig. 4); C, Ammonites aemilianus Stol. (11, pl. 70, fig. 7); D, Ammonites mudiasinus Stol. (11, pl. 70, fig. 1); E, Kossmaticeras (Madrasites) bhavni var. densicostatum Kilian and Reboul (46, pl. 15) (measured from a cast kindly presented by Professor Kilian); F, Kossmaticeras (Madiasites) bhavani var. densi-costatum K. & R. (46, pl. 18, fig. 1): G, Kossmaticeras tenuicostatum Marshall (49, pl. 33, fig. 1); H, Jacobites waitapuensis n. sp., Whangaroa, N.Z.; J, Maorites, suturalis n. sp. Batley, Kaipara Harbour, N.Z.: K, L, Kossimaticeras bhavani Kossmat (28. p. 145); M, N, Ammonites bhavani Stol. (11, p. 138).

The table shows clearly how very similar the dimensions of this species are to those of Kilian and Reboul's form Kossmaticeras (Madrasites) bhavani var. densicostatum; but the suture-line of that species is said by the authors to be quite similar to the suture-line drawn by Kossmat of the form that he identified with K. bhavani Stol., and in that case it must be radically different from the suture-line of this species, which, as will be seen on Plate 24, fig. 1, is quite Puzosid in form. From inspection of the cast of his species that Professor Kilian was good enough to send me, however, it seems to me that the suture-line if developed would be similar to that of this species.

Ornamentation: There are numerous fine sharp ribs narrower than the intervening furrows, and narrower and sharper than in the closely allied species Maorites suturalis. Ribs at first bent forwards, but at about one-third of the flank they bend backwards and, becoming strictly radial, cross periphery without interruption and without any bend. Most of ribs start from tubercles at top of umbilical wall, but there are several interstitial ribs as well. A few deep constrictions, most noticable in last whorl. They have a moderate bolster-rib before and behind, and as they have a strong forward inclination they intersect several of the ribs, many of which fork close to posterior bolster. Constrictions cross periphery with a pronounced forward curve. Umbilical tubercles are numerous and are elongated radially.

Suture-line much less complex than that of M. tenuicostata and M. suturalis. Median saddle not so much divided. External saddle deeply bifid, but, as usual, exterior portion is a good deal larger than interior. First lateral saddle unsymmetrically bifid. Second lateral quite small. and auxiliaries rapidly diminish in size and are steeply inclined. External lobe wide, but not nearly as deep as first lateral lobe, which has the usual want of symmetry, is wide and deep, and extends over almost half of length of suture-line. Second lateral lobe much smaller, and auxiliaries small and unimportant. The inclination of these elements causes suture-line to form a deep umbilical lobe. It is noticeable that, as is also the case in the other species of Maorites, the external saddle is exactly the same

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width as the first lateral lobe; while in Puzosia it is about three-quarters of the width, and in Desmoceras the two are about equal. The external lobe in this genus is of greater importance than in other genera, and this makes the external saddle appear much more upright than in Puzosia.

One specimen only has been found, and it comes from Batley. The Indian species to which it is closely allied come from the Aryalur group, except K. bhaiani Stol. This comes from the Trichinopoly group, which is classed in the Lower Senonian, while the former are placed in the Upper Senonian. The New Zealand species has a rather broader and more rounded periphery than the Seymour Island specimen which it so closely resembles.

Maorites suturalis n. sp. (Plate 23, fig. 3; Plate 43, fig. 1; Plate 45, fig. 5.)


  • Ammonites kandi (11, p. 140, pl. 70, fig. 4).

  • Ammonites aemilianus (11, p. 141, pl. 70, fig. 7).

  • Ammonites madrasinus (11, p. 139, pl. 70, fig. 1).

  • Kossmaticeras (Madrasites) bhavani var. densicostatum (46, p. 30, pl. 18, fig. 1; pl. 15, fig. 4).

Dimensions :—

[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]

A. B. C. D. E.
Diameter 74 100 55 100 60 100 92 100 63 100
Height of last whorl 35 47 40 50 42 30 47
Width of last whorl 22 30 34 36 21 18 29
Umbilicus 18 24 33 16 29 13 21

A, Maorites suturalis, type, Batley, Kaipara Harbour, N.Z.; B, Ammonites kandi (11, p. 140, pl. 70, fig. 4); C, Ammonites aemilianus (11, pl. 70, fig. 7); D, Ammonites madrasinus (11, pl. 70, fig. 1); E, Kossmaticeras (Madrasites) bhavani var. densicostatum (46, pl. 15, fig. 4).

It is evident that the dimensions of this species are very like those of Kilian and Reboul's species from Seymour Island.

Shell of moderate size, strongly involute (about two-fifths) and a good deal compressed. Umbilicus has a nearly vertical wall, and at the top of it the whorl has its greatest thickness. The sides are almost flat and the periphery is well rounded.

Ornamentation: Well-marked constrictions arise at intervals—about five in a revolution. They are inclined slightly forward and interfere some-what with costation. They have a thick bolster before and behind. Ribs numerous, high, steep-sided, and narrowly rounded. Some of the ribs begin at umbilicus and nearly always bifurcate at top of the wall and interstitial ribs frequently begin at this point. Ribs at first incline forward, but at a distance of one-third of height of flank they bend back and become almost exactly radial. They pass straight over umbilicus without any diminution in size and without any bend. Constrictions have such a pronounced bend forward that they may interfere with as many as seven ribs; they are nearly straight, but bend slightly forward at about the middle, and pass over periphery with a strong forward curve. There are about fourteen umbilical tubercles in a half-whorl, and most of the ribs originate from them.

Suture-line is very much divided but is clearly of the type of Dalmasiceras, though the external lobe is of far greater importance. It can also be compared with the Desmoceras difficile and beudanti type, which Kilian and Reboul regard as derived from a Hoplitid ancestry (46, p. 17, footnote). First lateral lobe of great depth and importance, and markedly inequilateral hough bifid. There are two prominent secondary saddles, one on each

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side of median line of lobe. The external of these as well as the other elements of the lobe on external side are much larger than those on internal side. Second lateral saddle small and much affected by development of umbilical tubercles. Auxiliary saddles small and progressively inclined, and thus suture-line forms a prominent and deep umbilical lobe. None of the specimens show internal portion of suture-line. Saddles all deeply bifid but not quite symmetrically. In external saddle exterior side is more strongly developed than interior side. In the first lateral saddle, however, the reverse is the case, interior portion being larger and more developed than exterior. Median saddle considerably frilled.

This species is clearly quite closely related to Kossmaticeras (Madrasites) bhavani var. densicostatum, which Kilian and Reboul have described from Seymour Island, but costation is a little coarser and umbilicus somewhat wider. The suture-line of that species has not yet been drawn, but reasons have been given on a previous page for thinking that it should not be placed in the genus Kossmaticeras, and that it is closely related to Puzosia.

The type specimen was sent to Professor Kilian, and after a brief examination of it he classed it as Kossmaticeras (Madrasites) bhavani var. densicostatum; but it appears to me that the costation and diameter of umbilicus are sufficient to distinguish it from the species bhavani. Further, the nature of the suture-line is entirely different from that of all the New Zealand species of Madrasites, and is certainly most distinct from that of M. bhavani given by Kossmat (28, pl. 8, figs. 5, 6), of which, however, Kilian and Reboul say, “La ligne suturale de notre espéce est bien conformé a celle du type” (46, p. 29).

This species is quite distinct from K. haumuriensis Hector as described and figured by Woods (57, p. 34, pl. 19, fig. 5a; pl. 20, fig. 1), for the suture-line of that species is of the K. bhavani (Stoliczka in Kossmat) type; the costation also is of a type different from that shown in the present species. No species that seems to be closely related to this one has yet been recorded from the other countries in which the Indo-Pacific fauna of Ammonites has been described, unless the species Desmoceras hoffmani from California be regarded as similar. It seems, however, that this species, in spite of its compressed form, is a true Desmoceras. Only one good specimen of Maorites suturalis has yet been found, though with it have been discovered a few fragments of other specimens at Bull's Point, the typical locality.

Puzosia Bayle.

Ammonites mayorianus was taken by Bayle as the genotype of this genus. Of it de Grossouvre (19, p. 171) says, “I keep the name Puzosia for the forms of a group of ammonites (subplanulata) which have a moderate umbilicus the whorls of which present transverse furrows and sickle-shaped ribs, which, however, are absent near the umbilicus and are marked only on the external region of the flanks. The suture-line offers a complete similarity to that of Desmoceras but is more reduced, and the first lateral lobe is longer than the ventral lobe. The lobes are trifid, and are somewhat more upright than in Desmoceras. The saddles, which are almost all similar, are divided into two parts by a deep lobule.” He takes the suture-line of D. subplanulatum as typical. Sarasin (29A, p. 793) says that, as the genus had not been clearly defined, it had been little adopted by palaeontologists. He characterizes it as follows: “La coquille est

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moyemment involutée avec les tours arrondis ou légèrement aplatés sur les côtés, toujours arrondis sur le pourtour externe, le pourtour de l'ombilic n'est jamais caréné et generalment arrondi, les tours sont marqués de nombre variable des constrictions droits ou flexueuses et regulièrement espacées entre les sillons. La coquille est tantôt Iissée tantôt ornée de fines côtes, attenuée sur la partie interne des tours. Les cloisons sont toujours très découpées, le lobe ventral est le même longueur ou un peu plus court que le premier latéral; la selle ventral trés reserrée à sa base s'élargit à la partie supérieure qui est profondement divisée par un lobe accessoire. Le premier lobe latéral est symétrique ou subsymétrique; la première selle latérale est généralment un peu plus elevée que la selle ventrale; sa partie interne est presque constamment plus haute que sa partie externe; le second lobe latéral, moins long et large que le premier, est toujours très disymmétrique; la deuxième selle latérale est moins haute que la précédente, elle est encore trés découpée. Ensuite viennent trois à cinq lobes auxiliaires qui sont tantôt droits ou tantôt au contraire très obliques.”

He classifies Desmoceras in four groups—(1) D. difficile, (2) D. beudanti, (3) D. emmerici, (4) D. mayorianum d'Orb. = Amm. planulatus Sow. The last two of these divisions are considered by him to constitute the subgenus Puzosia.

Zittel (11A, p. 465) divided the genus Desmoceras into five groups—(1) D. beudanti, (2) D. difficile, (3) D. emmerici, (4) D. planulatum, (5) D. gardeni. Kilian and Reboul refer the beudanti and difficile groups to the family of Hoplitidae, the mayoriana and angladei groups to the Phylloceratidae.

Kossmat in 1897 (28, p. 106) distinguishes between three groups of Desmoceratids: (1) Desmoceras emmerici Rasp. with a very regular suture-line. The separate lobes and saddles decrease little by little and extend to the umbilicus. (2) D. planulatum Sowerby (Puzosia Bayle). A distinctly-developed depressed umbilical lobe. The external lobe is almost always a good deal shorter than the first lateral lobe, and this makes it appear as though the external saddle were inclined outwards. The various lobe elements are not arranged with any regularity. He states that this group is closely related to Ammonites beudanti d'Orb. on the one hand, and with Holcodiscus through H. pondicherryianus on the other, as well as with Pachydiscus. (3) D. gardeni (Hauericeras de Grossouvre) includes keeled forms from the Upper Chalk, which in their other characters come very close to the group Ammonites planulatus.

Kihan and Reboul make the following comment (46, p. 17): “On réunit souvent á tort nous semble-t-il, en une même groupe, les Desmoceras s. str. (groupe beudanti-difficile) et les Puzosia (P. angladei, P. mayoriana, etc.) ces deux rameaux ne semblent cependant avoir entre eux aucun rapport genetique; le premier dérive sans doute des Holplitides (sensu lato) specialement de Leopoldia, et le second peutêtre des Phylloceratides par l'intermediare de Sowerbyceras (Tortisulcati) et de Silesites?”

Zittel in 1895 and Hyatt in 1913 included Puzosia and Hauericeras in the Desmoceratidae. This relationship was the one adopted also by Yabe in 1903 (37, p. 30).

Pervinquière (“Sur quelques Ammonites du Crétacé Algerien,” Mem. Soc. Geol. Fr., vol. 17, 1910, p. 31) appears to use Puzosia in the sense defined by Sarasin.

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Jacob (44, p. 26) divides Desmoceras into four groups, the last of which is Puzosia, with the genotype P. mayoriana, which has a symmetrical lobe. This author (p. 55) places several of Sarasin's second group of Desmoceras in Sonneratia, which is referred to the Hoplitidae.

Spath in 1922 makes the following remark (54, p. 120): “The genera Pachydiscus, Parapachydiscus, Parapuzosia, Kossmaticeras, and the many allied developments can all be derived from Desmoceratid stocks which persisted during the Upper Cretaceous.”

In spite of the weight of these authoritative opinions, and with comparatively a small amount of material to study, the author, relying on the character of the suture-line (both external and internal), as well as on the form of the shell, is inclined to regard Puzosia, Hauericeras, and Maorites n. gen. as derived from the Hoplitidae sensu lato.

Puzosia angusta n. sp. (Plate 22, fig. 5; Plate 41, fig. 1.)



Ammonites durga Forbes, in Stol. (11, p. 143, pl. 71, figs. 6, 7).


Puzosia compressa Koss. (28, p. 119, Taf. 18, fig. 4 = Ammonites durga Forbes, in Stol.).

Dimensions :—

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A. B.
Diameter 143 100 190 100
Height of last whorl 52 36 64 34
Width of last whorl 18 12.5 34 18
Umbilicus 53 37 74 39

A, Puzosia angusta n. sp., Bull's Pomt, Kaipara Harbour, N.Z.; B, Puzosia compiessa Koss. (28, p. 119).

The dimensions given above emphasize the feature of the high and narrow whorl, which seems to be more extreme than in any other species of this genus of which descriptions are available. Involution nearly one-half, and whorls rapidly increasing in height. Flanks nearly flat and periphery sharply rounded, with the broadest point about two-thirds of distance from umbilicus to periphery. Umbilical wall short but vertical, passing into flank over a sharp angle of a little more than 90°.

Ornamentation: Lower two-thirds of whorl without costation, but upper third with numerous low rounded ribs arising without any tubercles or projections on the shell. First they are directed slightly forward, but they soon bend to a much more acute angle, and, though they diminish in size towards periphery, they distinctly pass over it with a sharp forward loop. Prominent constrictions, apparently five in a revolution, take their rise from umbilicus, and each of them has a high bolster behind. At first the constrictions slope forward at a moderate angle, and near top of flank they bend farther forward and become parallel with ribs and pass over periphery with a sharp curve forward.

Suture-line of the typical Puzosid form, not dissimilar from that of P. compressa, but more finely divided. In the only specimen that shows a suture-line the external saddle, unfortunately, cannot be seen. First lateral lobe deep and, as usual, wanting in symmetry. First lateral saddle almost equally bifid, and auxiliary lobes and saddles, few in number, form a deep umbilical lobe.

This species is certainly closely related to Puzosia compressa Koss., which comes from the Utatur formation of India, regarded as of Cenomanian

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age. It appears to be confined to this formation, for the small specimen figured by Stoliczka is classified by Kossmat as Hauericeras rembda.

Three fragmentary specimens only have been found, and all come from the north side of Bull's Point.

Parapuzosia Novak, 1913.

Novak states that the inner whorls have the typical Puzosia form with constrictions. These disappear in the outer whorls and ribbing is developed (45, p. 350).

Novak established this genus for the Puzosia demsoni group; but, as Spath points out, Novak described and figured P. daubréei only, and consequently Spath takes P. daubréei as the genotype of Parapuzosia. This genus, he says, has the course of the radial line and the strongly projected constrictions of the P. subplanulata type, different from the straight ornamentation of the Campanian (54, p. 127). He includes in this genus the Indian species P. gaudama Forbes and P. indopacifica Koss. In both these species the ribs cross the periphery and there are no tubercles.

At the same time Spath established the genus Kitchinites, including therein K. pondicherryanus Koss., K. japonicus Spath = Desmoceras gaudama (Forbes) Yokoyama, as well as K. darwini Phil., all rather coarse-ribbed types, with the ribs crossing the periphery though they are absent on the lower part of the flanks. There are no tubercles. The first of these three species has a typical Madrasites suture-line. K. darwini, however, has a suture-line of Puzosid type, and the strong projections of the constrictions of that type.

The New Zealand species Parapuzosia brevicostata appears to be closely related to P. gaudama on the one hand and to Kitchinites darwini on the other; but I am inclined to retain it in the genus Parapuzosia, as explained later.

Parapuzosia brevicostata n. sp. (Plate 24, fig. 3; Plate 43, fig, 2.)



Ammonites gaudama Forbes (1, p. 113, pl. 10, fig. 3).


Ammonites planulatus Sowerby, in Stol. (11, p. 134, pl. 67, fig. 1).


Desmoceras gaudama Forbes, in Yokoyama (17, p. 184, Taf. 18, fig. 14; Taf. 19, figs. 5 a, b) = Puzosia indopacifica Koss. (28, p. 117) = Kitchinites japonicus Spath (54, p. 127).


Puzosia sp. Kilian and Reboul (46, p. 19).


Puzosia darwini Steinmann (26, p. 73, Taf. 5, figs. 3 a, b, c, fig. 4) = Kitchinites darwini in Spath (54, p. 127).


Parapuzosia gaudama Forbes, in Spath (54, p. 126).

Dimensions :—

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A. B. C. D.
Diameter 56 100 89 100 56 100 190 100
Height of last whorl 26 46 37 42 23 41 64 34
Width of last whorl 13 26 27 30 19 34 34 18
Umbihcus 19 34 25 28 18 32 74 39

A, Parapuzosia brevicostata, specimen from Nedler's, Whangaroa, N.Z.; B, Parapuzosia gaudama (28, p. 115); C, Puzosia planulata (28, p. 112); D, Puzosia compressa (28, p. 119).

Periphery gently rounded, the curve passing rapidly into a long flat flank. A short curve at the edge of umbilicus passes quickly into a vertical umbilical wall.

Ornamentation: A number of low rounded ribs commence at umbilicus and are at first directed radially, but towards outer side of flank they curve

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forward and become thinner, and are almost lost at periphery. At the point where the forward bend commences a secondary rib arises between each pair of primary ribs. Deep constrictions arise at intervals, but only two in a half-whorl. They are at first radial, but soon bend forward more strongly than ribs, and are continuous across periphery, which they cross in a strong almost V-shaped curve.

Suture-line has a moderately deep external lobe, which is deeper than is usual in Puzosia, though little more so than in Parapuzosia gaudama (28, pl. 16, fig. 4) and Puzosia crebrisulcata (28, pl. 18, fig. 2). First lateral lobe, as usual, is of great importance, but in this species is rather more symmetrical than usual. External saddle rather wide, bifid, with a rather wide median lobe. Second lateral lobe very small, and three auxiliaries much inclined, forming a deep umbilical lobe.

This species differs from P. gaudama in its narrower form, distinction between primary and secondary ribs, and in the failure of the ribs to maintain their strength across periphery. The same characters distinguish it from P. crebrisulcata, and in addition the small number of constrictions it shows. Kitchinites darwini from Quiriquina has rather stouter ribs which cross periphery, a broader form, more numerous constrictions, and a suture-line more finely divided. The variety P. gaudama var. intermedia does show primary and secondary ribs, and, while it is not so narrow as P. compressa, it has a smaller umbilicus. P. compressa, also, has little sculpture (28, p. 119).

Suture-line shows a general similarity to that of P. gaudama, but its lobe elements are more nearly symmetrical, the external lobe is more transgressive, and median saddle is more frilled. P. stoliczkai Koss. has a median saddle that is nearly simple, and its ribbing consists of wide low rounded ribs.

A single specimen only, in indifferent state of preservation, has been found at Whangaroa.

Parapuzosia ordinaria n. sp. (Plate 24, fig. 4; Plate 31, figs. 6, 7.)



Ammonites bhima Stol. (11, p. 137, pl. 69, figs. 1–3).


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A. B. C.
Diameter 28 100 48 100 130 100
Height of last whorl 12.5 45 41 46
Width of last whorl 9 32 33 34
Umbilicus 8 28 30 23

A, Parapuzosia ordinaria Batley, Kaipara Harbour, N.Z.; B, Ammonites bhima Stol. (11 p. 137); C, Ammonites bhima Stol. (11, p. 137).

Shell small; involution nearly one-half; whorls much higher than wide and rapidly increasing. Umbilical wall at first steep, then gently sloping, with the thickest part of whorl rather more than half the distance from umbilicus to sharply-rounded periphery. Umbilicus shallow.

Ornamentation: Surface of shell practically destitute of ornament except for very fine striations or growth-lines, which at first bend backward but on the flank curve forward and pass over periphery with a sharp forward curve. This, however, does not prevent the shell from having a bright and polished appearance, and a strong lens is required to make the striations distinct. Seven constrictions in the last whorl, bordered behind by a rounded bolster rib. They are directed forward at first, and on upper

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part of flank they bend still more forward, finally passing over periphery in a sharp forward curve.

Unfortunately the suture-line is not well preserved. External lobe shallow and narrow. First lateral lobe a good deal deeper and very wide, and nearly symmetrical. Saddles bifid but not deeply dissected. Second lateral saddle already on umbilical slope. Three auxiliary saddles all strongly inclined and forming a deep umbilical lobe.

The sloping nature of umbilical wall, the smooth surface, as well as the relatively simple suture-line, distinguish this from all other species of Parapuzosia of which descriptions can be found in the available literature. It seems to be closer to Puzosia bhima Stol. (11, p. 137, pl. 69, figs. 1–3) than to other species. However, this species comes from the Cenomanian, and is apparently related to Ammonites octosulcatus in the Grey Chalk in the Isle of Wight. It is probable that the resemblance is more apparent than real.

A single specimen has been found at Batley.

Tainuia n. gen.

Genotype: Tainuia aucklandica n. sp.

Shell with flat flanks, especially in the inner whorls. Costation strong and coarse, with ribs strongly inclined forwards. Five constrictions in a whorl running parallel to ribs. Umbilical and siphuncular tubercles on each rib. On each rib a number of rounded tubercles. Suture-line much dissected. Median saddle rather frilled. First lateral lobe bipartite but not quite symmetrical, much deeper than external lobe. Saddles bifid and suture-line sloping backward rapidly from edge of umbilicus.

At first sight the ornamentation of this form suggests that it belongs to the genus Acanthoceras; but the constrictions distinguish it from that genus, and the distinction is emphasized by the suture-line with its deep unforked first lateral lobe. On the other hand, Kossmat has a seventh group of species of Acanthoceras (28, p. 25), “VII Arten aus der Verwand-schaft des Acanthoceras vicinale,” which has the main characters of form and ornamentation (except the constrictions) and the deep slightly unsym-metrical bipartite first lateral lobe. This new genus Tainuia is evidently close to this group, the members of which are far from typical species of Acanthoceras. This is also true of the compressed variety of Acanthoceras rotomagense Stol. (11, pl. 34, figs. 5 a–c), which, however, is placed by Kossmat under A. gothicum, a much less compressed form with a very different suture-line (28, p. 198, Taf. 25, figs. 3 a, b).

In external form A. pseudodeverianum Jimbo (22, p. 32, Taf. 5, figs. 1 a, b) closely resembles the genotype, but here again the suture-line is very distinct.

The genus resembles Parapuzosia, in respect of its compressed form, inclination of the constrictions, and in the nature of the suture-line itself. On the other hand, it is wholly different from Parapuzosia in the character of its ornamentation. In the absence of specimens of the Indian species referred to above it is not possible to offer any suggestions as to whether any of them should be included in this genus. The suture-lines as drawn by Stoliczka and Kossmat do not show sufficient detail to make any close comparison, but it is probable that they are far less dissected than that of Tainuia. In the three forms Acanthoceras discoidale Koss., Acanthoceras vicinale Stol., and the compressed variety of Acanthoceras rotomagense

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de France in Stol. (non A. gothicum Koss., 28, p. 198, which has a very different suture-line), there is often an absence of the median line of tubercles. This, however, must not be regarded as too important a difference. Kossmat, for instance, says, “das Fehlen der siphonalen Knotenreihe bilden ebenfalls keinen durchgreifenden Unterscheid.”

A fragment of a specimen of the only species was sent to Professor Kilian, who notes, “une forme adulte [of Gunnarites antarcticus] prenant outre les crénelures des côtes, des tubercles siphonaux produisant une convergence vers le type Acanthoceras” (55, p. 175). I find myself unable to adopt this view.

This genus is certainly similar in some respects—so far, at any rate, as ornamentation is concerned—to Mortoniceras (Meek), a genus which is diagnosed by de Grossouvre as follows: “Ce sont en général des coquilles dont le section des tours est plus élevé que large de forme subquadrangu-laire, à bord externe large et ornée sur la ligne siphonale d'une petite quille arrondie très peu saillante. La cloison rassemble par certains traits à celle des Acanthoceras sauf que le premier lobe latéral est arrondi à son extrémité et termine par des digitations avant tout à peu prês la même valeur de sorte que l'on n'y distingue pas, ou tout ou moins très peu nettement la fourche terminale caractéristique des Acanthoceras et des Stohczkaia.”

Whilst in general form not far distant from Mortoniceras, there is no keel, but a well-defined row of ventral tubercles. However, von Heupen (52, pp. 42, 43) states that in Peroniceras and Acanthoceras a keel may be replaced by a row of tubercles. Thus this distinction falls to the ground. In this genus there are five constrictions in a whorl. The suture-line, however, is much more divided than that of Mortoniceras as figured by Spath and von Heupen. The median saddle, in particular, is frilled. The first lateral lobe is bifid, not forked. There are three auxiliary saddles, and the suture-line falls backward considerably from the umbilicus. On the other hand, the suture-line is very similar to that drawn by Crick of Mortoniceras soutoni (Baily) (53, pl. 20, fig. 4). Spath says of this, “The suture-line of an example of M. soutoni Baily, very close to Baily's type, with small umbilicus and comparatively smooth outer whorl, is given for comparison, since it differs from that figured by Woods and from the original drawing by Baily.”

“Tainui” is the name of one of the canoes in which the ancestors of the Maori navigated the Pacific Ocean to New Zealand.

Tainuia aucklandica n. sp. (Plate 24, fig. 2; Plate 34, fig. 3; Plate 46, figs. 1–3.)


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A. B. C. D.
Diameter 195 100 125 100 121 100 210 100
Height of last whorl 71 36 43 34 42 35 90 43
Width of last whorl 56 29 34 27 35 29 72 34
Umbilicus 74 38 42 34 42 35 78 37
E. F. G. H.
Diameter 65 100 134 100 139 100 76 100
Height of last whorl 46 66 49 60 43 32 42
Width of last whorl 41 38 28 38 27 31 41
Umbilicus 26 24 18 38 27 26 34

A, B, Tainuia aucklandica from Bull's Point, Kaipara, N.Z.; C, Tainuia aucklandica from Whangaroa, N.Z.; D, Acanthoceras pseudo-deveiianum Jimbo (22, p. 32); E, Ammonites compressus Stol. (11, p. 66, pl. 34, fig. 5); F, Ammonites vicinale Stol. (11, p. 84, pl. 44; 28, p. 200, Taf. 25, fig. 2); G, Acanthroceras discoidale Koss. (28, p. 201, Taf. 25, fig. 1); H, Acanthroceras gothicum Koss. (28, p. 198, Taf. 25, fig. 3).

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It will at once be seen from the table given above that Tainuia aucklandica in its dimensions comes quite close to Acanthoceras pseudo-deverianum Jimbo, and among the Indian species it comes closest to Acanthoceras discoidale Koss.

Shell large, discoid, with whorls much higher than wide. Involution about five-eighths. Umbilical wall steep, but sides soon rounded off into flanks, which are almost flat and slope gradually into a curve passing rather sharply over periphery. The thickest part is about one-third of the height from umbilicus.

Ornamentation: The shell is strongly ribbed—about twenty-one in each half-revolution. Most of ribs commence as moderate rather lamellar tubercles at top of umbilical wall, and pass over flanks almost radially or with a slight forward inclination, and over periphery with a distinct forward curve. There are four knots on each rib between umbilical tubercle and siphuncle, which increase gradually and uniformly in size towards siphuncle. The first is situated at about one-third of height from umbilical tubercle; the others divide the remaining space about equally, but are not quite regular in position. On median line there is an additional row of tubercles. Between each pair of ribs which arise from tubercles there is always one interstitial rib, which commences just below first tubercle of flank and has tubercles similar to those on ordinary ribs. Six constrictions in a revolution, and each of these has a strong bolster before and behind, relatively slightly tuberculated. The posterior of these intersects one interstitial rib. Some indistinct striations between ribs on body-chamber, which is rather more than half a whorl in size.

Suture-line much dissected. Median saddle somewhat frilled, and a great deal shorter than external saddle, which is conspicuously bifid. First lateral lobe very deep and conspicuously bifid, though the two sides are not symmetrical. Second lateral lobe not nearly so deep, and auxiliary lobes small and a good deal inclined. The other saddles as far as second auxiliary are also bifid. Four auxiliary saddles only are present. Taking a broad view, it may be said that the suture-line is much more complicated than that of Acanthoceras; the first lateral lobe is much deeper and of more importance than in Madrasites, but has these qualities in a less degree than Maorites. The external lobe is not so deep as in Parapachydiscus, and there is a more decided slope towards the umbilicus. The first lateral lobe is distinctly wanting in the forked form that is so characteristic of Acanthoceras and Mortoniceras, to which genera the external form of Tainuia shows a good deal of resemblance, though the umbilicus is noticeably smaller and the periphery is far more rounded.

Three specimens of this form have been found: the type comes from Whangaroa; the other two from Bull's Point, Kaipara Harbour. The type is in the Auckland Museum.

Parapachydiscus Hyatt emend Spath (54, p. 122).

Genotype: P. gollevillensis D'Orb.

I have been unable to find a diagnosis of the characteristics of this genus, and give the following short abstract of de Grossouvre's description of the genotype (19, p. 214, pl. 29, fig. 10; pl. 31, fig. 9):—

Shell discoid compressed, involution more than one-half. Umbilicus of moderate size with vertical walls. Whorls much higher than wide, and

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in the young forms quite smooth. Umbilical ribs few (nine in a whorl). Ribs of peripheral region short, and become less prominent in the more mature individuals. Suture-line much the same as in Pachydiscus. Median saddle rather frilled. External lobe not quite so deep as first lateral lobe, which is symmetrical. Saddles nearly symmetrically divided by a deep secondary lobe. Frilling of suture-line extremely complex and intricate.

Spath in a critical survey of this genus establishes a large number of subgenera, but it seems that this species should be included in his Parapachydiscus sensu stricto (54, p. 122).

All the species of this genus are found in the Campanian or the Maestrictian.

Parapachydiscus rogeri n. sp. (Plate 25, fig. 2; Plate 47, figs. 1, 2.)



Parapachydiscus aff. wittekindi Sch., in Spath (53, p. 229, pl. 24, fig. 1).


Parapachydiscus tweenianus Stol. (11, pl. 54, fig. 3).


Parapachydiscus tweenianus Stol., in Koss. (28, p. 102).

The specimens of this large species are unfortunately so imperfect that approximate measurements only can be given. The most complete specimen represents perhaps only one-eighth part of a complete individual, but it measures 350 mm. by 300 mm. by 120 mm. and weighs about 40 kilograms. This fragment is entirely septate, and it is probable that the complete individual with the body-chamber would be as much as 774 mm. in diameter. Involution considerable, but the proportion cannot be stated. Flanks gently rounded, the curve gradually increasing as the periphery is approached. Lower part of umbilical wall is almost vertical, but afterwards slopes off gradually to periphery. The surface has large low rounded ribs. In the portion nearest to periphery that is shown in the specimens the distance from the crest of one rib to that of the next is 46 mm. The ribs arise at base of umbilical wall and are at first directed backwards at a sharp angle, but they soon bend, and at about one-third of flank they become radial and maintain this direction as far as can be seen. Here and there a faint striation can be seen parallel to the ribs. The shell matter close to umbilicus is as much as 5 mm. in thickness.

Suture-line: Unfortunately the greater part of the specimen is considerably eroded, and it is not possible to see the details of the external lobe or of the external saddle. First lateral lobe deeper than external lobe and nearly symmetrical. Second lateral lobe also deep, but first auxiliary lobe is far shallower; beyond that the suture-line cannot be seen. Secondary lobe in saddles extends rather less than half-way to base, and the two divisions into which it splits the saddle are not quite equal. In external saddle exterior half is larger than interior, but in first lateral saddle the reverse is the case. Both saddles and lobes are complexly divided.

Ornamentation resembles that of the species from Zululand compared by Spath with P. wittekindi Schluter (53, p. 229, pl. 24, fig. 1). In Parapachydiscus rogeri, however, the ribs are carried right down into umbilicus with a strong forward sweep, whereas in the Zululand species they disappear on the edge of umbilical slope. Suture-line of this species is not very different from that of P. quiriquinae Phil. (26, p. 74, pl. 6, fig. 3 a, b, text-fig. 5), though it cannot be seen whether the inner whorl is

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tuberculate as in that species, which also has more numerous ribs, that do not decrease much towards periphery. It resembles P. tweenianus Stol. (11, p. 107, pl. 54) more closely than any of the other of the Indian species. P. tweenianus is said by Kossmat to be similar to a species, apparently unnamed, from Vancouver (28, p. 122).

Only two fragments of this species have been found, both at Bull's Point, one at each end of the exposure of Cretaceous rocks at that place.

Named in honour of the finder, Roger T. Marshall.

Nowakites Spath (54, p. 124).

Genotype: N. carezi de Grossouvre.

The following diagnosis is drawn from de Grossouvre's description of the genotype (19, p. 190):—

Shell discoid compressed. Spire formed of involute whorls which increase rather slowly in height and width. Flanks slightly convex, and external border rounded with a small umbilicus. Ribs very slightly oblique, leaving the umbilical tubercles in pairs. Between the pairs there are usually one or two auxiliary ribs. Occasionally a transverse constriction, wide but not deep. Mature individuals show shorter and shorter auxiliary ribs. Suture-line not known. Occurs in the Coniacian of Corbières.

The genus Nowakites is a division of the genus Parapachydiscus of Hyatt.

Nowakites denticulatus n. sp. (Plate 25, fig. 3; Plate 38, figs. 5, 6.)

Shell in too imperfect a condition to allow of measurements. Shell of moderate size, with aperture a good deal higher than wide. Numerous ribs with high sharp crests and wide rounded intervals. The sharp crests are finely denticulated, but it is only where the state of preservation of the shell is particularly good that this feature can be seen. Ribs arise in pairs from tubercles on edge of umbilicus, but between every pair are one or two interstitial ribs. All of these pass straight over periphery without any noticeable curvature, except where the specimen has been slightly distorted by pressure. Periodic constrictions indistinct.

Only a small portion of suture-line can be seen. It is distinctly of the Parapachydiscoid type, complex and much divided, and with a deep straight first lateral lobe.

The sharp costation and fine denticulations are the dominant features of this species, which in general respects somewhat resembles Kossmaticeras antarcticum var. bhavaniformis Kilian and Reboul from Seymour Island (46, p. 33, pl. 15, fig. 2). I am tempted to compare it with Koss-maticeras (Grossouvrites) gemmatum Hupe from Quiriquina, a specimen of which has been recorded by Trechmann (56, p. 387) from the Selwyn Rapids, in the South Island.

In virtue of its ornamentation, form, and suture-line, K. gemmatum ought, I think, to be placed in the genus Parapachydiscus. Kilian and Reboul remark that it approaches very close to Pachydiscus in the character of the suture-line (46, p. 23, also p. 42). So far as European species are concerned, I am inclined to say that P. sayni de Gross. (19, p. 181, pl. 39, fig. 2) comes nearest to the present species.

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One specimen only has been collected. It comes from the northern side of Bull's Point, Kaipara Harbour, close to the spot where Parapachydiscus rogeri, Ancanthoceras ultimum, and Gaudryceras subsacya were obtained.

Hauericeras de Grossouvre, 1893.

The genus is described by its author as follows: Shells with a large umbilicus. Whorls thin and high, with a keel on the margin. Flanks slightly convex or even flat, without sculpture but showing constrictions more or less clearly. The suture-line is quite analogous to that of Desmoceras, Puzosia, and Pachydiscus. The first lateral lobe is as long as the external lobe (19, p. 219).

Kossmat says that Hauericeras is separated from Puzosia by the sharp external keel, but remarks later that in a young example of Hauericeras rembda the keel is not developed, and that the same is true of young examples of H. gardeni. There is the same development of auxiliary lobes as in Puzosia, which has the external lobe always distinctly shorter than the first lateral lobe, and the first lateral lobe and external saddle are inclined. In Hauericeras the first lateral lobe is the same length or only slightly longer than the external lobe. The external saddle is straight and not regularly bifid (28, p. 122).

Van Hoepen says that in the South African examples the first lateral lobe is distinctly longer than the external lobe. In Puzosia planulata the external saddle reaches farther forward than the first lateral, but in Hauericeras the reverse is the case. He points out that in the smaller whorls the umbilical surface is perpendicular to the plane of symmetry, while in the older whorls it is inclined (52, p. 27).

Yokoyama says that in the Japanese example the lobes are quite irregularly trifid and not so deep as in the Indian example. This asymmetry is noticeable in the South African example (52, text-fig. 5) and in the Indian specimens (11, pl. 33, fig. 4), but not in the Nanaimo example figured by Kossmat (28, Taf. 18, fig. 10).

Hauericeras ngapuhi n. sp. (Plate 24, fig. 5; Plate 43, fig. 3; Plate 45, fig. 3.



Ammonites gardeni Baily, Quart. Journ. Geol. Soc., vol. 11, p. 456, pl. 9, fig. 3.


Ammonites gardeni Baily, in Stoliczka (11, p. 61, pl. 33, fig. 4)


Ammonites gardeni Baily. in Whiteaves (7, p. 102).


Desmoceras gardeni Baily, in Yokoyama (17, p. 184, pl. 20, fig. 10).


Hauericeras gardeni Baily, in Woods (40, p. 332).


Desmoceras (Hauericeras) gardeni Baily, in Kilian and Reboul (46, p. 18).


Hauericeras gardeni Baily, in Spath (53, p. 238, text-fig. 12).


Hauericeras gardeni Baily, in Spath (59, p. 50, table).


Hauericeras gardeni Baily, in van Hoepen (52, p. 27, text-fig. 15).


Hauericeras gardeni Baily, in Spath (54, p. 129).

The dimensions given below show that the species is closely related to the widely-occurring species Hauericeras gardeni Baily, which has been recorded from so many countries that lie on the border of the Pacific Ocean. At the same time, the differences are certainly great enough to be

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of specific value. In all the three specimens found there is no keel, but this may be due to their immaturity.


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A. B. C. D. E. F. G.
Diameter 22.2 100 23.5 100 27 100 120 100 26 100 100 79 100
Height of last whorl 9 40 10 42 12 44 30 10 38 35 32 40
Width of last whorl 6.8 30 7 30 8 30 5.75 22 19
Umbilicus 7 31 7 30 8 30 42 10 38 39 27 37

A, B, C. three specimens of Hauericeras ngapuhi from Whangaroa, N.Z.; D, Ammonites durga Forbes = Puzosia compressa Koss. (11, p. 143); E, Hauericeras gardeni Spath (54, p. 130); F, means of measurements of thirty specimens of Hauericeras gardeni by Crick, Spath (54, p. 129): G, Hauericeras welschi de Grossouvre (19, pl. 35, fig. 9).

Shell small, discoid and much compressed. Whorls high and narrow. Umbilical wall nearly vertical, with a sharp angle to the almost flat flank. Periphery evenly rounded, and involution rather more than a third. Umbilicus of moderate size.

Ornamentation: The surface of shell quite smooth except for conspicuous constrictions, which number six in last whorl. They commence at bottom of umbilicus and, directed strongly forward, pass in a straight line to periphery, over which they pass in a sharp forward curve.

Suture-line not very highly divided. First lateral saddle a good deal higher than the others, and auxiliary saddles fall away posteriorly to a distinct umbilical lobe. There are in all eight saddles. First lateral lobe deeper than external lobe and symmetrical. Second lateral lobe straight but much shorter, and auxiliary lobes much inclined. Saddles are rather more regularly bifid than is usual in this genus.

The shape and ornamentation of the three specimens have caused me to place the species in this genus. Unfortunately the shell matter is not preserved on the periphery, and consequently no keel is to be seen—probably it was not developed; though, as pointed out, the small size of the specimens and their probable immaturity may account for this. The table of dimensions shows that Hauericeras ngapuhi has a higher whorl than the related species, but at the same time it is wider and has a smaller umbilicus. As shown in the table on pages 19697, Hauericeras gardeni has a very wide occurrence in the countries that border the North Pacific.

Three specimens, one from Bull's Point and two from Whangaroa.

The Ngapuhi is a famous Maori tribe of New Zealand.

Schluteria de Grossouvre.

This genus was established by de Grossouvre to include some species of the Upper Chalk closely allied to Phylloceras by their form and ornamentation, but their suture-line clearly approaches that of Puzosia and Pachydiscus, and differs entirely from that of Phylloceras in the absence of the large foliaceous terminations (19, p. 216). De Grossouvre, however, as Spath has pointed out (53, p. 46), included in this genus certain species of Phylloceras related to P. nera. Spath has therefore emended the genus, and has taken as the genotype Schluteria larteti Seunes.

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Schluteria rarawa n. sp. (Plate 19, fig. 10; Plate 32, figs. 7, 8.)



Ammonites diphylloides Forbes, Trans. Geol. Soc., ser. 2, vol. 7, p. 105, pl. 8, fig. 8.


A. diphylloides Stol. (11, p. 119, pl. 59, figs. 8, 9).


Desmoceras diphylloides Koss. (28, p. 109, Taf. 19, figs. 8 a, b, c, 9 a, b, c)


[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]

A. B. C.
Diameter 11 100 22 100 34.6 100
Height of last whorl 6 55 11 50 21.1 58
Width of last whorl 6 55 11 50 16.3 47
Umbilicus 4 18 2.8 08

A, Schluteria rarawa, Kaipara Harbour, N.Z.; B, Schluteria (Desmocerds) inanis Koss. (28, p. 107); C, Schluteria (Desmoceras) crassa van Hoepen (52, p. 21).

Shell small, completely involute, with whorls rapidly increasing. Wall of umbilicus sloping steeply at first but soon rounding off into the flanks, which maintain the same curve. The whole aperture, therefore, is nearly circular.

Ornamentation: Very fine ribs can be distinguished on umbilical slope, the specimens being extremely well preserved. About every fifth is stronger than the others. They curve sharply backward first, then forward, but they disappear about half-way up the flanks. A strong lens is required to make them visible, as to the naked eye the surface appears smooth and polished. Six constrictions in a whorl, which are shallow and bordered behind by a low but wide bolster rib.

Suture-line with nine saddles decreasing gradually and uniformly from periphery to umbilicus. Exterior and first lateral saddle bifid. Lobes symmetrically bifid and much wider than saddles. Lobes uniformly of same depth, and the whole suture-line is linearly radial in direction.

Schluteria simplex from Pondoland has a narrower whorl than this species, but Schluteria crassa from the same country comes nearer to it in form. The species certainly comes close to Desmoceras loryi from Seymour Island, described by Kilian and Reboul (46, p. 18, pl. 1, figs. 4, 5), though all the specimens that have been found in New Zealand are much smaller. This species is much more inflated than S. diphylloides Forbes and than S. inane Forbes. The suture-line is spaced out like that of S. phyllimorphum Koss., but the form and size are quite different. The suture-line is not unlike that of S. diphylloides and S. inains.

A specimen in the collection of the Geological Survey of New Zealand, which was found by McKay near Awanui, on the east coast of the North Island, comes very close to this species, but it is badly preserved. Several specimens have been obtained from Batley and Bull's Point.

Importance of Suture-lines.

The suture-lines that are represented in Plates 1925 have been drawn with great care, and are thought to be nearly exact, though the projection on to a flat surface offers difficulty, and certainly gives rise to some error. As far as practicable, mechanical methods were employed, and errors in proportions are thereby greatly reduced. It is recognized that too much reliance cannot be placed on the details of the form of the suture-line for purposes of identification: ornamentation and proportions of the shell must have full consideration.

Suture-lines of the following species:—

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Fig. 1.—Baculites rectus n. sp. Bull's Point.
Fig. 2.—Ptychoceras zelandicum n. sp. Whangaroa.
Fig. 3.—Diplomoceras wakanene n. sp. Bull's Point.
Fig. 4.—Phylloceras nera Forbes. H. 19; W. 9. Batley.
Fig. 5.—Phylloceras bistriatum n. sp. H. 19; W. 19. Bull's Point.
Fig. 5a.—Internal suture-line.
Fig. 6.—Phylloceras forbesianum d'Orb. H. 17; W. 16.5. Batley.
Fig. 7.—Phylloceras radiatum n. sp. H. 22; W. 15. Bull's Point.
Fig. 7a.—Internal suture-line.
Fig. 8.—Phylloceras minimum n. sp. H. 10.5; W. 9.
Fig. 8a.—Internal suture-line.
Fig. 9.—Zelandites kaiparaensis n. sp. H. 7.6; W. 6. Bull's Point.
Fig. 9a.—Internal suture-line.
Fig. 10.—Schluteria rarawa n. sp. H. 4.75; W. 5.25. Batley.

Suture-lines of the following species:—

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Fig. 1.—Pseudophyllites indra Forbes. H. 22; W. 21. Bull's Point.
Fig. 2.—Pseudophyllites whangaroaensis n. sp. H. 16; W. 18. Whangaroa.
Fig. 3.—Gaudryceras politissimum Koss. H. 9; W. 7.5. Bull's Point.
Fig. 4.—Gaudryceras propemite n. sp. H. 7; W. 9.
Fig. 4a.—Internal portion of suture-line. Bull's Point.
Fig. 5.—Tetragonites margaritatus n. sp. H. 5.5; W. 6.25. Batley.
Fig. 6.—Tetragonites latus n. sp. H. 6.25; W. 10. Batley.
Fig. 6a.—Internal portion of suture-line. H. 4; W. 8.
Fig. 7.—Gaudryceras particostatum n. sp. H. 7; W. 6.5. Bull's Point.
Fig. 7a.—Internal portion of suture-line. H. 8; W. 8.
Fig. 8.—Gaudryceras subsacya n. sp. H. 5; W. 7.5.
Fig. 8a.—Internal portion of suture-line. H. 7; W. 9.
Fig. 9.—Vertebrites murdochi n. sp. H. 5; W. 10. Hokianga.
Fig. 9a.—Internal portion of suture-line.
Fig. 10.—Gaudryceras crenatum n. sp. H. 4.75; W. 6.5. Bull's Point.
Fig. 11.—Tetragonites simplex n. sp. H. 5.75; W. 5.5. Batley.
Fig. 11a.—Internal portion of suture-line. H. 3; W. 3.

Suture-lines of the following species:—

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Fig. 1.—Jacobites whangaroaensis n. sp. H. 44; W. 35. Whangaroa.
Fig. 2.—Jacobites angularis n. sp. H. 21; W. 22.5. Bull's Point.
Fig. 3.—Jacobites anderssoni K. & R. H. 12; W. 11.
Fig. 3a.—Internal portion of suture-line. H. 5; W. 5.
Fig. 4.—Neomadrasites nodulosus n. sp. H. 7.5; W. 9.25. Bull's Point.
Fig. 4a.—Internal portion of suture-line.
Fig. 5.—Jacobites minimum n. sp. H. 3; W. 2.2. Batley.
Fig. 6.—Madrasites multicostatus n. sp. H. 9.5; W. 10. Bull's Point.
Fig. 7.—Madrasites regularis n. sp. H. 10; W. 10. Bull's Point.
Fig. 8.—Gunnarites nordenskjoldi K. & R. H. 5; W. 5. Batley.
Fig. 9.—Brahmaites rotundus n. sp. H. 5; W. 8. Batley.
Fig. 10.—Tetragoniles tetragonus Koss. H. 16; W. 17. Bull's Point.
Fig. 11.—Pseudophyllites whangaroaensis n. sp. Internal portion of suture-line. H. 6.5; W. 7.5. Whangaroa.
Fig. 12.—Madrasites fortior n. sp. Whangaroa.

Suture-lines of the following species:—

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Fig. 1.—Gunnarites inflatus K. & R. H. 38; W. 32. Batley.
Fig. 1a.—Internal portion of suture-line H. 35; W. 31. Bull's Point.
Fig. 1b.—Suture-line of juvenile form. Bull's Point.
Fig. 2.—Gunnarites zelandicus Marshall. H. 24; W. 22. Specimen somewhat eroded.
Fig. 3.—Gunnarites antarcticus Stuart Weller. H. 35; W. 27. Batley.
Fig. 4.—Internal suture-line of Jacobites angulans n. sp. H. 21; W. 25.
Fig. 5.—Puzosia angusta n. sp. H. 25; W. 9.

Suture-lines of the following species:—

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Fig. 1.—Maorites tenuicostatus Marshall. H. 51; W. 35.5. Batley.
Fig. 1a.—Internal portion of suture-line. H. 39; W. 35.
Fig. 2.—Jacobites waitapuensis n. sp. H. 34; W. 30. Whangaroa.
Fig. 3.—Maorites suturalis n. sp. H. 26; W. 17. Bull's Point.

Suture-lines of the following species:—

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Fig. 1.—Maorites densicostatus K. & R. H. 21; W. 13. Bull's Point
Fig. 2.—Tainuia aucklandica n. sp. H. 43; W. 34. Whangaroa.
Fig. 3.—Parapuzosia brevicostata n. sp. Whangaroa.
Fig. 4.—Parapuzosia ordinaria n. sp. H. 8; W. 6. Batley.
Fig. 5.—Hauericeras ngapuhi n. sp. H. 9; W. 6. Bull's Point.
Fig 5a.—Internal portion of suture-line.

(Through an unfortunate error tubercles are indicated in fig. 3 in place of fig. 1.)

Suture-lines of the following species:—

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Fig. 1.—Acanthoceras ultimum n. sp. Bull's Point.
Fig. 1a.—Internal portion of suture-line. H. 9.
Fig. 2.—Parapachydiscus rogeri n. sp. H. 220; W. 270 (?). Bull's Point.
Fig. 3.—Nowakites denticulatus n. sp. H. 39; W. 38 (?). Bull's Point.

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Fig. 1.—Phylloceras nera Forbes. × 1 ½.
Fig. 2.—Phylloceras nera Forbes: section from periphery to umbilicus.
Fig. 3.—Phylloceras radiatum n. sp. × 1 ½.
Fig. 4.—Phylloceras radiatum n. sp.: section from periphery to umbilicus.
Fig. 5.—Phylloceras minimum n. sp. Bull's Point. × 2 ½.
Fig. 6.—Phylloceras minimum n. sp.: section.

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Fig. 1.—Phylloceras bistriatum n. sp. Bull's Point. × 2.
Fig. 2.—Phylloceras bistriatum n. sp.: section.
Fig. 3.—Phylloceras forbesianum d'Orb. × 2.
Fig. 4.—Phylloceras forbesianum d'Orb.: section.

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Fig. 1.—Gaudryceras politissimum Koss. Batley. × 1 ½.
Fig. 2.—Gaudryceras politissimum Koss.: section.
Fig. 3.—Gaudryceras propemite n. sp. Bull's Point. × 1 ½.
Fig. 4.—Gaudryceras propemite n. sp.: section.

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Fig. 1.—Gaudryceras subsacya n. sp. Bull's Point. × 1 ¼.
Fig. 2.—Gaudryceras subsacya n. sp.: section.
Fig. 3.—Pseudophyllites indra Forbes. × 1 ¼.
Fig. 4.—Pseudophyllites indra Forbes: section.
Fig. 5.—Pseudophyllites indra Forbes: flank, showing suture-line.
Fig. 6.—Tetragonites epigonus Koss. × 1 ½.
Fig. 7.—Tetragonites epigonus Koss.: section.

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Fig. 1.—Vertebrites murdochi n. sp. × 1 ½.
Fig. 2.—Vertebrites murdochi n. sp.: section.
Fig. 3.—Gaudryceras particoslatum n. sp. × 1 ½.
Fig. 4.—Gaudryceras particoslatum n. sp.: section.
Fig. 5.—Tetragonites margaritatus n. sp. × 1 ½.
Fig. 6.—Tetragonites margaritatus n. sp.: section

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Fig. 1.—Zelandites kaiparaensis n. sp. Bull's Point. × 1 ½.
Fig. 2.—Zelandites kaiparaensis n. sp.: section.
Fig. 3.—Gaudryceras crenatum n. sp. Bull's Point. × 1 ½.
Fig. 3a.—Gaudryceras crenatum n. sp.: section.
Fig. 4.—Brahmaites rotundus n. sp. Batley. × 1 ½.
Fig. 5.—Brahmaites rotundus n. sp.: section.
Fig. 6.—Parapuzosia ordinaria n. sp. Batley. × 1 ¼.
Fig. 7.—Parapuzosia ordinaria n. sp.: section.
Fig. 8.—Diplomoceras wakanene n. sp Bull's Point. × 1 ½.
Fig. 9.—Diplomoceras wakanene n. sp.: section.

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Fig. 1.—Tetragonites latus n. sp. Batley. × 1 ½.
Fig. 2.—Tetragonites latus n. sp.: section.
Fig. 3.—Tetragonites simplex n. sp. Bull's Point. × 1 ½.
Fig. 4.—Tetragonites simplex n. sp.: section.
Fig. 5.—Periphery of Pseudophyllites whangaroaensis n. sp., showing sculpture
Fig. 6.—Pseudophyllites whangaroaensis n. sp.: section.
Fig. 7.—Schluteria rarawa n. sp. Batley. × 1 ½.
Fig. 8.—Schluteria rarawa n. sp.: section.
Fig. 9.—Baculites rectus n. sp. Batley. × 1 ¼.
Fig. 10.—Baculites rectus n. sp.: section.
Fig. 11.—Ptychoceras zelandicum n. sp. Bull's Point. × 1 ½.
Fig. 12.—Ptychoceras zelandicum n. sp.: section.

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Oxybeloceras sp. Mangamuka River. Hokianga. × 3/2.

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Fig. 1.—Acanthoceras ultimum n. sp. Bull's Point. × 1 ¼.
Fig. 2.—Acanthoceras ultimum n. sp.: section.
Fig. 3.—Tainuia aucklandica n. sp.: showing development of tubercles. × 1 ¼.

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Fig. 1.—Madrasites multicostatus n. sp. Bull's Point. × 2.
Fig. 2.—Madrasites multicostatus n. sp.: section.
Fig. 3.—Madrasites regularis n. sp. Bull's Point. × 2.
Fig. 4.—Madrasites regularis n. sp.: section.

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Fig. 1.—Neomadrasites nodulosus n. sp. Bull's Point. × 1 ½.
Fig. 2.—Neomadrasites nodulosus n. sp.: section.
Fig. 3.—Neomadrasites nodulosus n. sp.
Fig. 4.—Jacobites angularis n. sp.: Bull's Point. × 1 ½.
Fig. 5.—Jacobites angularis n. sp.: section.
Fig. 6.—Gunnarites nordenskjoldi K. & R. Batley. × 1 ½.

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Fig. 1.—Jacobites whangaroaensis n. sp. Whangaroa. × ½.
Fig. 2.—Jacobites whangaroaensis n. sp.: section.
Fig. 3.—Jacobites anderssoni K. & R. Bull's Point. × 1 ½.
Fig. 4.—Jacobites anderssoni K. & R.: section.

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Fig. 1.—Madrasites sulcatus n. sp. Batley. × 2.
Fig. 2.—Madrasites sulcatus n. sp.: section.
Fig. 3.—Jacobites minimus n. sp. Batley. × 2.
Fig. 4.—Jacobites minimus n. sp.: section.
Fig. 5.—Nowakites denticulatus n. sp. Bull's Point. Nat. size.
Fig. 6.—Nowakites denticulatus n. sp.: showing costation. × 2.

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Fig. 1.—Gunnarites zelandicus Marshall. Batley. × 1 ½.
Fig. 2.—Gunnarites zelandicus Marshall: section.
Fig. 3.—Gunnarites antarcticus Stuart Weller. Bull's Point. × 1 ½.
Fig. 4.—Gunnarites antarcticus Stuart Weller: section.

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Fig. 1.—Gunnarites inflatus K. & R. Batley. Nat. size.
Fig. 2.—Gunnarites inflatus K. & R.: section.
Fig. 3.—Flank of Vertebrites murdochi n. sp., showing change in costation. × 5.

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Fig. 1.—Puzosia angusta n. sp.: showing costation. Bull's Point. × 2.
Fig. 2.—Puzosia angusta n. sp.: section.
Fig. 3.—Madrasites fortior n. sp. Whangaroa. × 2.
Fig. 4.—Madrasites cumshewaensis? Whiteaves. Whangaroa. × 2.

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Fig. 1.—Maorites tenuicostatus Marshall. Batley. Nat. size.
Fig. 2.—Maorites tenuicostatus Marshall (juv.). Bull's Point. × 2.
Fig. 3.—Maorites densicostatus K. & R. (juv.). Bull's Point.
Fig. 4.—Maorites suturalis n. sp. (juv.). Bull's Point. × 2.

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Fig. 1.—Maorites suturalis n. sp. Bull's Point. × 1 ¼.
Fig. 2.—Parapuzosia brevicostata n. sp. Whangaroa. × 2,
Fig. 3.—Hauericeras ngapuhi n. sp. Bull's Point. × 2.

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Fig. 1.—Jacobites waitapuensis n. sp. Whangaroa. × 1 ¼.
Fig. 2.—Maorites densicostatus K. & R. Bull's Point. Nat. size.

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Fig. 1.—Maorites tenuicostatus Marshall: section.
Fig. 2.—Jacobites waitapuensis n. sp.: section.
Fig. 3.—Hauericeras ngapuhi n. sp.: section.
Fig. 4.—Maorites densicostatus K. & R.: section.
Fig. 5.—Maorites suturalis n. sp.: section.
Fig. 6.—Parapuzosia brevicostata n. sp.: section.

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Fig. 1.—Tainuia aucklandica n. sp. Whangaroa. Nat. size.
Fig. 2.—Tainuia aucklandica n. sp.: fragment showing ornamentation. × 1 ½.
Fig. 3.—Taunuia aucklandica n. sp.: section.

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Fig. 1.—Parapachydiscus rogeri n. sp. Bull's Point. × ⅛. Fig. 2.—Parapachydiscus rogeri n. sp.: showing costation. × ⅓.

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So far as this collection is concerned, it has been found that the suture-line in nearly all cases has given clear indication of the generic position that a species should have, though it hardly enables one to distinguish between such genera as Madrasites, Jacobites, and Gunnarites. It is thought that the time that has been taken in drawing these lines in detail has not been wasted.

It seems that little attention has hitherto been given to the nature of the internal portion of the suture-line. In few researches on collections of ammonites have any drawings been made of them. So far as can be ascertained, Kossmat is the only author who has made any use of them for purposes of classification, and he appears to have done so in one instance only—in the distinction between Gaudryceras and Tetragonites.

In this paper drawings of the internal suture-line of several species will be found, and it is suggested that the form of it may be of great use in assigning species to their proper position in doubtful cases. Spath, however, attaches little or no importance to it: “The stretching-out of the auxiliary elements may only be the result of the adaptation of the suture-line to wider sides, and the raising of the umbilical portion is often found in younger developments” (53, p. 239). This, however, does not hold generally: compare, for example, Gaudryceras and Pseudophyllites.

The form of the internal suture-line is little varied in time and in the widely scattered species of a genus. The internal suture-line of a species of Phylloceras from the Dogger (Zittel, Handbuch, vol. 2, fig. 609) differs but little from those of New Zealand specimens from the Upper Cretaceous. The difference is almost restricted to a reduction in the number of saddles. It is certainly striking that the New Zealand species have the same number of saddles as in Sowerbyceras, and the suggestion thus offered that the New Zealand species should be placed in the latter genus is supported by some other details. Further material for study is, however, needed.

The genus Gaudryceras has the same unusual form of this suture-line in species from India, Japan, South Africa, and New Zealand. The importance of this line that is here suggested is supported by the great variety of form that it presents.

A summary of the variations that have been found in these New Zealand specimens is considered worthy of a special statement:—

Phylloceras: Internal saddle club-shaped. Two lateral saddles, the second with an almost spherical termination. No auxiliaries.

Gaudryceras: A single saddle of uniform width, which is typical in G. subsacya. Those species of Gaudryceras, such as G. semileve, which have a single saddle, increasing greatly in width at the base, should perhaps be placed in a separate genus.

Tetragonites: Two well-developed saddles, as in T. epigonus. Such a species as T. simplex has an additional saddle.

Pseudophyllites: Two saddles, the external one with much greater development than the first lateral.

Vertebrites: Five or six saddles, gradually decreasing in size towards the umbilicus.

Zelandites: A single saddle, wide at the base.

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Jacobites and Gunnarites both have two large saddles, which in the latter genus are much dissected; the antisiphonal lobe is very deep. This form appears to indicate a descent from Acanthoceras.

Madrasites, so far as the New Zealand specimens are concerned, has an internal suture-line but little different from Jacobites, and thus differs much from that figured by Kossmat.

The suture-line of Pachydiscus as drawn by Yabe is quite different from those of the genera mentioned, but it could not be seen in any of the New Zealand specimens.

Maorites, Puzosia, and Hauericeras have the same type of internal suture-line. The lateral saddles and the auxiliaries are situated on the outer slope of the internal saddle, and thus give the appearance of a single complex much-divided saddle. This type of internal suture-line appears as far back as Dalmasiceras, and is continued in Perisphinctes, and is supposed to indicate a relationship to the Hoplitid stock. It is much regretted that the material that was available did not allow of further study.

Geographical Relationship.

In order to make it possible to draw any geographical conclusions from the distribution of the species of ammonites that have been found in New Zealand it is necessary to summarize the facts that have already been recorded in regard to their occurrence in different circum-Pacific countries.

The following percentages of the ammonite fauna of the various countries are represented in New Zealand by identical or closely allied species:—

Number of Species. Number of Related Species in N.Z. Percentage of Species in N.Z. Percentage of N.Z. Species in each.
Aryalur 53 18 34 34
Trichinopoly 26 4 15.5 8
Utatur 89 11 12.5 20
Seymour Island 41 16 39 31
South Patagonia 20 2 10 4
Quiriquina 10 5 50 10
Chico 52 7 13 14
Nanaimo 17 2 11 14
Queen Charlotte Island 27 7 8 4
Japan 47 10 21 19
New Caledonia 12 4 33 8
Pondoland 54 12 22 22
Madagascar 49 5 10 10

It is unfortunate that the Upper Cretaceous ammonites of New Caledonia have not been more fully collected and identified, though Piroutet has done excellent work. The fauna that has been collected by him suggests

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that it is rather similar to that of New Zealand, but in the meantime this statement is somewhat hazardous.

As shown in this table, the Seymour Island fauna has remarkable similarities with that of New Zealand, and the conclusion is suggested that there was a continuous coast-line between these countries. This is the belief of Wilckens and others. On the other hand, the Patagonian fauna contains the Hoplites-Placenticeras elements that are absent from New Zealand, Antarctica, and Chile, and in this last country the small fauna that has been discovered shows a closer resemblance to that of New Zealand than any other that has been recorded. No country between Chile and California has yet yielded an Upper Cretaceous ammonite fauna, but in the latter there is the well-known Chico formation. In the lower series of this formation there is a fairly rich fauua, but it is not closely related to that of New Zealand. In the Upper Chico series there are fewer ammonites, and they show a closer resemblance than those of the lower series.

In British Columbia the Queen Charlotte Islands have yielded a fauna that has little resemblance to the one described in this paper, but it is probably of Middle Cretaceous age. On the other hand, the Nanaimo beds are of the Upper Cretaceous period, but still the species that have been obtained from them show little similarity, on the whole, to the New Zealand species.

In the highest Upper Cretaceous ammonite-bearing strata of South India—that is, the Aryalur—a close relationship with New Zealand is again displayed; but in the beds below them—the Trichinopoly series of Lower Senonian age—the resemblance is much less pronounced, and the specific resemblances are fewer. The Utatur beds, which are considered to be of Cenomanian age, have also several species that are closely related to New Zealand forms. The amount of resemblance is, however, decidedly less in the lower than in the upper series.

I have no access to a complete statement of the species that have been found in Madagascar. The works of Spath, Woods, and van Hoepen give full information of the ammonite fauna of Pondoland. It is rather remarkable that these beds contain a fauna that is much more closely related to that of New Zealand than is the fauna of the beds of the same age at Umzamba Hill, in Zululand.

The general geographical impression that is produced is, as might be expected, that the ammonite fauna of this Upper Cretaceous age in the countries closer to New Zealand is more closely related than that of the countries that are more remote. Patagonia is, of course, a striking exception to the general application of this statement, while the species that have been found in Chile and New Caledonia are too few to allow of a satisfactory comparison.

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Table showing the Occurrence of Representative
South Africa. Madagascar. India (Aryalur and Valudayur). India (Trichinopoly). India (Utatur).
1 Phylloceras nera Forbes P. woodsi (Pondoland) P. nera
2 Phylloceras radiatum n. sp. P. umsambiense P. velledae (Cenomanian) P. forbesianum
3 Phylloceras forbesianuim d'Orb. P. forbesianum (Cenomanian) P. forbesianum
4 Phylloceras minimum n. sp. P. whiteavesi
5 Phylloceras bistriatum n. sp.
6 Vertebrites murdochi n. sp. G. kayei
7 Gaudryceras propemite n. sp. * G. cintum (Pondoland) G. multiplexum (Coniacian) G. multiplenum
8 Gaudryceras particostatum n. sp. G. varicostatum (Pondoland)
9 Gaudryceras subsacya n. sp. G. sacya
10 Gaudryceras politissimum Koss. G. politissimum G. politissimum
11 Tetragonites epigonus Koss T. virgulatus (Pondoland) T. epigonus (Cenomanian) T. epigonus
12 Tetragonites latus n. sp. T. superstes (Pondoland) T. timotheanus
13 Tetragonites simplex n. sp. T. nuperus T. cala
14 Tetragonites margaritatus n. sp. T. sigcau
15 Pseudophyllites indra Forbes Ps. indra Ps. indra Ps. indra
16 Baculites rectus n. sp. (B. vagina)
17 Diplomoceras wakanene n. sp. D. rugatum
18 Ptychoceras zelandicum n. sp. Pt. sipho
19 Acanthoceras ultimum n. sp. Ac. newboldi (Cenomanian) Ac. newboldi
20 Madrasites regularis n. sp. (M. africanus) (M. muravia-turensis)
21 Madrasites multicostatus n. sp. M. buddhaicus
22 Gunnarites antarcticus St. Wel. G. kalika
23 Gunnarites zelandicus Mar. G. kalika
24 Gunnarites inflatus K. & R. G. kalika
25 Gunnarites nordenskjoldi K. & R.
26 Jacobites anderssoni K. & R.
27 Jacobites whangaroaensis n. sp.
28 Jacobites waitapuensis n. sp.
29 Brahmaites rotundus n. sp. Brahmaites sp. Brahmaites brahma
30 Maorites tenuicostatus Mar. ? M. bhavani
31 Maorites densicostatus K. & R. M. aemilianus
32 Maorites suturalis n. sp. M. kaudi
33 Puzosia angusta n. sp. F. compressa (Cenomanian) P. compressa
34 Parapuzosia brevicostata n. sp. P. gaudama (Coniacian) P. gaudama
35 Parapuzosia ordinaria n. sp. P. bhima
36 Tainuia aucklandica n. sp. Ac. vicinale (Cenomanian) Acanthoceras vicinale
37 Parapachydiscus rogeri n. sp. P. aff. wittelkindi P. tweenianus P. tweenianus
38 Hauericeras ngapuhi n. sp. H. gardeni H. gardeni H. gardeni
39 Schluteria rarawa n. sp. S. compactum S. diphylloides
40 Grossouirites gemmatus Huppé §
41 Madrasites haumuriensis Hect. M. bhavani
42 Turrilites circumtaeniatus Koss. T. circumtaeniatus
Number of species in fauna 54 49 53 26 89
Number of allied species in New Zealand 12 5 18 4 11

[Footnote] * Represented in Europe by G. Mite, Lower Senonian.

[Footnote] † Represented in Europe by Ac. rotomagense, Cenomanian.

[Footnote] ‡ Represented in Europe by P. octosulcata, Senonian.

[Footnote] § Recorded by Trechmann, Geol. Mag, dec. 6, vol 4, p. 338, 1917.

[Footnote] ∥ Recorded by Woods, N. Z. Geol. Surv. Pal. Bull. No 4, 1917.

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Forms of the New Zealand Ammonites.
Antarctica, Seymour Island, Snow Hill Island. South Patagonia. Chile (Quiriquina). California. Vancouver. Japan. New Caledonia.
P. nera P. nera (P. surya) 1
P. ramosum P. ramosum P. ramosum P. velledae P. velledae 2
P. whiteavèsi 4
P. esoense 5
G. kayei G. kayei G. kayei (Mt. Diablo) G. kayei 6
G. multiplexum? 7
G. tenuiliratum 8
G. sacya G. sacya G. sacya, G. tenuiliratum var. ornatum 9
? Gaudryceras 10
T. epigonus T. epigonus T. aft. epigonus Tetragonites sp. 11
T. timotheanus T. timotheanus T. glabrus 12
Ps. indra Ps. indra Ps. indra 15
B. chicoensis B. chicoensis ? Baculites 16
? Anisoceras 17
Pt. glaber 18
Ac. turneri (Mt. Diablo) Ac. rotomagense 19
? M. cumshewaense ? M. cumshewaense ? M. cumshewaense 21
G. antarcticus 22
G. antarcticus 23
G. inflatus 24
G. nordenskioldi 25
J. anderssoni 26
? J. anderssoni 27
? M. bhavani var. seymouriana 28
? M. bhavani 29
? M. bhavani var. densicostatus 30
? Puzosia sp. P. darwini P. gaudama ? P. gaudama 34
Ac. pseudodeverianum 36
P. quiriquinae Pachydiscus sp. P. haradai 37
H. gardeni H. gardeni H. gardeni 38
S. loryi 39
G. gemmatus G. gemmatus 40
41 20 10 52 27 + 17 47 12
16 2 5 7 2 7 10 4 ?

The following New Zealand species have no representatives in other countries: Gaudryceras crenatum n. sp., Oxybeloceras sp., Madrasites sulcatus n. sp., M. fortior n. sp., Jacobites angularis n. sp., J. minimus n. sp., Neomadrasites nodulosus, Parapachydiscus denticulatus n. sp., Pseudophyllites whangaroaensis. The following additional species have been recorded from New Zealand: (1) by Woods, Gaudryceras sacya Forbes (? G. subsacya), G. baculities cf. vagina (? B. rectus); (2) by Spath (53, p. 299), Tetragonites sp. (? T. simplex), Kossmaticeras bhavani Stol., K. cumshewaense (?Whiteaves) K. & R., K. (Gunnarites) bhavaniforme K. & R.

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Geological Age Of Circum-Pacific Cretaceous Ammonite Horizons.

Whether all these faunas are strictly of the same age may well be doubted. Spath, for instance, says, “The chances then are that a fauna like that of Pondoland which does not quite agree with (e.g.) either the Upper Campanian of Galicia or the Valudayur of southern India, if homogeneous at all, may belong to a hitherto unrecognized horizon or horizons. Since non-sequences are possible, these horizons are, perhaps, not even consecutive” (54, p. 116). Paulcke also, in speaking of the Patagonian ammonites, says, “We find ourselves in this dilemma: Either we have to deal with Lower Senonian strata containing Pachydiscus which shows Upper Senonian characters, or we must suppose that the stratification is wrongly interpreted” (42A, p. 73). Wilckens, who described the lamellibranchs that were collected in the same beds as the ammonites, classes the strata that contain Pachydiscus in the Cenomanian or Turonian age, and thus supports the first alternative. (Lamell. der oberen Kreide Sud-Pat., 1907, p. 60).

Kilian and Reboul, in their work on the ammonites of Seymour Island and Snow Hill, distinguish three different horizons, extending from Cenomanian to the Upper Senonian. Wilckens, however, in his work on the lamellibranchs and gasteropods of the same region, is emphatic that the collections from all of the localities indicate the same age, though he excepts from this statement the localities 2 and 6 of Snow Hill, whence nothing but ammonites was obtained. Wilckens concludes, “Ebenso wie die ubrige molluskan Fauna einheitlich ist. Sie ist obersenonisch. Die Hill Schichten für alter als die alteren Seymour Schichten zu halten liegt kein Grund vor; beide sind obersenonen.”

It is thus apparent that those who have closely examined the faunas from Pondoland, Seymour Island, and Patagonia are at least in grave doubt about their actual age, or, at any rate, differ considerably in their opinions on this question. There is even doubt whether in any locality the strata represent more than one division of the Upper Cretaceous. A tabulated statement of the opinions that have been expressed in regard to the faunas that have been described from the Indo-Pacific countries will make this statement clearer.

[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]

Pondoland Upper Campanian-Maestrictian Spath, 54, p. 116.
Upper Santonian van Hoepen, 52, p. 45.
Senonian Woods, 40, p. 345.
Zululand Campanian-Maestrictian Spath, 53, p. 223.
Seymour Island and Snow Hill Cenomanian—Upper Senonian Kilian and Reboul, 46, p. 58.
Upper Senonian Wilckens, 47B, p. 114.
South Patagonia Upper Senonian Paulcke, 42A, pp 72, 73.
Cenomanian—Upper Senonian Wilckens, 39. pp. 57—58.
Quiriquina Upper Senonian Steinmann, 26, p. 27.
Upper Senonian Wilckens, 38, p. 284.
California Upper Chico = Lower Senonian Anderson, 33, p. 62.
Lower Chico = Turonian Anderson, 33, p. 62.
Nanaimo ? Whiteaves.
Lower Senonian Anderson, 33, p. 62.
Japan Cenomanian ? Yokoyama, 17, p. 170.
Middle Cretaceous Jimbo, 22, p. 11.
Diff. Upper Cretaceous divisions Yabe, 37, p. 5.
Senonian Bohm, in Yabe, 37, p. 4.
Aryalur and Valudayur Upper Senonian Kossmat, 28, p. 102.
Trichinopoly Lower Senonian—Upper Turonian Kossmat, 28, p. 102.
Utatur Cenomanian—Lower Turonian Kossmat, 28, p. 102.
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In the general account in his Geology, however, Haug classes all those strata that contain the important fossils Gaudryceras kayei and Pseudo-phyllites indra in the Maestrictian formation. These strata are in general those that have been classed as Senonian or Santonian or Maestrictian by various authors. This point at least is apparent: All those strata that contain the two fossils named above are in all probability of the same age; and, since those authorities that are familiar with the species consider that they are closely related to Gaudryceras planorbiforme and Gaudryceras colloti, it becomes equally certain that this formation is very high in the Cretaceous succession, perhaps even in the Maestrictian. Included in these are the Aryalur, Valudayur, Nanaimo, Lower Chico, Quiriquina, Seymour Island, New Zealand, and Pondoland Upper Cretaceous formations. This has naturally led to wide generalizations, but it is perhaps right to call attention to the present state of our knowledge in regard to the distribution of such animals as ammonites. Information is, of course, meagre and indefinite, as we know nothing of the larval period, the swimming-powers of the animals, or their requirements as regards food or temperature. It appears, however, to be certain that Baculites and Ptychoceras were mud-borers in shallow water, and in this case, unless the larval period was long, they must have required a continuous shore-line with considerable areas of sheltered water in order to have become so widely distributed. Diplomoceras, also, must have had the most limited power of movement on the sea-floor. Species of other genera, such as Pachydiscus, perhaps had habits not very different from those of the present Nautilus, and, given suitable food and temperature-conditions, may have wandered rapidly along a variety of coasts.

There appears to be a general belief that the lytoceratids, in part at least, were stenothermal. Thus Spath speaks of the stenothermal genera Phylloceras and Lytoceras (53, p. 53). He also says that the Pondoland and Zululand faunas are connected by the presence in both of Pseudo-schoenbachia—probably an active swimmer—and of benthonic crawlers and mud-boring Baculites, the stenothermal Lytoceratidae being dependent upon deeper water or warmer currents. The oxycone developments of the Upper Senonian, that might have been thought to be active swimmers, are often curiously restricted, and, like other marine organisms, may have a limited horizontal distribution. He notes, however, that Buckman differs, and thinks that it is wrong to generalize from our knowledge of the habits of Recent forms in regard to extinct groups.

Haug states on this subject, “La presénce du genre Phylloceras et les Lytoceratidae au Japon, dans la Colombie Britannique, en Californie, au Chile, en Patagonie, et dans la Terre de Graham montre avec evidence que l'on n'est pas dans tous ces pays en présence de formations littorales mais qu'il s'agit de formations bathyales, déposées dans un géosynclinal étroit ou s'accumulaient de grandes épaisseurs de dépot” (Haug, Traité de Géologie, p. 1349). He therefore includes much of the area of New Zealand in his “géosynclinal circumpacifique” (p. 1359).

If Baculites and Ptychoceras were mud-borers and Diplomoceras was a benthonic crawler, it is clear that in Upper Cretaceous times the coast-line round the Pacific was nearly continuous, if we are justified in assuming that these genera had a comparatively brief larval period. Since it has already been recorded that species of Baculites showing an Upper Cretaceous character occur in India, Pondoland, Chile, California, British Columbia, New Zealand, and New Caledonia, it is unnecessary to imagine that great swimming-powers were possessed by Phylloceras or the Lytoceratidae in order to account for the presence of species of these genera in the New Zealand fauna of that period.

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The same conclusion is arrived at when the nature of the strata and the fossils embedded in them are considered. The strata certainly give evidence of deposition near a coast-line, for they are often sandy, and sometimes even pebbly and conglomeratic. Fragments of wood and leaves of several species of plants are not uncommon in most of the localities. Such fossils are, of course, most frequently preserved in strata that have been deposited in sheltered waters into which some rivers flow—certainly not in those deposited on an open coast-line nor on the floor of a geosynclinal area. As Phylloceras and species of Lytoceratidae are found in association with them, there is no reason to suppose that the presence of species of those groups is, in this case at least, any evidence of the existence of bathyal conditions in the New Zealand area.

The occurrence of identical or very similar species of Gaudryceras and Phylloceras in countries that now have such different climatic conditions as India, British Columbia, Chile, Patagonia, Seymour Island, and New Zealand, as well as Algeria and western Europe, appears at first sight to support the idea that these genera were stenothermal. However, it is found that closely related species of Puzosids, Pachydiscus, Baculites, Madrasites, &c., have an almost equally wide occurrence, and this term based on such an occurrence would be as applicable to them. It seems perhaps more reasonable to suppose that in Upper Cretaceous times the ammonites as a family were stenothermal, or that climatic zones were far less defined and extreme, and that temperature conditions over the world were far more uniform.

It may be positively stated that none of the formations in which Cretaceous fossils have yet been obtained in New Zealand give any evidence that they have been deposited in any deep synclinorium—even though the formation that in so many localities rests on the fossiliferous rocks of Cretaceous age is a greensand, succeeded by a forminiferal (mainly Globigerina) limestone sometimes siliceous and containing diatoms and radiolaria that may be of deep-water origin. Reasons have been given elsewhere, however, which seem to prove that this limestone is of Eocene age. It is a striking fact that similar material of wide occurrence in New Caledonia has lately been shown to be of Eocene age by A. Heim and A. Jeannet (Bull. Geol. Soc. Geol. de France, 1922, pp. 246—53).

It is noticeable that, with one or two exceptions, such as Baculites chicoensis in California and B. reetus in New Zealand, there is no evidence of a closer relationship of the New Zealand species of ammonites with those of distant lands than with those of nearer countries. In other words, it appears that the New Zealand species are more closely related to those that have been described from countries close to New Zealand than to those more distant, though exception must always be made of Patagonia. This general relation seems to indicate that the circum-Pacific coast-line was much the same in its position in Upper Cretaceous times as it is at the present day. In making this statement, however, it must be borne in mind that the Seymour Island and Chilian coast-lines were certainly not separated from New Zealand by such wide oceanic stretches as now. It is thought by Wilckens that there was a connection between these lands, either continuous or slightly interrupted, a good deal to the north of the present coast-line of the Antarctic Continent. So far as the ammonites of these three countries are concerned, it is almost certain that, though the species found in them are closely similar to those found in India, they are still more closely related to one another. The faunas differ from those of India in the absence of

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species of such general and important genera as Schloenbachia, Scaphites (one species of Scaphites in Quiriquina, Chile), Placenticeras, &c., as well as in having such an unusual development of Madrasites, Jacobites, and Gunnarites.

As to the westward development of New Zealand in Upper Cretaceous times, we have at present no information. Australia seems to have no marine sediments of that age, and it was presumably above the sea-level at that time. Though this, too, was probably the case with the country on the west coast of the South Island of New Zealand, there appears to be no reason to believe that the basin of the Tasman Sea was then a land area, as has been lately suggested by Benson.

The ammonite fauna of Pondoland has been shown by Spath to have some distinct Indo-Pacific characteristics, as well as others of a distinctly Atlantic nature, in the presence of Schloenbachia, Mortoniceras, Sphenodiscus, and Lenticeras. The absence of these important genera in the faunas of Seymour Island and New Zealand seems to me to indicate that South Africa derived its Indo-Pacific elements directly from India along the coast of the western Indian Ocean, and that there was no association with the countries at the south of the Pacific Ocean.

It appears that the New Zealand ammonite fauna of Upper Cretaceous age is of such a nature that it indicates that the general position of the Pacific coast-line was much the same as at the present time. New Zealand was, however, more continuously united with Seymour Island and with India, though we do not at present possess enough information to show us exactly where this coastal connection extended.

Stratigraphical Features of the Batley Series.

The stratification of the Upper Cretaceous rocks of North Auckland is extremely difficult, if not impossible, to unravel with the amount of information at present at my disposal. At Batley there is practically no outcrop of the rock that has not suffered from superficial slipping. The same is true of the occurrence at Whangaroa; and at both of these places the small extent of the fossiliferous beach shows that the actual exposure of the Cretaceous sediments is small. At Bull's Point the stratification is almost vertical, and the thickness of the formation exposed is perhaps 300 ft. Here again, however, surface slipping and soil covering allow very little of the stratification to be seen. In all three localities no fossils have been found actually in situ in the strata. In nearly all cases they have been extracted from concretionary boulders, often of large size, lying on the foreshore. In a few cases only they have been extracted from the boulders by natural agencies, and were lying in the small rock-fragments on the beach.

It has already been stated that much of the fossiliferous rock is sandy or even pebbly, and it is therefore unlikely that the total thickness that is exposed represents much lapse of time. It is not reasonable to suppose that the interval from the Cenomanian to the Maestrictian should be represented by such a small thickness of sediments, which appear to have been deposited rather quickly. It was, of course, hoped and expected that definite species of ammonites would be found to be restricted to certain stratified divisions, and thus make it possible to define exact zones in the stratigraphical sequence. However, as the work proceeded and the collection became more extensive, it was found that no species of ammonites could be regarded as restricted in their occurrence to any particular portion of the beach.

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The question then arose as to whether this apparent mixture of zones could be due to the transport of material along the beach by wave-action. Full consideration of the conditions that actually exist, however, show that wave-transport must be small. If, for instance, the locality of Bull's Point be taken—for in this place the conditions would favour wave-transport more than elsewhere—the following facts must be noted :—

By far the greater number of the specimens were found on the side of the exposure extending west to east for about 100 metres on the north side of Bull's Point. At almost the same spot on the beach were found the following species: Acanthoceras ultimum, Baculites rectus, Gaudryceras subsacya, Gaudryceras particostatum, Gunnarites inflatus, Tainuia aucklandica, Gaudryceras propemite, Maorites tenuicostatus, Phylloceras radiatum, Phylloceras forbesianum, Puzosia angusta, Parapachydiscus rogeri, Nowakites denticulatus, Zelandites murdochi, Pseudophyllites indra, Tetragonites epigonus, Tetragonites latus, Ptychoceras zelandicum.

Several of these species appear to have distinct affinities with species from the Indian Utatur formation, which is, of course, definitely correlated with the Cenomanian. In this respect the following are most important: Tainuia aucklandica, Acanthoceras ultimum, Phylloceras radiatum, Puzosia angusta, Gaudryceras subsacya. Of these the species of Acanthoceras is particularly noticeable; for in the Indian ammonite fauna, which has 167 species, there are twenty-four of Acanthoceras, all of which were found in the Utatur series. It is obvious, therefore, that among the specimens found on the northern side of Bull's Point there is an important Utatur element, and it is certain that if any association of the species named had been found alone it would have been necessary to assign the formation to the Upper Cenomanian age. Associated with these species, however, as is shown in the list given above, there is a very distinct element of the highest ammonite horizon, for Haug has taken the species Pseudophyllites indra and Gaudryceras kayei (of the latter of which Vertebrites murdochi is merely a local representative) as typical of the Maestrictian in all those countries from which they have been recorded. In addition, Diplomoceras wakanene, Ptychoceras zelandicum, and Baculites rectus are indicative of a high horizon. The species of Gunnarites, Madrasites, and Jacobites also belong to an horizon distinctly higher than the Utatur. Thus on the north side of Bull's Point alone species are found that, judging by the Indian succession, would indicate that a great lapse of time is represented by the strata in a very small thickness of rock.

It must be added that a similar association of species has been found in each of the main collecting localities—viz., Bull's Point, Batley, and Whangaroa—though, since the collections made at both of the last two localities are much smaller than that made at the other, the variety of types that has been found is distinctly less. At Batley Pseudophyllites indra, Vertebrites murdochi, Gunnarites inflatus, Maorites tenuicostatus, Baculites rectus, Tetragonites epigonus, Tetragonites latus, Ptychoceras zelandicum, Phylloceras forbesianum, and Gaudryceras subsacya are relatively common. All of these are found at Bull's Point, though Zelandites murdochi was represented by one specimen only. At Whangaroa Vertebrites murdochi and Ptychoceras zelandicum are far more common than elsewhere, and perhaps Hauericeras ngapuhi also. On the other hand, Pseudophyllites indra is less common, as is Baculites rectus. Phylloceras forbesianum occurs, as well as Maorites tenuicostatus. It is thus clear that a similar association of ammonites occurs in each of the localities. The same is true also of the lamellibranchs and gasteropods, though it has not yet been found possible

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to study them fully. Lastly, a species of Araucarites has been found quite commonly at each of the localities, represented by fragments of wood as well as leaves.

Geological Age of the Batley Series.

It is proposed to call this the Batley series, after the locality where these northern New Zealand ammonites were first found. The similarity of the faunas shows that the same horizon or horizons are represented in each of the localities, and it is extremely unlikely that in such a small outcrop as that which is found at Batley and at Whangaroa there should be the same association of higher and lower strata as at Bull's Point, where the outcrop is a good deal larger. It therefore appears to be probable that either the association of species is different from any that has been found in India, or else this New Zealand horizon is intermediate in age between the Utatur and Aryalur horizons of India, but different from the Trichinopoly horizon. Any suggestion of correlation with the Trichinopoly horizon appears to be discounted by the occurrence of such forms as Pseudophyllites indra, Vertebrites murdochi, and Diplomoceras ngapuhi, all of which—or, rather, their local representatives—are restricted to the highest of the Indian ammonite horizons.

If it is agreed that all of these fossils are derived from a small range of horizons, we encounter a great deal of difficulty in correlating the New Zealand formation with equivalents elsewhere. It is necessary to take the Indian series as a guide. The table given on page 194 shows of the fifty-three Aryalur and Valudayur species there are eighteen representatives in New Zealand—that is, a percentage of 34; of the Trichinopoly fauna of twenty-six species there are four representatives, a percentage of 15.5; while of the large Utatur fauna of eighty-nine species there are only eleven representatives, a percentage of only 12.5. It is, then, clear that as regards the Indian ammonites the greatest affinity of this northern New Zealand Cretaceous fauna is with the Aryalur.

The same result may be arrived at in a different manner. If it is admitted that one horizon alone is represented, it is clear that it must contain either Utatur survivals or Aryalur antecedents. Even granting that the isolation of New Zealand was not nearly so extreme in the Upper Cretaceous times as now, it still seems that the former alternative is more probable than the latter.

It is therefore concluded that the Upper Cretaceous rocks of New Zealand are the equivalent of the Aryalur and Valudayur of India. All authorities are agreed in correlating this with a very high horizon in the Cretaceous of Europe. Kossmat takes the Upper Senonian; Spath for probably equivalent formations in Pondoland prefers the Upper Campanian plus Maestrictian, though van Hoepen regards these beds as Santonian. The Upper Senonian, however, is accepted by Kilian and Reboul, and generally by all of those who have described any series of Upper Cretaceous ammonites in the Indian Pacific region. On the other hand, Haug has adopted the Maestrictian equivalent for the Aryalur, and has correlated most of the circum-Pacific ammonite faunas with it. He appears to take as the criterion for this age the occurrence of the species Pseudophyllites indra and Gaudryceras kayei. According to this criterion, the Kaipara and Whangaroa ammonite fauna should be placed in the Maestrictian. My own opinion is that this is rather too high an horizon, and I prefer to take the Upper Santonian or the Lower Campanian as the more probable European equivalent.

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Comparison with other Cretaceous Circum-Pacific Districts.


As a basis for comparison, Kossmat's classification of the Indian formations is taken, and is quoted below. It will be noticed that there are 158 species in the Indian fauna, while there are only fifty-three in New Zealand.

[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]

Trichinopoly. Pondicherry.
Danian c. Ninnyur. c. Nerinaea beds.
Upper Senonian Aryalur Group, 53 spp. b. Kulmodu. b. Trigonoarca beds.
a. Otacod, Aryalur, Karapady. a. Anisocei as beds =Valudayur (Blan.)
Lower Senonian b. Upper Trichinopoly.
Turonian Trich. Group, 26 spp. a. Lower Trichinopoly.
c. Cunum beds.
Cenomanian Utatur Group, 89 spp. b. Acanthoceras beds.
a. Muraviatur, Odium, Utatur.
Table showing the Generic Arrangement of Species in India and New Zealand.
Phylloceras. Gaudryceras. Tetragonites. Pseudophyllites. Turrilites Hamites (D'plomoceras, &c.) Baculites. Neoptychites. Placenticeras. Splenodiscus. Ferbesiceras. Schloenbachia. Sch. subgenera. Stoliczkaia. Acanthoceras. Olcostepl anus. Scaphites Holcodiscus Brahmaites. Pachydiscus Desmoceras Puzosia. Hauericeras.
Aryalur 4 3 1 1 0 9 2 0 0 1 0 0 1 0 0 0 2 8 2 11 3 0 2
Trichinopoly 0 2 1 0 1 0 1 0 1 0 0 0 2 0 0 0 4 7 0 5 1 2 0
Utatur 4 8 2 0 6 9 4 2 1 0 2 10 0 4 28 2 3 3 0 1 3 8 0
New Zealand 5 8 4 2 1 3 1 0 0 0 0 0 0 0 1 0 0 12 1 2 1 2 1

In this table Hamites includes Diplomoceras, Ptychoceras, and Oxybeloceras; while Holcodiscus includes Madrasites, Gunnarites, Jacobites, and Maorites, the last for comparative purposes only, without any suggestion as to affinities.

This table shows that the species which have been found in New Zealand belong in general to those genera which in India have a considerable range in time, and actually, with two exceptions, are represented in the formation which is correlated with the Upper Senonian in Europe. The exceptions are found in Pseudophyllites, Turrilites, Puzosia, and Acanthoceras. The only species of Turrilites that has been found in New Zealand came from the Clarence Valley, in the South Island. The two species of Puzosia are perhaps survivals, and this is more remarkable in regard to Acanthoceras ultimum, which is closely related to the well-known Acanthoceras rotomagense.

It is noticeable that twenty-eight species of this genus have been found in the Utatur formation of India. Turrilites also is represented by several species in India. The third of these genera—Puzosia—has species in the Senonian of Pondoland and Chile, the Lower Chico of California, and the Urakawa of Japan.

The species that it is considered do not belong to previously established genera are included in the following new genera: Maorites, probably a development of Dalmasiceras or of Sonneratia, a Hoplitid genus, and

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probably related to some species that have been placed in Puzosia, a Desmoceratid genus; Tainuia, related to Acanthoceras vicinale; and Neomadrasites, derived from Madrasites; also Vertebrites and Zelandites, related to Gaudryceras. In the opinion of the writer these genera indicate a younger rather than an older horizon.

The outstanding features of this ammonite fauna when compared with that of India are most notably the complete absence of Schloenbachia and Scaphites. Another noticeable feature is the abundance of Gunnarites and Jacobites, both of which genera are included under Holcodiscus in the preceding table, which is based on the study of the Indian ammonites by Kossmat. It is also apparent that the genera Phylloceras are Gaudryceras are well represented. If the three species of Maorites are considered as closely related to the genus Puzosia it is clear that it has an unusual representation.

So far as a general statement can be made of the generic features, one is justified in saying that the fauna follows very closely on the main features of the Indian fauna. When the species are compared it is found that there is a considerable number of the Indian species that are extremely close to some in New Zealand. The following species appear to be almost, if not quite, identical: Phylloceras nera, Phylloceras forbesianum, Vertebrites murdochi cf. Gaudryceras kayei, Gaudryceras politissimum, Gaudryceras subsacya, Pseudophyllites indra, Tetragonites epigonum, Hauericeras ngapuhi cf. gardeni, Gunnarites antarcticum cf. Holcodiscus kalika, Brahmaites rotundum, Ptychoceras zelandicum, and Diplomoceras wakanene. This great similarity seems to indicate either that a coast-line was almost continuous from India to New Zealand, or that the species had remarkable powers of dispersion. In connection with this it is well to remember that there seems to be a general agreement that the Phylloceratidae and Lytoceratidae had such free-swimming powers that they were distributed by ocean currents. It is, however, almost impossible that such relatively fixed forms as Baculites and Ptychoceras could have had such means of distribution.


This fauna appears to be much less closely related to the ammonite fauna of Japan than to that of India. As in New Zealand, no species of Schoenbachia have as yet been recorded from Japan. Madrasites, however, is not well represented, but species of Jacobites, Gunnarites, and Baculites have not yet been discovered. On the other hand, the species of Phylloceras, Gaudryceras, and Tetragonites show a good deal of resemblance to New Zealand forms, though it may be said that the species of the two last genera, having little ornamentation, have not so much opportunity of displaying striking differences.

The numerous Japanese species of Desmoceras constitute a striking feature, and it seems as though there are no forms in Japan that show analogy with the species of Maorites. The genera Placenticeras, Olcostephanus, Crioceras, and Scaphites, which are absent from the New Zealand fauna, have been found in Japan, as well as a great variety of Turrilites, which, however, come from the lower horizons. The widely distributed species Pseudophyllites indra and Gaudryceras kayei appear to be unrepresented in Japan, while the Gaudryceras sacya of Yokoyama and Jimbo is referred by Yabe to a new species, G. tenuiliratum.

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The Chico formation is well known to include at least one horizon which is universally admitted to be of Senonian age. Anderson's list of the Upper Chico (Lower Senonian) fauna on page 27 includes twelve species of ammonites only. Of these, eight belong to the genera Ancyloceras, Helicoceras, Schloenbachia, and Hoplites, none of which genera are represented in the New Zealand fauna as at present known. Of the remainder, Baculites chicoensis is the only species which shows close relationship to the New Zealand forms. The small number of species does not allow a comparison of any great value to be made. The Lower Chico, however, which is of Turonian age, has a much larger and more representative fauna, and some forty-four species of ammonites are included in it. Among these are twelve species of Schloenbachia and six of Scaphites, which again are genera unknown at present in New Zealand. Of five species only can it be said that there is a definite relationship.

At Mount Diablo there is a fossil-bearing formation, the exact age of which, and relation to the other formations, has not yet been satisfactorily settled. This horizon is of special interest, for it contains Gaudryceras kayei and Acanthoceras turneri, both of which are rather close to New Zealand species of ammonites.

From these statements it would seem that the Californian fauna had certain affinities with the Atlantic fauna that were not shared by New Zealand. Schloenbachia, though so common in Europe and Africa, and extending to India, is nevertheless absent from the Upper Senonian fauna of many circum-Pacific countries. Mortoniceras also is represented in California, though absent from all other Indo-Pacific faunas. (This genus has lately been recorded from Japan.)

British Columbia.

In this country Whiteaves has described two different series of Cretaceous ammonites. The lower of these two formations was found in the Queen Charlotte Islands, and the fossils contained in them are considered by Haug to be indicative of the Albian or the base of the Cenomanian. However, Gaudryceras sacya, Tetragonites timotheanus, and perhaps Hoplites beudanti, show some affinities with members of the New Zealand ammonites. On the other hand, the presence of the genera Schloenbachia, Sphenodiscus, Olcostephanus, and Ancyloceras establish a complete difference from the New Zealand fauna, as well as from that of Seymour Island and of Quiriquina, and perhaps also from that of South Patagonia.

The Nanaimo fauna, however, which is considered by Haug to be of Maestrictian age, and by other authorities to be of Upper Cretaceous age, shows many points of resemblance to the present fauna. The presence of such species as Pseudophyllites indra, Gaudryceras kayei, Baculites chicoensis, Phylloceras velledae, and Hauericeras gardeni at once establishes a relationship that must be regarded as a close one.


Paulcke has identified twenty species from South Patagonia. These display a considerable difference from those of Seymour Island, Chile, New Zealand, and Japan, especially in the presence of a large series of Hoplites plasticus sp. This series Paulcke himself and Haug compare with North European species, and it thus appears that South Patagonia may have had an Atlantic connection that did not extend to the other

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countries mentioned. The other Patagonian genera are all represented in New Zealand and in other circum-Pacific countries. Pachydiscus and Baculites have four and three species respectively, but Gunnarites and Jacobites, which are common in Seymour Island and New Zealand, have no Patagonian species. Again, there are no species of Puzosia, Hauericeras, or Desmoceras; and it is noticeable that Vertebrites murdochi has no representative species, though it has such a wide Indo-Pacific occurrence.

The species of Hoplites give this fauna a most distinctive character when compared with the other South Pacific ammonite faunas. In other respects, however, all the elements of the fauna might well be found amongst those species that have been found in the north of Auckland.


The only locality from which ammonites have been recorded is the island of Quiriquina, in the Bay of Concepcion. Nine species only have been recorded. All the genera, with the exception of Scaphites, are represented in New Zealand. The species that have been recorded show a considerable resemblance to New Zealand species. Though this is a small collection, it seems to show a closer affinity with New Zealand species than that of any other country except Seymour Island.

Seymour Island.

The New Zealand ammonites show rather a close relationship to those described by Kilian and Reboul from Seymour Island, a point to which Kilian has already drawn attention. It is in the genera Madrasites, Gunnarites, Jacobites, and perhaps Maorites, that this is most markedly shown, for these genera have a fuller representation in both countries than in any other. The species of Phylloceras, Gaudryceras, and Tetra-gonites are also rather similar, but perhaps no more so than in the ammonite faunas of the other circum-Pacific countries where Upper Cretaceous ammonites have been described. On the other hand, the absence of Baculites and Ptcychoceras, especially the former, is rather surprising. On the whole, as previously remarked and as shown in the tables on previous pages, the Seymour Island fauna is more closely related to New Zealand than that of any other country.

Fossils Found With The Ammonites.

Remains of a large number of organisms have been found in association with the ammonites.

The plant-remains are described by Mr. W. N. Edwards in the present volume (pages 121128).

An echinoid was sent to Dr. Bather; he forwarded it to Professor Hawkins, who was good enough to examine it, and he remarks that it is probably a true Hemiaster.

Dr. Smith Woodward was good enough to examine the fish-scales, and detected some that were characteristic of Cladocyclus, a fish of the Cretaceous family Ichthyodectidae.

Two species of brachiopods were sent to Dr. J. A. Thomson, who forwarded them to England, but no report has yet been received.

There are about twelve species of gasteropods, which include a form close to Amberleya spinigera Wilckens, from Seymour Island.

The lamellibranchs include a species of Thyasira close to T. townsendi Wilckins, from Seymour Island. It is hoped to classify the species of gasteropods and lamellibranchs next year.

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