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Volume 82, 1954-55
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Fossil Leaves, Fruits and Seeds from the Wanganui Series (Plio-Pleistocene) of New Zealand.

[Read by title before Wellington Branch, March 30, 1954; received by Editor, March 31, 1954.]

Summary

Previous systematic work on New Zealand plant macrofossils from the Wanganui Series is reviewed Seven new species are described: Podocarpites tolleyi, Pipei erewhonensis, Rumex pachyperianthus, Proteoides ohukaensis. Betulites couperi. Nothofagus searellae, Phyllites kaiwakaensis. Oliver's Ormond flora has been revised, the number of species present is reduced from 31 to 26, and the number of extinct species from 18 to 9.

Introduction

The only plant macrofossils so far described from the Plio-Pleistocene Wanganui Series in New Zealand are fossil leaves from the Waipaoa series at Ormond, Poverty Say, described by Oliver (1928).

Two fossil fruits were described by Berry as Pliocene in age, but the beds containing them are now considered to be Miocene by Couper (1952; 346). Oliver (1936) described a flora from Kaikorai, considering it lower Pliocene in age and listed similar leaves from Great Barrier Island (in Couper, 1953); both these localities are considered Miocene (Couper, 1953).

During the past three years the writer has studied collections of fossil leaves, fruits and seeds obtained by the New Zealand Geological Survey from the Wanganui Series in the North Island, from Pokeno in the north to Halcombe in the south. (See list p. 4.)

The present paper contains descriptions of the known extinct species from these collections and a revision of the species described by Oliver (1928) from Ormond. Seven new species are described from beds ranging from Opoitian to Castlecliffian in age. The many Recent species in these collections have been listed by Couper and McQueen (1954).

Material

The types of new species are in the macropaleobotanical collection of the New Zealand Geological Survey, Department of Scientific and Industrial Research, Wellington.

The list below gives in the following order: N. Z. Geological Survey Register number, sheet fossil number, collectors and year, grid references, lithology, and stratigraphic position. Molluscs from the N133 and N124 One-Mile Sheet districts were identified and ages determined by Mr. O. P. Olson, N.Z. Geological Survey.

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B163: N133/534 “Erewhon” Station, probably same as N133/509. J. Park 1886. ? 349339.*. Yellow brown micaceous siltstone, moderately hard.

B195: N98/503 Ormond, Waipaoa Valley, H. Hill, M. Ongley 1910–1916. 320510. Creamy yellow tuffaceous siltstone, soft. Castlecliffian (Oliver, 1928: 287).

B206, 209, 215: N124/501, 503, 510 Esk Valley, on powerline road, E. of Te Pohue P. R. and D. R. McQueen, 1950. 214649. Light yellow brown siltstone, soft. Interbedded with Nukumaruan molluscs.

B226: N135/506 Rangitawa Stm. Halcombe P. R. and D. R. McQueen, 1951. 965625. Carbonaceous grey sandy siltstone, very soft Interbedded with Upper Castlecliffian molluscs (Te Punga, 1952·23).

B228: N133/522 Rangitikei R. road cutting 1/2 mile S. of Matawhero Rd. bridge P. R. and D. R. McQueen, A. P. Druce, 1951. 351331.* Yellow brown micaceous siltstone, moderately hard. Conformably beneath Opoitean molluscs.

B230: N133/509 Rangitikei R 1 mile downstream from Inland Patea Road Bridge. P. R. and D. R. McQueen, A P. Druce, 1951. 349339.*Yellow brown micaceous siltstone, moderately hard. 1–200ft. conformably below Opoitean molluscs.

B249: N47/514 Railway cutting south of Pokeno Station. D. Kear and D. R. McQueen. 1952. 549122 Brown carbonaceous mudstone, soft Castlecliffian (Couper and McQueen, 1954).

B323: N55/522 Mouth of Ohuka Ck Raglan Co D. Kear, D. R. McQueen, 1952 277788. Purple brown pumiceous siltstone, soft, ? Nukumaruan (Couper and McQueen, 1954).

B328: N52/562. Cutting on present main trunk railway, about 1 mile north of Whangamarino Railway Station. P. R. and D. R. McQueen, D. Kear, 1952 Light brown siltstone, soft. ? Lower Wanganui Series.

Systematics

Family Podocarpaceae

Genus Podocarpites Andrae

Podocarpites tolleyi n.sp (Text-fig. 1, Figs. 1–3)

Length (incomplete specimen) 8.5 mm.

Diameter (on plane of dehiscence) 6 mm.

Seed ovoid, apex acuminate, base mammiliform. Two longitudinal ridges diametrically opposite along junctions of two halves, ridges more pronounced at apical end. Surface coarsely longitudinally striate Apparently dehiscent into two equal halves along ridges Structure of walls along dehiscence three layers, an outer, hard, longitudinally striate thin integument (0.2 mm. thick), a thick middle softer palisade layer (0.8 mm. thick), and a hard, smooth surfaced interior integument (0.1 mm. thick) Inner integument attenuated into the apex and leading to micropyle.

Locality. Rangitawa Stm. B226/8, 128, 129, Paratypes.

Age. Upper Castlecliffian (Pleistocene).

[Footnote] * Grid references in terms of Lands and Survey Department Map of the Ruahine Ranges (1951).

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Text-Fig 1.—New Species of Fossil Seeds from the Wanganui Series. 1—Podocarpites tolleyi n.sp external view of one-half (B226/128) × 10. 2—Podocarpites tolleyi n.sp. apical view (B226/129) × 10. 3—Podocarpites tolleyi n.sp. section along dehiscence (B226/130) × 10.4—Rumex pachyperianthus n.sp. (B226/131) × 25. 5—Rumex pachyperianthus n. sp. apical view (B226/132) × 25. 6—Proteoides ohukaensis n.sp. external view (B323/31) × 13. 7—Proteoides ohukaensis n.sp. partial section (B323/21) × 13

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This seed differs from all known living New Zealand species; the general morphology, wall structure and presence of a micropyle suggests inclusion in the family Podocarpaceae. As all Recent Podocarpaceae have not been examined this new species is placed in the form genus Podocarpites. The specific name is an acknowledgment to Mrs. W. P. Tolley, who drew the illustration on Text-fig. 1, and who first noticed the micropyle.

Family Lauraceae

Genus Beilschmiedia Nees

Beilschmiedia ovata W. R. B. Oliver.

1928. Beilschmiedia ovata W. R. B. Oliver, Trans. N.Z. Inst. 59: 293, Fig. 6.

1928. Parsonsia obtusa W. R. B. Oliver, Trans. N. Z. Inst. 59: 293, Fig. 20.

Localities. Ormond B195/18, 195/31 (Holotype here designated); Pokeno B249/18,/19 Rangitikei River B228/5,/6 Whangamarino B328/31.

Age. Opoitean-Castlecliffian (Pliocene-Pleistocene).

The leaf Oliver described as the type specimen of Parsonsia obtusa matches his Beilschmiedia ovata in shape, venation and tertiary reticulation. It is slightly smaller and the secondaries are closer than in the type of B. ovata; it is possible this leaf may have come from an exposed portion of a tree.

Family Piperaceae

Genus Piper L.

Piper erewhonensis n.sp. (Pl. 27, Fig. 1)

40 mm. long, 27 mm. wide. Lower and middle sections of a probably ovate leaf. Margin entire. Surface flat with lightly impressed veins. Midrib straight 0.7 mm. wide at base. Venation camptodromous. Secondary veins alternate, arising singly, except for basal two on each side which leave the midrib at the same point, the lower at 90°, the upper at 60°. Upper secondaries leaving midrib at 40° and curving towards apex. Major secondaries branching 1/3 way to apex. Tertiary venation consisting of prominent intersecondaries, with fine interconnecting venules.

Locality. “Erewhon” Station B136/30 Holotype.

Age. Opoitean (Pliocene).

This general form of venation is found in the New Zealand Macropiper excelsum Miq. but the shape of the leaf agrees far more with specimens of Piper (Yuncker, 1952). The species name refers to the locality name used by Park (1886), taken from the name of the sheep station at the Rangitikei River.

Family Polygonaceae

Genus Rumex L.

Rumex pachyperianthus n.sp. (Text-fig. 1, Figs. 4, 5.)

Outer portion of fruit, irregularly ovoid, base cuneate, apex acuminate, trigonous, faces flat to convex. Shortly pedunculate and dehiscing along angles, dehiscence extending three-quarters of length of fruit. Surface irregularly cristate. Wall 0.2 mm thick, amorphous in structure Interior smooth.

Measurements of twenty specimens:

Average Range Holotype (B226/131)
Length 2.0 mm 1 5–2.2 mm. 2.2 mm.
Greatest Diameter 1.5 mm. 1.4–1.8 mm. 1.5 mm.
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Plate 1—Retouched photographs of fossil leaves from the Wanganui Series 1—Piper crenhou cusis nsp Holotype B163/30 × 2 2—Belulites couperi n.sp Holotyupe B163/9 × 2 3—Nothofagus searellae nsp B209/30 × 4 4—Nothotagus searellae nsp Holotype B206/7 × 4 5—Notholagus of Chifor fusea B206/2 × 3 6—Phyllires lamahsis nsp Holotype B209/1 × 3.

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Locality. Rangitawa Stm. B226/132–150 Paratypes.

This portion of a fruit is closest to the enlarged perianths of the Polygonaceae, and is similar to those of Rumex It is thicker than any named New Zealand species.

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Text-Fig. 2 —Knightia fossilis Oliver. Sketch of B195/29 showing dentation (in solid line) exposed by further clearing. The outline used as a basis for Oliver's description is here shown dotted. (X 0 75). del W. P. T.

Family Proteaceae

Genus Knightia R. Br.

Knightia excelsa R. Br. (Text-fig. 2)

1810. Knightia excelsa R. Br. in Trans. Linn. Soc. × 194, t 2.

1925. —— Cheesem. Man. N.Z. Flora (2nd Ed.) 386.

1928. K. fossilis W. R. B. Oliver Trans. N.Z. Inst. 59: 295, Fig. 11.

1928. Quintinia waipaoaensis W. R. B. Oliver. Trans. N.Z. Inst. 59: 300, Fig. 17.

Locality. Ormond B195/29, B195/12.

Age. Castlecliffian (Pleistocene).

The type leaf of Knightia fossilis has been further cleaned to expose more of its margin, which is more regularly dentate than described. The minor irregular teeth near the base are matched in leaves of adult Knightia excelsa. The nature of the margin was Oliver's reason for distinguishing the fossil species from the living one.

The type of Quintinia waipaoaensis was compared with recent Quintinia spp., and with Knightia excelsa The margin is unlike that of the living New Zealand Quintinia spp., but is far closer to that of Knightia excelsa. The venation also matches Knightia and the size is comparable.

Family Proteaceae

Genus Proteoides Heer.

Proteoides ohukaensis n.sp. (Text-fig. 1, Figs. 6, 7.)

Fruit, orbicular to ovate in outline, bilaterally compressed, with slightly convex sides. Length 5 mm., greatest width 3 mm., least width 1.5 mm. Apex mucronate, base boss shaped 1 mm. in diameter.

Surface irregularly granular, possibly weathered. Wall of two fused layers, an outer thin layer less carbonized, and an inner layer carbonized to give a glossy fracture. Loculi two, fused for about ¼ of their length and surrounded by the inner carbonized wall which separates in the upper section of the fruit Each loculus lined with a thin glossy cellular layer.

The fruit has shrunk, possibly since the matrix dried after collection—a latex mould of the cavity when dried measured 4 mm. across the widest plane.

Locality. Mouth of Ohuka Ck B323/21 and 31, Paratypes.

Age ? Nukumaruan (Pleistocene)

This has been compared with Australian and South American Proteaceae, but has not been satisfactorily placed in a genus.

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Family Pittosporaceae

Genus Pittosporum Banks

Pittosporum cf. colensoi Hook. f.

1853. Pittosporum colensoi Hook. f. Fl. Nov. Zel. i 22.

1925. —— Cheesem. Man. N.Z Fl. (2nd Ed.) 487 (with synonomy).

1928. Pittosporum oblongum W. R. B. Oliver Trans. N.Z. Inst. 59: 299. Fig. 14.

Locality. Ormond B195/32.

Age. Castlecliffian.

In comparing the Waipaoa specimen with P. colensoi, Oliver found that the fossil had a broader basal angle and wider angles of divergence of the secondary veins. Leaves of recent specimens of P. colensoi are extremely variable and are hard to differentiate from other species of the group P. tenuifolium Banks et Sol, P. colensoi Hook, f., and P. fasciculatum Hook, f., of Cheeseman (1925).

The fossil is very similar to specimens of P. colensoi in the Herbarium of the Dominion Museum, and the basal angle is close to plants from Bluff (coll. 1945) and Banks Peninsula (coll. Armstrong). The angles of divergence of the secondary veins is close to those of the Banks Peninsula specimens.

Family Myrtaceae

Genus Leptospermum Forst.

Leptospermum ericoides A. Rich.

1832. Leptospermum ericoides A. Rich. Fl. Nouv. Zel. 338.

1925. —— Cheesem. Man. N.Z. Fl. (2nd Ed.) 589 (with synonomy)

1928. L. pliocenicum W. R. B. Oliver. Trans. N.Z. Inst. 59. p. 298.

Locality Ormond B195/33.

Age. Castlecliffian (Pleistocene).

Oliver separated the Waipaoa fossil from L. ericoides because the capsules are solitary; capsules are frequently solitary in many recent specimens of L. ericoides where the flowers are usually fascicled, but with often only one capsule reaching maturity. The fossil is thus classified under Leptospermum ericoides. Under the International Rules of Botanical Nomenclature 1948 (Art. 39), Oliver's name is invalid as the description lacked an illustration.

Family Malvaceae

Genus Plagianthus Forst.

Plagianthus betulinus A. Cunn.

1839. Plagianthus betulinus A. Cunn Precur n. 606.

1925. Plagianthus betulinus Cheesem Man N. Z. Fl. (2nd Ed) 562.

1928. Nothofagus fusca W. R. B. Oliver. Trans. N.Z. Inst. 59: 292. Fig. 4.

Locality. Ormond B195/17.

Agez Castlecliffian (Pleistocene).

The fossil leaf from Ormond identified as Nothofagus fusca by Oliver (1928) shows two features foreign to those in New Zealand Nothofagus leaves with dentate margins: the marginal teeth form re-entrant angles of about 90°, and the secondary veins branch ¼ of the distance between the midrib and the margin.

These features and a size range of 2.5–7 cm. length are found in leaves of Plagianthus betulinus.

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Family Papilionaceae

Genus Carmichaelia R. Br.

Carmichaelia sp.

1928. Carmichaelia australis W. R. B. Oliver in Trans N.Z Inst. 59. 299. Fig. 28.

Locality. Ormond B195/24.

The specimen identified as Carmichaelia australis is not well enough preserved to identify specifically.

Family Betulaceae

Genus Betulites Göppert

Betulites couperi n.sp. (Plate 27, Fig. 2.)

Portion of an ovate leaf, base slightly uneven, with each margin meeting the midrib at 90°. Length 22 mm., width 22 mm. Petiole 1 mm. long, 1 mm. wide. Margin slightly crenate. Midrib slightly sinuate, with fine groove down centre, running into base of each vein. Venation of moderate relief, craspedodromous with more than five pairs of alternate veins leaving the midrib. Lower veins leaving midrib at 60°-70°, upper veins at 30°-40°, intermediate veins at intermediate angles Several veins branching once or twice about ¾ distance to margin Tertiary venation of numerous closely set intersecondaries, branching frequently.

Locality “Erewhon” Station, Rangitikei R. B163/9 Holotype.

Age. Opoitean (Pliocene).

Although the leaf is incomplete the nature of the secondary venation, and the type of intersecondary venules is very close to these features of leaves of members of the Betulaceae—e.g., Betula verrucosa Elub Members of the Betulaceae grow in the Andes and Argentina (Willis, 1948), but have not previously been recorded from New Zealand.

Family Fagaceae

Genus Nothofagus Blume

Nothofagus searellae n. sp. (Pl 27, Figs. 3, 4)

Lanceolate leaf with entire margin, basal angle 150° Length 22.5 mm., width 10 mm. Remnant of petiole 0.75 mm. wide, 0.25 mm. long. Midrib straight and well defined to apex, 0.75 mm. wide at base, 0.1 mm. wide at apex. Venation craspedodromous with seven pairs of alternate secondaries leaving midrib at 50° -60°. Each secondary branching of way to margin into two equal short veins merging into the tertiary reticulation.

Tertiary reticulation a roughly rectangular mesh with mterstices about 0.5 mm. across.

Locality Esk Valley B206/7 Holotype. and Esk Valley localities tabulated below.

Age. Nukumaruan.

Other specimens have the following dimensions:

Length Width Basal Angle
B206/7 28 mm 6 mm c 150°
B215/94 24 mm. 8 mm.
B215/5 28 mm. 10 mm.
B209/10 25 mm 8 mm
B209/30 24 mm 9 mm
B209/25 25 mm. 11 mm. 120°
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215/5, 209/10, 209/30 are incomplete at the base, but if symmetrical their basal angles exceed 90°. B206/7 is long and narrow, but B209/25 and /30 are much wider. There are 6 or 7 pairs of alternate veins, all branching in the same way. Tertiary venation is generally not well preserved. The leaves of N. searellac are similar in shape to the group N. solandri-cliffortioides but are distinguished by their larger size and more numerous secondary veins.

This species is named after my wife, who helped considerably in the location and collection of the Esk Valley fossils.

Nothofagus cf. cliffortioides × fusca (Pl. 27, Fig. 5)

Leaf ovate, with rounded apex and tapering base, left half of lamina joining petiole slightly below the right side Margin entire, slightly downcurved.

Lamina 24 mm. long, 13 mm. wide, petiole 3 mm. long, 1 mm. wide, basal angle 75°.

Midrib straight near base, sinuate above, persistent almost to apex. Venation craspedodromous, 8 pairs of alternate and subopposite secondaries leaving midrib at angles varying from 35° at base to 60° nearer apex. Secondaries forking once between ¾ and ⅞ of distance to margin. Tertiary venation of oblong reticulae about 0.3 mm. across.

Locality. Esk Valley B206/2 and /3 (counterparts).

Age Nukumaruan.

This specimen is similar to leaves of hybrids between Nothofagus fusca and cliffortioides, that Kirk named N. blairii (see Cockayne, 1926) N. cliffortioides leaves have been identified from B206 but no N. fusca leaves were found.

Family Rutaceae

Genus Melicope Forst

Melicope ternate Forst.

1776. Melicope ternata Forst Char Gen 56.

1925. —— Cheesem Man S.Z. Fl (2nd Ed.) 539.

1928. Melicope elliptica W. R. B. Oliver. Trans N.Z. Inst 59: 300, Fig. 16.

Locality. Ormond B195/19.

Age. Castlecliffian (Pleistocene).

The specimen figured by Oliver is stated to differ from M. ternata only in the shape of the leaf. Living M. ternata leaves vary considerably in shape and some are identical with the fossil.

Family Rubiaceae

Genus Coprosma Forst

Coprosma australis (A. Rich.) Robinson.

1832. Ronabea australis A Rich. Fl. Nouv Zel. 187.

1853. Coprosma grandifolia Hook. f Fl. Nov Zel I 104.

1910. C. australis Robinson, Proc. Am. Acad. Arts and Science, 45. 40S.

1925 C. grandifolia Cheesem. Man N.Z. Fl (2nd Ed.) 858.

1928 C. vulcanica W. R. B. Oliver, Trans N. Z. Inst. 59: 303, Fig. 23.

1935 C. australis W. R. B. Oliver, Bishop Mus Bull. 132: 107. Fig. 59A, Fig. 35.

Locality. Ormond B195/25.

Age. Castlecliffian (Pleistocene).

The fossil described by Oliver (1928) as C. vulcanica is stated to differ from C. australis in having a wider angle between midrib and secondaries, in being

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less cuneate at base and apex, and in its emarginate apex. Recent specimens vary in all these characters to such an extent that the fossil cannot be systematically separated.

Family Araliaceae

Genus Nothopanax Miq.

Nothopanax edgerleyi Harms (Text-figs. 3–6)

1853. Panax edgerleyi Harms Hook f. Fl Nov. Zel i 94.

1925. Nothopanax edgerleyi Cheesem. Man. N.Z. Fl (2nd Ed.) p. 635 (with synonomy).

1928. Nothopanax reticulatum W. R. B. Oliver, Trans. N. Z. Inst. 59, p. 301, Fig. 18.

1928. Apocyanophyllum novae zelandiae W. R. B. Oliver, Trans. N. Z. Inst. 59, p. 296, Fig. 18.

1928. A. inaequilaterale W. R. B. Oliver, Trans N.Z Inst. 59. p. 298, Fig. 15.

Locality. Esk Valley B209/103, Ormond B195/1, 195/23, 195/37.

Age Nukumaruan-Castlecliffian (Pleistocene).

The shape and venation of Oliver's three fossil species can be closely matched amongst the leaves of the living Nothopanax edgerleyi. Nothopanax reticulatum is stated to differ from N. edgerleyi in the abruptly narrowed base, and absence of regular loops between the ends of the secondary veins, but the basal outline and fine reticulation can be matched in herbarium specimens of N. edgerleyi. The fossil has regular secondary loops similar to those of N. edgerleyi.

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Text-Figs 3–6.—3—Nothopanax edqerleyi Miq. Living specimen from Endeavour Inlet. (Herb. Dom. Museum) × 0 65 4—Nothopanax reticulatum Oliver. B195/1 (After Oliver, 1928) × 0.75. 5—Apoonophyullum Oliver B195/37 (After Oliver, 1928) × 0. 75. 6—A. inaequilaterale Oliver. B195/23 (After Oliver, 1928).

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The unequalsidedness of the base of Apocyanophyllum inaequilaterale is matched by recent specimens of N. edgerleyi; the venation of the fossil compares well with that of Nothopanax edgerleyi.

Apocyanophyllum novae zelandiae is identical in venation with Nothopanax edgerleyi, but smaller than A. inaequilaterale. Both fall within ranges of size and shape shown by Nothopanax edgerleyi.

Plantae Incertae Sedis
Genus Phyllites Sternberg

Phyllites kaiwakaensis n.sp (Pl. 27, Fig. 6)

Ovate leaf, 34 mm. long, 22 mm. wide, entire margin. Venation craspedodromous, midrib and secondaries deeply impressed. Midrib 2 mm. wide at base, 0.5 mm. wide at apex. Seven secondaries on one side, six on the other, opposite below and alternate above the third pair, all leaving midrib at 50°-60° and curving up distally, parallel to the margin. Tertiary venules short at right angle to secondaries, prominent for about ¼ distance across primary areas.

Locality. Esk Valley B209/1, Holotype.

Age. Nukumaruan (Pleistocene).

This specimen could not be compared with anything in the recent New Zealand flora and is assigned to the form genus Phyllites. It is named P. kaiwakaensis after the name of the original Esk Valley sheep station.

Acknowledgments

The writer's thanks are extended to all who helped collect the fossils described in this paper, to Mrs W. P. Tolley (N. Z. Geological Survey) for her drawings of the seeds and fruits, and to Mr. B. G. Hamlin (Dominion Museum) for help with problems of nomenclature.

Literature Cited

Berry, E. W., 1926. Cocos and Phymatocaion in the Pliocene of New Zealand. Amer. Journ. Sci., Series 5. vol. 12. no. 69: 181–184.

Camp, W. H., Rickett, H. W., Weatherby C. A. 1947 International Rules of Botanical Nomenclature. Brittonia 6:1.

Cockayne, L., 1926. Monograph on the New Zealand Beech Forests, Part 1. N.Z. State Forest Bull., No. 4.

Couper, R. A., 1952 The Spore and Pollen Flora of the Cocos-bearing Beds, Mangonui, North Auckland. Trans. R. Soc. N.Z., 79.3 and 4: 340–348

— 1953. Plant Microfossil Dating of some New Zealand Upper Tertiary Volcanic Rocks N.Z.J.S.T (B.) 34 No. 5: 373–377.

— and McQueen, D. R., 1954. Phocene and Pleistocene Plant Fossils of New Zealand and Their Climatic Interpretation. N.Z.J S.T.(B) 35 No. 5: 398–420

Oliver, Oliver. W. R., 1928. The Flora of the Warpaoa Series (Later Pliocene) of New Zealand Trans. N.Z. Inst 59 (2). 287–303.

— 1936. The Tertiary Flora of the Kaikorai Valley, Otago, New Zealand. Trans. Roy. Soc. N.Z., 66 (3): 284–304.

Park, J., 1886 Reports of Geological Explorations, XVIII 38

T. E. Punga, M. T., 1952. The Geology of Rangitikei Valley, N. Z. Geol. Surv. Mem 8. 13.

Willis, J. C., 1948. A Dictionary of Flowering Plants and Ferns. 6th Edition, C.U.P.

Yuncker, T. G., 1952. New Species of Peruvian and Colombian Piperaceae, Am Journ. Bot Vol. 39.