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Leaves of Middle and Upper Cretaceous Pleridophytes and Cycads from New Zealand

By D. R. McQueen,
New Zealand Geological Survey

[Received by the Editor, August 1, 1955.]

Abstract

Seventeen specres of middle and upper Cretaceous pteridophytes and cycads are recorded from eleven New Zealand localities, and the following new species described. Sphenopteris mackayi, Cladophlebis wellmann, Pterophyllum clarencianum, Ptilophyllum seymouicum. The Paparoa Coal Measures, which have never been satisfactorily dated, contain plant fossils suggesting Senonian age.

Introduction

In New Zealand, as elsewhere, the Cretaceous was marked by the first appearance of angiosperms and their rapid increase in number and diversity. At the same time the dominant plants of the lower Mesozoic, chiefly pteridophytes and cycads, were declining in importance, and pteridophytes similar to those of to-day were appearing for the first time.

Text-fig. 1, based partly on unpublished work, shows the change in the composition of New Zealand Cretaceous floras so far known It shows that the pteridophytes decreased in importance after the Neocomian, and that the cycads decreased in the later Cretaceous after a slight increase in the Turonian. Conifers increased slightly during the Cretaceous. Angiosperms are first known about the Turonian and become dominant before the Senonian.

This paper deals with the systematics and stratigraphie distribution of the pteridophytes and cycads from the middle and upper Cretaceous of New Zealand.

Picture icon

Text-Fig. 1—Diagram to show changes in dominance of plant groups in New Zealand during the Cretaceous, based on percentages of species in collections from Waikato Heads (Neocomian),† Seymour River coal measuies ( [ unclear: ] Turonian), Paparoa coal measures (Lower Senonian), and from Shag Point and Pakawau (Upper Senonian).

Now with Department of Forestry, Limbe, Nyasaland

[Footnote] † In 1954 ammonites from the Puaroan Stage of Kawhia were provisionally identified as Neocomian and the Waikato Heads plant berts were also classed as Neocomian when the author wrote this paper. New evidence on sequence at Kawhia has since led to revision of this dating, and it now seems unlikely that the Waikato plant beds are younger than Tithonian C. A. Fleming (Note added in proof, June, 1956)

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Acknowledgments

I should like to express my thanks to those in the Geological Survey who have helped in the preparation of this paper: Mrs. W. P. Tolley, for the drawings, Dr. C. A. Fleming and Dr. H. W. Wellman for their help with stratigraphic problems.

Previous Work

In 1886 Hector gave the name Taeniopteris stipulata to a fossil leaf, figured without a description, from Pakawau. In the following year Ettingshausen described and illustrated one fern from Shag Point and four from Pakawau. A Neocomian collection from Waikato Heads described by Arber (1917) and revised by Edwards (1934) includes two species that are now known to range up into higher parts of the Cretaceous. Arber also figured a slab from Pakawau, bearing pinnae of Cladophlebis prisca (Ett.) and Taeniopteris stipulata Hect, which had been labelled “Waikato Heads” in error (McQueen, 1955). In 1926, Edwards described and illustrated Sphenopteris sp. and Taeniopteris batleyensis n.sp from the upper Cretaceous ammonite beds of Kaipara Harbour.

Age

All the plant fossils described in this paper are from beds that have long been considered middle and upper Cretaceous. The collections examined are listed below, grouped according to their known or inferred ages:

New Zealand Cretaceous Fossil Plant Localities.

N.Z. Geol. Surv. Age and Locality.
Register.
Albian (Clarence Series).
B470 Tributary of Cover Creek, Coverham, from concretion containing
Albian Mollusca (S35/617, grid reference 155463).
? Turonian (Raukumara Series)
The following collections are from the Seymour Coal Measures conformably overlain by marine sediments containing basal Raukumar [ unclear: ] (? lower Senonian) mollusca:
B125 Quail Flat, Clarence River (Gs560 = S41/500).
B166 Quail Flat, Clarence River (Gs568 = S41/492).
B351 West bank, Seymour River (S41/549, grid reference 743147).
B356 East bank, Seymour River (S41/537, grid reference 734091).
B358 Quail Flat, Clarence River (S41/550, grid reference 722145).
Lower Senonian (Paparoa Coal Measures)
B23 Ford Formation, drillhole 241 (S44/731, grid reference 832973).
B172 Waiomo Formation, Paparoa mine (S44/483, grid reference 878001)
B52
B428 Rewanui Formation, Strongman mine dump (S44/752, grid reference 800000).
B19 Goldlight Formation, drillhole 266, Brunner Bridge (S44/727, gird reference 830904).
Upper Senonian (Sub-Piripauan)
B160 Shag Point (Gs414 = S146/519, additional specimens in Otago Museum).
B161 Pakawau (Gs410 = S3/492, grid reference 910070; additional spec [ unclear: ] mens in Canterbury Museum).
B530 Mikonui Stream, coast south of Oaro River, Marlborough (S56/510, grid reference 799762).
B160 and B530 are from coal measures underlying fossiliferous Piripauan
Maestrichtian.
B182 Bulls Point and Batley, Kaipara Harbour, from concretions containing Maestrichtian ammonites (N28/626, grid references 806390 and 843365).
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The collections from Coverham (Albian) and Kaipara (about Maestrichtian) are dated from associated marine fossils. Those from Shag Point and Mikonui Stream are from coal measures conformably underlying Piripauan (upper Senonian) marine sediments and are probably themselves upper Senonian. The Pakawau coal measures have been correlated with those at Shag Point on the evidence of plant microfossils (R. A. Couper, pers. comm.).

The collections from Seymour River and Quail Flat (middle Clarence Valley) are from coal measures conformably underlying marine sediments containing Trigonia glyptica, T. meridiana (Woods, 1917) and fragments of a giant Inoceramus which is the zone fossil of the basal Raukumara Series as redefined by Wellman (1955). Wellman (1955) has correlated the base of the Raukumara Series with the base of the Senonian. The Seymour River coal measures are thus either basal Senonian or probably Turonian. They are here classed as ? Turonian.

The Paparoa Coal Measures have never been dated from satisfactory paleontological evidence and are variously classed as lower Cretaceous (?) (Wellman, 1950), possibly pre-Cretaceous (Cotton, 1951), probably upper Cretaceous (Gage, 1952) and lower Cretaceous (Couper, 1953).

The fossil cycads and ferns described below provide the following additional evidence for dating the rocks in which they occur:

The Seymour River coal measures contain leaves of Ptilophyllum and Pterophyllum, genera recorded in New Zealand from the Jurassic but not previously from the Cretaceous. Pterophyllum does not range above the Jurassic in other parts of the world (Seward, 1933: 332) and is apparently an archaic element in the New Zealand Cretaceous. Ptilophyllum ranges up to the middle Cretaceous in other countries (Seward, 1933, 413).

In New Zealand, Taeniopteris stipulata apparently replaced the Jurassic and Neocomian Taeniopteris spatulata during the Lower Cretaceous. No beds are known to contain both species. T. spatulata is present in the Styx River Series of Queensland, tentatively classed as Cenomanian by David and Browne (1950), but T. stipulata is unknown in Australia. If a change from T. spatulata to T. stipulata occurred at the same time in Australia and New Zealand, the Seymour River coal measures (which contain T. stipulata) are probably post-Cenomanian and may well be Turonian as suggested above.

Four species from the Paparoa Coal Measures are unknown in the upper Senonian of New Zealand, but range back to middle Cretaceous or older strata (Sphenopteris mackayi, Phyllopteris lanceolata, P. expansa, and Coniopteris ? lobata). On the other hand, the Paparoa Coal Measures lack the genera Pterophyllum and Ptilophyllum that persist into the Seymour River coal measures and contain more angiosperms than known pre-Senonian floras in New Zealand. They are thus probably older than upper Senonian but younger than Turonian, that is, lower or middle Senonian.

The Shag Point and Pakawau Coal Measures have been correlated from their plant microfossils (R. A. Couper, pers comm.). They seem to be younger than the Paparoa Coal Measures because they lack the older Mesozoic species of Phyllopteris, Coniopteris and Sphenopteris (mackayi) and contain three species of fern unknown in Paparoa or older beds.

As the Shag Point and Mikonui coal measures underlie Piripauan (upper Senonian), they and the Pakawau coal measures may be slightly older than the plants from Kaipara, collected from concretions containing ammonites probably of Maestrichtian age (Mason, 1953: 356). The Kaipara beds apparently lack the

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common upper Cretaceous species Taeniopters stipulata and contain Taemopteris batleyensis which has not been found at other horizons and thus may be an index species for the Maestrichtian, although it is a pteridophyte (Edwards. 1934) and thus not the descendant of T. stipulata.

The known ranges of pteridophytes and cycads in the Cretaceous of Australia and New Zealand, on which the above conclusions are based, are summarised in Table I.

  • Cladophlebis australis (Morr.)

  • Coniopteris? lobata (Oldham)

  • Taeniopteris spatulata MacClell.

  • Phyllopteris lanceolata Walk.

  • P. expansa Walk.

  • Zamites takuraensis Walk.

  • Lycopodium of volubile Forst. f.

  • Sphenopteris mackayi n. sp.

  • Taeniopteris stipulata Hect.

  • Pterophyllum (clarencianium n. sp.)

  • Ptilophyllum (seymouricum n.sp.)

  • Cladophlebis wellmanii n. sp.

  • C. prisca (Ett.)

  • C. obscura (Ett.)

  • Sphenopteris pteriodies (Ett.)

  • Sphenopteris sp. (Edwards 1926)

  • Taeniopteris batlcyensis Edw.

Waikato Heads1 Neocomian
Burrum, Queensland2 Aptian-Albian
Coverham Albian
Styx. Queensland3 Cenomanian
Seymour R. coal measures Turonian
Ford
Waiomo
Rewanui
Goldlight Paparoa Coal Measures
Lower Senonian
Pakawau4
Shag Point4
Mikonui Upper Senonian
Kaipara5 Maestrichtian
Table I—Stratigraphic Distribution of New Zealand and Australian Cretaceous Pte [ unclear: ] dophytes and Cycads. 1Arber 1917, Edwards 1934; 2Walkom 1919 David and Browne 1950, 3Walkom 1919; 4Ettingshausen 1887, Suggate and Couper 1952, 5Edwards 1926.

Systematics
Order Lycopodiales

Genus Lycopodium Linn

Lycopodium cf. volubile Forst. f. (Text-fig. 2, Fig. 14).

1786. Lycopodium volubile Forster f., Prodi n. 482.

1925.——Cheeseman. Man. N. Z. Flor. 2nd Edn 104 (with full synonymy)

Description. Apex of a branch preserved with dorsal side upmost. Midrib straight, bearing falcate leaves tapering at apex only Leaves with prominent midribs.

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Locality. Seymour River, B351/10.

Age. Turonian to Recent.

This fragmentary specimen has somewhat larger leaves than Recent specimens but agrees in all other characters.

Order Filicales

Genus Aspidium Linn

Aspidium cretaceozelandicum Ettingshausen.

1887. Aspidium cretaceozelandicum Ettingshausen, Denkschr. k Akad. Wissen Wien (Math-Nar. Cl.) 53 (1). 143.

1890.—Ettingshausen, Trans. N. Z. Inst 23: 50, Pl XXX, Figs. 2, 3.

Locality. Pakawan (Canterbury Museum Zf 2 77, Syntype, possibly Pl. XXX, Fig. 2.)

Age. Upper Senoman.

Examination of the syntype at Canterbury Museum has shown that its classification as a fern is doubtful. There is no trace of the secondary veins of the pinnules shown in Ettingshausen's figure. It may belong to the Proteaceous genus Dryandra, known from the Paparoa Coal Measures, from the upper Senonian of Pakawau (as Dryandroides, Ettingshausen, 1887. 68) and from possibly upper Senonian coal measures from Trelissick Basin.

Genus Sphenopteris Brongniart

Sphenopteris mackayi n. sp. (Text-fig. 3, Figs. 1-3.)

Description. Pinnately branched frond, 6 cms long, with winged rachis and pronounced central vein Pinnae deltoid, arising at acute angles, distal part often split, margin entire, apex truncate Venation of pinnae. one prominent vein leaving the rachis and dichotomising three or four times.

Localities. Quail Flat B166/70, (Holotype); Paparoa Coal Measures (Waiomo Formation) B172/5.

Age. Turonian to lower Senonian.

No species matching this has been seen in other New Zealand Mesozoic floras. It is similar in general shape and venation to Sphenopteris (Ruffordia) goepperin Dunker (Arber, 1917. 42; Seward, 1894. 81) but is larger and its secondary veins dichotomise more frequently.

Sphenopteris pterioides (Ettingshausen)

1887. Dicksonia pterioides Ettingshausen, Denkschr k. Akad. Wissen Wien (Math-Nat Cl) 53 (1) 143.

1887. Aspidium otagoicum Ettingshausen, Denkschi k. Akad Wissen Wien (Math-Nat Cl) 53 (1). 143.

1890. Dicksonia ptetioides Ettingshausen Trans. N.Z. Inst. 23: 51. Pl. XXX, Figs. 4-6.

1890. Aspidium otagoicum Ettingshausen, Trans. N. Z. Inst. 23: 15. Pl. XXIV, Fig. 3.

Localities Pakawan (Canterbury Museum Zf 279, a syntype, original of Pl XXX, Figs. 6 6a); Shag Point (Otago Museum C48 68, holotype of Aspidium otagoicum, Pl XXIV, Fig. 3).

Age. Upper Senonian.

As the specimen seen has no trace of sori and as Ettingshausen's other figures are also based on sterile fronds, this species is transferred to Sphenopteris. Two other specimens labelled “Dicksonia pteriodes Ett”, in the Canterbury Museum (Zf 278 and Zf 280), neither figured by Ettingshausen, are fragmentary, and may not be conspecific.

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Picture icon

Text-Fig. 2.—Figs. 1.4—Cladophelebis australis (Morris). 1 B52/1 Strongman Mine, Lower Senonian; 2. B161/6 Pakawau. Upper Senonian: 3 B166/69 Quail Flat, ? Turonian, 4, B530/7 Mikonui Stream, Upper Senonian. Fig. 5—Cladophlcbis [ unclear: ] llmann n. sp. B428/23, holotype, Strongman Mine. Lower Senonian Fig. 6—Cladophlebis obscura (Ettingshausen) B161/7 Pakawan Upper Senonian Figs. 7.9—Coniopteris [ unclear: ] lobata (Oldham) 7. B125/4 Quail Flat, [ unclear: ] Turonian, 8, B52/5, 428/28 Strongman Mine, Lower Senonian, 9. B166/69 Quail Flat, [ unclear: ] Turonian Figs. 10.12—Phyllopte [ unclear: ] s expansa Walkom. 10, 11, B125/34, 61 Quail Flat, ? Turonian, 12, B19/39, Brunner Bridge, Lower Senonian. Fig. 13—Phyllopteris lanceolata Walkom. B23/1 Ford Formation, Lower Senonian. Fig. 14—Lycopodium of volubile Forst. B351/10 Seymour River, ? Turonian All Drawings natural size.

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Genus Cladophlebis Brongniart

Cladophlebis australis (Morr.) (Text-fig. 2, Figs. 1-4.)

1845. Pecopteris australis Morris, in Strzelecki, Phys. Desci. N. S. Wales: 248, Pl. VII, Figs. 1, 2.

1917. Cladophlebis austrahs Arber, N.Z. Geol. Surv. Pal. Bull. 6: 29. Pl. IV, Figs. 1. 5, 7, 8; Pl. XIV (with full synonomy).

1919. C. ausiralis Walkom, Queensland Geol. Surv. Publ. 263: 13, Pl. 3, Figs. 6, 7. Dimensions. See Table II.

Table II.-Dimensions of New Zealand Cladophlebis.
Length of Pinnules Width of Pinnules
Average angle of divergence Average angle of divergence of Number of Secondaries per cm at
Locality and No. Cm cm
of Pinnules Secondaries margin.
Cladophlebis australis
Waikato Heads B77/2 1.7-3.3 0.7
70° 57° 16-18
Strongman Mine B52/1 1.6-1.9 0.8
63° 67.6° 12-14
Pakawau B161/6 1.7-1.9 0.6-0.7
74° 53° 14
Quail Flat—B166/68 0.7-1.1 0.4
67.5° — —
B125/33 1.7-1.8
0.35-0.4 75.5° —
—
B166/69 1.0-1.4
0.4-0.5 58.5° —
—
B166/67 0.9-1.2
0.4-0.5 60.5° —
—
B166/65A 1.2
0.4 62.5° —
—
1.1 0.35
58.2° — —
Mikonui Stream—B530/1 >1.2 0.5
56° 59° 20
B530/4 1.1
0.35 — 55°
—
B530/2 >0.9
0.5 62° 63°
20
B530/5 —
— 68° 61°
—
B530/7 1.1
0.4 60° 49°
—
Cladophlebis obscura
Pakawau B161/7 2.5-3.0 0.5-0.6
85° 56° 14
Cladophlebis prisca
Pakawau B161/10 2.5 0.6-0.7
67° 43° 14-16

Localities. Waikato Heads B65/2; Pakawau B161/6; Strongman Mme (Rewanui Formation) B52/1; Quail Flat B166/30; Mikonui Stream B530/7.

Age. Rhaetian to Upper Senonian in New Zealand.

Cladophlebis australis was originally described from New South Wales, and is a widely distributed Southern Hemisphere fern, known from South Africa, Patagonia, Australia and Antarctica (Arber, 1917: 30, 31).

The pinnules are all more or less falcate, with entire margins. There are certain differences in the mode of attachment of pinnules to the rachis. In the Rhaetian specimen of Cladophlebis australis from Owaka Creek (Arber, Pl. IV, Fig. 5) the pinnules are fused basally producing a wing along the rachis. In one specimen from the Neocomian of Waikato Heads (B77/2) the pinnules are free to their junction with the rachis, but in another specimen (Arber's Pl. V, Fig. 8) they are winged. The specimens figured by Walkom (loc. cit.) from Burrum (Aptian or Albian) have similar wings. The specimens from the Paparoa Coal Measures all have winged bases, but those from Pakawau have the pinnules free basally. Those from Quail Flat and Mikonui Stream are distinguished by their small size and often have straight midribs in their pinnules.

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Picture icon

Text-Fig. 3—Figs. 1-3—Sphenopteris mackay [ unclear: ] n. sp. Quarl Flat, [ unclear: ] Turonian Holotype B166/70 (1) and paratypes (B166/29 and B166/34). Figs. 4-7—Taeniopte [ unclear: ] s Stipulata Hector. 4. B161/24 Pakawau, lectotype, Upper Senoman; [ unclear: ] B160/105 Shag Point, Upper Senoman; 6 B161/30 Pakawau: 7, B358/8 Quail Flat, ? Turonian. Figs. 8 9—Ptilophllum seymouricum n. sp. Seymoui River.? Turonian. Holotype B356/30 (8) and paratype B356/4. Fig. 10—Pterophullum d [ unclear: ] e [ unclear: ] n. sp. B106/60 Quail Flat,? Turonian, holotype. Fig. 11—Zamites [ unclear: ] tak [ unclear: ] raensts Walkom. B470/1 Coverham, Albian.

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Cladophlebis prisca Ettingshausen.

1887. Blechnum priscum Ettingshausen, Denkschi. k. Akad. Wissen Wien (Math-Nat.Cl) 53 (1): 143.

1890. ——Ettingshausen. Tians. N.Z. Inst. 23: 50, Pl. XXX, Figs. 1, la.

1917. Cladophlebis austialis Arber, N.Z. Geol. Surv. Pal. Bull. 6, Pl. 4, Fig. 1; Pl. XIV.

Type. In the Canterbury Museum, according to Ettingshausen (1890), but not located at the time of writing.

Locality. Pakawau B161/10. Arber (1917, Pl. XIV) figured a specimen of Cladophlebis prisca as C. australis on a slab with angiosperm leaves, reputedly from Waikato Heads (Neocomian) but actually from Pakawau (McQueen 1955).

Age. Upper Senonian.

As Ettingshausen's species is completely sterile, it is better classed under Clodophlebis. It is characterised by the dilated bases of the pinnules and by the secondary veins diverging more acutely than those of C. australis (see Table III).

Cladophlebis obscura (Ettingshausen) (Text-Fig. 2, Fig. 6.)

1887. Gleichenia obscuia Ettingshausen, Denkschi. k. Akad. Wissen Wien (Math-Nat. Cl) 53: 143.

1890. Gleichenia obsura Ettingshausen, Trans. N. Z. Inst. 23. 52, Pl. XXX. Figs. 7, 7a: Locality. Pakawau, B161/7.

Age. Upper Senonian.

Ettmgshausen's type from Pakawau should be m Canterbury Museum, but has not been recognised there. The Pakawau specimen figured here agrees with Ettingshausen's figure of Gleichenia obscura in its pinnae, which diverge at a wide angle, but differs somewhat in venation.

Cladophlebis wellmanii n.sp. (Text-fig. 2, Fig. 5.)

Description. Rachis striate. Fronds pinnate. Pinnae widely set, each attached by the whole base, parallel sided, tapering in last quarter. Margins finely serrate. Midrib with a median groove; secondary veins curving away from midrib, dichotomising once basally and once medially.

Dimensions. Width of midrib, 1.5 mm; length of pinna, 4.5 cm; width of pinnae, 1.4 cm; secondaries 20 per cm at margin (holotype). An incomplete paratype (B 428/2) has pinnae 16 mm wide and secondaries 14 per cm at margin.

Locality. Rewanui Formation, Paparoa Coal Measures, Strongman Mine, B428/23 (holotype) and B428/2.

Agf. Lower Senonian.

The pinnae resemble those of Blechnum in form. They are the same size as those of B. procerum (Forst. f.) Labill but differ in the attachment to the rachis. As no fertile fronds are known, the new species has been placed in Cladophlebis. C. wellmann is quite distinct from C. australis and similar species.

Genus Coniopteris Brongniart

Coniopteris [ unclear: ] lobata (Oldham)? (Text-fig. 2, Figs. 7-9.)

1863. Pecopteris (?) (Asplenites) Lobata Oldham and Morris Fossils of the Gonduana system 1 (1): 52, Pl. 28. Fig. 1; Pl. 29, 30; Pl. 36, Fig. 3.

1913. Conropteris ? lobala Halle, Schued. sud Polar-Exp. 111 (140: 22, Text-fig. 3; Pl. 1. Fig. 27 ?; Pl 3, Fig. 13 ? (with full synonomy)

1934. Coniopteris ? lobala Edwards, Ann. and Mag. Nat. llist. Series 10, 13: 81, 93, Text-fig. 3.

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Description. Fronds tri-pinnate, pinnae opposite or subopposite. Pinnules close set and joined by wing along rachis. Margins entire, lobed in basal pinnules. Venation: a single vein leaving the midrib, sometimes dichotomising thrice, but one of the veins resulting from the first dichotomy usually remaining undivided.

New Zealand Localities. Waikato Heads (British Museum of Natural History No. V 23276). Quail Flat B125/4, B166/69. Strongman Mine B52/5, B428/8.

This species was first described from the Upper Gondwana of India and has been recorded from the Liassic of Austria and the Jurassic of Antarctica (Halle, 1913: 24).

Age. Liassic to Lower Senonian. (Neocomian to Senonian in New Zealand.)

Specimens from Strongman Mine (Rewanui Formation) are similar in shape to Halle's specimen (loc. cit., Text-fig. 5). The fragments from Quail Flat are tentatively placed in C. ? lobata because they resemble in shape and venation the apical portions of C. ? lobata from Strongman Mine.

Order Cycadales

Genus Zamites Brongniart

Zamites cf. takuraensis Walkom. (Text-fig. 3, Fig. 11.)

1919. Zamites takuraensis Walkom, Queensland Geol. Surv. Publ. 263: 31, Pl. 2; Fig. 1.

Description. Portion of a frond with subopposite pinnae set obliquely to axis; method of attachment obscure. Pinnae elliptic, slightly narrowed basally, acuminate, with entire margins. Surface finely striate. Venation parallel and simple, not converging at apex.

Dimensions. Length 6-7 cm, breadth 1.8-2.2 cm; there are 10 veins per cm at the widest part of the pinnae.

Locality. Coverham B470/1, B470/2.

Age. Aptian-Albian in Australia, Albian in New Zealand.

Walkom's figure shows only the basal portion of the pinna, but his description allows the New Zealand material to be compared with Z. takuraensis. Apart from its somewhat more widely spaced veins, the New Zealand material agrees with Walkom's description and figure of Zamites takuraensis. Other known Southern Hemisphere species of Zamites are considerably smaller than Z. takuraensis.

Genus Phyllopteris Brongniart

Phyllopteris lanceolata Walkom. (Text-fig, 2, Fig. 13.)

1919. Phyllopteris lanceolata Walkom, Queensland Geol. Surv. Publ. 263: 22, 56. Pl. 3, Figs. 1, 2, 13.

Description. Apical and central portion of an oblong pmna, basally distorted. Margin entire, midrib straight and not persisting to apex, secondary veins leaving midrib at an acute angle, arched and dichotomising once near base.

Dimensions. Length 2.5 cm, width 0.75 cm.

Locality. Paparoa Coal Measures: Ford Formation B23/1, Waiomo Formation B172/5.

Age. Aptian or Albian (Australia), Cenomanian (Australia), Lower Senonian (New Zealand).

The specimen figured agrees well with Walkom's species which is known from both the Burrum (Aptian or Albian) and Styx River Series (Cenomanian).

Phyllopteris expansa Walkom. (Text-fig. 2, Figs. 10-12.)

1919. Phyllopteris expansa Walkom, Queensland Geol. Suiv. Publ. 263: 24, Pl. 5. Figs. 1, 2.

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Description. Detached pinnae, ovate to elliptic. Margins slightly dentate or entire. Midrib straight, not persisting to apex. Secondary veins leaving midrib at acute angles, dichotomising three times, basally, medially and distally. Veins resulting from dichotomies arched apart at junctions.

Dimensions. Length 3 cm, width 2 cm (B19/39); length 3 cm, width 1.5 cm (B125/61); length 5.5 cm, width 1.75 cm (B125/34).

Localities. Quail Flat B125/34, B125/61; Paparoa Coal Measures (Goldlight Formation) B19/39, 40.

Age. Walkom records this species only from the Burrum Series (Aptian and Albian) and not from the Styx Series (Cenomanian); the New Zealand specimens are ? Lower Senonian.

The specimens agree reasonably well with Walkom's figures. There are fewer secondaries per tooth than in Walkom's figures, but the margins are badly preserved and cannot be compared easily.

Order Benettitales

Genus Taeniopteris Brongniart

Taeniopteris stipulata Hector. (Text-fig. 3, Figs. 4-7.)

1886. Taeniopteris stipulata Hector, Det. Cat. Ind. and Col. Exhib.: 61, Fig. 24A, 3.

Hector's name was published with an inaccurate figure, without description, of a specimen from Pakawau, in the collection of the New Zealand Geological Survey, here chosen as lectotype. The specimen originally agreed in outline with Hector's figure but has now been cleared of matrix to show the base and part of the apex.

Description. Leaf elliptic lanceolate, margin entire. Midrib with smooth surface. Secondary veins leaving midrib at less than a right angle, rarely dichotomising, then only once and always basally.

Dimensions. Length 6.5 cm (incomplete); width 1.2 cm; midrib 0.125 cm width at base; average angle of divergence of secondaries 81°; 36 secondaries per cm at margin (Lectotype B161/24).

Localities. See Table III.

Age. ? Turonian to Upper Senonian.

Table III.—Dimensions of Taeniopteris Stipulata Hector.
Locality. Length of Lamina cm Width of Lamina cm
Midrib Width cm Divergences of Secondaries Secondaries/cm at margin
Shag Point (B165/75) 3.8->4.3 1.1-1.7 (1.5)
0.01-0.17 (0.13) 76° 18-40 (27)
Quail Flat (B125/2, /10, /37; 358/8) 6.5-8.2 (7.3) 1.2-3.2 (1.9)
0.15-0.5 (0.28) 76°-85° (79°) 26-40 (34.5)
Pakawau (B161/19, /24, /25) 3.2->6.5 1.2-2.7 (1.6)
0.1 -0.2 (0.14) 74° -87° (80°) 28-36 (31.5)
Pakawau (B161/24, lectotype) >6.5 1.2
0.12 S1° 36
Pakawau* (B161/30) - 0.1.8
0.1 73° 30

[Footnote] * B161/30 is the original of Arber (1917), Pi. XIV, formerly labelled Waikato Heads in error (McQueen 1955).

– 684 –

Specimens from different localities vary in size (Table III), and there are minor variations in morphology amongst those from one locality.

The midribs vary from perfectly smooth to finely striate and occasionally (B125/10, B125/62) have a median groove in the basal portions, a feature probably dependent on the condition of preservation of the fossil. The number of secondary veins that dichotomise varies in different specimens, but such dichotomy is always found in the basal portions of the secondaries.

This species does not match any other New Zealand Taeniopteris. It differs from T. spatulata McLelland, which has been recorded from several New Zealand upper Jurassic localities (Arber 1917, Edwards 1934) in the following characters:

The midrib is seldom grooved, the secondary veins diverge from the midrib at a smaller angle, the secondary veins are more closely spaced, and those which dichotomise do so only in their basal portion, whereas in T. spatulata more secondaries dichotomise, and may do so anywhere between midrib and margin.

T. stipulata resembles T. howardensis Walkom (1919), which, however, is characterised by secondary veins that dichotomise in the outer portions of the lamina. T. stipulata is also very similar to the Wealden species. T. beyrichii (Schenk.) illustrated by Seward (1894, Pi. IX, Fig. 3) but differs in its more closely spaced veins (36 per cm at the margin, compared with 20 per cm in T. beyrichii).

Taeniopteris batleyensis Edwards.

1926. Taeniopteris batleyensis Edwards, Trans. N.Z. Inst. 56: 124, Figs. 1, 2.

Edwards provisionally classified this leaf, which is obviously a fern, as Taeniopteris. Mrs. W. P. Tolley has unsuccessfully attempted to isolate spores from the highly calcified sori. Accordingly, no change is made in the generic classification.

Locality. Bulls Point or Batley, Kaipara Harbour, B182/24 (Holotype).

Age. Maestrichtian.

Genus Pterophyllum Brongniart

Pterophyllum clarencianum n. sp. (Text-fig. 3, Fig. 10.)

Description. Medial portion of a frond bearing alternate and opposite pinnae. Rachis, with faint median groove. Pinnae contiguous at base and often fused for a short distance, slightly tapering, with entire margins and obtuse rounded apices. Secondary veins simple, closely set.

Dimensions. Length of incomplete frond, 6 cm; length of pinnae, 1.4-1.6 cm, width of pinnae, 0.5-0.7 cm; veins 0.4-0.5 mm apart.

Locality. Quail Flat B166/60 (Holotype).

Age. ? Turonian.

The specimen differs from the only other New Zealand species of Pterophyllum, P. matauraensis Hector. (Jurassic, Mataura Falls), in the more regular shape and obtuseness of its pinnae and in the finer pattern of secondary veins Pterophyllum is not known above the Jurassic (Seward, 1933: 332) in other parts of the world. P. clarencianum is similar to some species of Nilssonia, which extends into the Cretaceous, but Seward's definition (1917: 548-9) of Pterophyllum fits the New Zealand Cretaceous specimen better than his definition of Nilssonia.

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Genus Ptilophyllum Morris

Ptilophyllum seymouricum n. sp. Text-Fig. 3, Figs. 8, 9.

Description. Portions of frond bearing alternately spaced pinnae, interspaces equal to the width of a pinna. Pinnae attached at a wide angle to the upper surface of a rachis by its entire base. Base parallel sided or slightly auriculate on upper edge, decurrent on lower. Pinnae tapering to point and bearing four to six parallel or slightly diverging veins, apparently all originating at rachis.

Dimensions
Length Length of pinnae Width at base
Veins per pinna
(cm) (cm) (cm)
356/12 3.3 1 -1.2
0.2 4
356/14 2.1 1.2
0.15 -0.2 4-5
356/4 6.2 0.7-1.2
0.175-0.25
356/30 — 1.3-1.6
0.2 -0.25 4-6

Loaclity. Seymour River, B356/30 (holotype) and B356/4 (figured paratype).

Age ? Turonian.

This species is similar in some respects to P. pecten (Phillips), a European Jurassic species recorded from the Burrum (Aptian-Albian) beds in Australia (Walkom, 1919: 30) and from Jurassic strata in many parts of both hemispheres (Seward, 1917: 524 et seq.). P. seymouricum is also similar to P. andersonii (Halle) and P. antarcticum (Halle) from the Jurassic of Grahamland. It is distinguished by the wide spacing of the pinnae, which are their own width apart as if forced apart by the insertion of the alternate pinnae on the rachis. It differs from the other New Zealand species, Ptilophyllum acutifolium Morris (middle Jurassic of Waikawa) in its venation and in the shape of its pinnae which are dilated at their bases and less tapering.

References

Arber, E. A. N, 1917. The Earlier Mesozoic Floras of New Zealand N.Z. Geol. Suiv. Pal Bull. No. 6

David T. W. Edgworth, and Browne, W. R., 1950 The Geology of the Commonnealth of Australia, Edward Arnold, London.

Cotton, C. A., 1951. Post-Hokonui Oiogeny, Eiosion and Peneplanation N. Z. Set Tech B33 (3): 173-8.

Edwards, W N., 1926. Cretaceous Plants from Kaipara, New Zealand Trans. N. Z. Inst 56: 121-8.

—— 1934. Jurassic Plants from New Zealand. Ann. and Mag Nat Hist Set 10, (8): Si et seq.

Ettingshausen, Cv E., 1887 Beitrage zur Kenntniss der fossilen Floia Neuseelands. Denkschr. k. Akad. Wissen Wien (Math.-Nat. Cl.) 53 (1): 143 et seq. Trans. N.Z. Inst. 23: 237 et seq (1890).

Hector, J, 1886. Det. Cat. and Guide to Geol. Exhibits, Ind. and Col. Exhib. London. N.Z Geological Survey.

Mason. A. P, 1953 The Geology of the Central Portion of Hokianga Country, North Auckland. Trans. Roy. Soc. N.Z. 81 (3): 356.

MCqueen, D. R, 1955. Revision of Supposed Juiassic Angiosperms from New Zealand. “Nature,” 175: 177.

Seward, A. C, 1894. The Wealden Floia Pt 1 Cat. Mes Pl in Dept. Geol Brit Mus Nat Hist.

—— 1933. Plant Life through the Ages Cambridge Univ. Press.

Suggate, R. P and Couper, R. A., 1952 The Stratigraphic Relations and Plant Microfossils of New Zealand Coal Measures N. Z. J. Sci. Tech. B 34: 2.

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Walkom, A. B., 1919. The Mesozoic Floras of Queensland Pt. III: The Floras of the Burium and Styx River Series, Queensland Geol. Surv. Publ. 263.

Wellman, H. W., 1950. Paparoa-Brunner Contact near Greymouth. N.Z. J. Sci. Tech B, 32 (1): 10.

—— 1955. A Revision of the Type Clarentian Section at Coverham, Clarence Valley, S.35. Trans. Roy. Soc. N.Z., 83.

Woods, H., 1917. The Cretaceous Faunas of the North-Eastern Part of the South Island of New Zealand. N.Z. Geol. Surv. Pal. Bull. 4.

D. R. McQueen, M.Sc.,

Department of Forestry,
Limbe,
Nyasaland.

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