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Mesozoic leaf genus

Taeniopteris at Port
Waikato and Clent Hills,
New Zealand
a a b
P.M. Blaschke & J.A. Grant-Mackie
a
Botany and Geology Departments ,
University of Auckland , New Zealand
b
Geology Department , University of
Auckland , Private Bag, Auckland, New
Zealand
Published online: 21 Dec 2011.

To cite this article: P.M. Blaschke & J.A. Grant-Mackie (1976) Mesozoic
leaf genus Taeniopteris at Port Waikato and Clent Hills, New Zealand,
New Zealand Journal of Geology and Geophysics, 19:6, 933-941, DOI:
10.1080/00288306.1976.10420747

To link to this article: http://dx.doi.org/10.1080/00288306.1976.10420747

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 N.Z. Journal of Geology and Geophysics Vol. 19, No.6 (1976): 933-41

MESOZOIC LEAF GENUS TAENIOPTERIS AT

PORT WAIKATO AND CLENT HILLS, NEW ZEALAND
P. M. BLASCHKE AND J. A. GRANT-MACKIE
Botany and Geology Departme1l!s, University of Auckland, New Zealand*

ABSTRACT

Populations of the leaf form-genus Taeniopteris Brongniart from two middle

Mesozoic localities in New Zealand were studied. Attempts to obtain leaf cuticles
from the little-metamorphosed deposits of the Huriwai Formation at Port Waikato
were unsuccessful, and the affinities of taxa included within the form-genus in New
Zealand remain unknown.
The population of Taeniopteris at Port \X'aikato (topmost Jurassic), although
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variable, falls within the limits of the species T. daintreei McCoy, described from the
Jurassic and lower Cretaceous in Eastern Australia. T. dainlreei has been separated
from T. spatulala McClelland (= Nipaniophyllum raoi Sahni) on anatomical grounds,
and neither of the latter two names should be accepted for the Port Waikato leaves
unless definite Pentoxylalean affinities were to be shown for them. T. daintreei
(reported also from the Southland Temaikan) was not found in the Clent Hills and
appears to be confined to middle Jurassic and younger beds in New Zealand. T.
thomsoniana Arber, found in the Clent Hills (middle to upper Jurassic), is given an
emended description.
The distribution of Taeniopteris species during the Jurassic between the localities
discussed possibly provides a further indication of different provenances for Torlesse
and Murihiku strata.

INTRODUCTION

T aeniopteris is the name given to a group of gymnosperm leaves occurring

very widely throughout the world during the Mesozoic, but particularly
characteristic of Jurassic floras. It refers to simple, elongate-oval to straplike
leaves up to 20 em in length, with a prominent midrib and numerous
bifurcating secondary veins running out to the margin at approximately
right angles to the midrib.
The genus was erected by Brongniart (1828) .and has been since reported
from many localities in Northern Europe, Asia, South Africa, Australia,
New Zealand, and the Americas. In New Zealand, T aeniopteris occurs at
almost all important Mesozoic plant fossil localities. Along with the fern-like
frond Cladophlebis, it is the most common early Mesozoic plant fossil,
occurring as impressions or as carboni sed compressions.
Arber (1917) published the only important work on T aeniopteris in New
Zealand, recognising five local species. All valid and invalid names given
to New Zealand members of the genus are listed by Mildenhall (1970).
Since the time of Arber's work, it has been generally realised that
T aeniopteris is not a natural grouping but includes representatives of several
different gymnosperm groups with similar leaf forms. These may be dis-

Received 11 March 1976.

*All communications should be addressed to ]. A. G-M, Geology Department Univer-
sity of Auckland, Private Bag, Auckland, New Zealand. '
 934 N.Z. JOURNAL OF GEOLOGY AND GEOPHYSICS VOL. 19

tinguished by precise study of anatomical features such .as epiderm~l ~nd

stomatal structure (Harris 1932; Douglas 1969). ThIs commu~l~atlOn
presents results from a preliminary study of the taxonomy and affi?-lt1es of
T aeniopteris in New Zealand, using some methods developed SInce the
time of Arber's pioneer studies.

METHODS

Fossils used in this study were collected from the northern end of Huriwai
Beach near Port Waikato, South Auckland (fossil locality no. N51/f659),
and from Haast Gully in the Clent Hills, central Canterbury (fossil locality
no. S81/f569) (Fig. 1).
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The best-preserved remains are found in fine-grained, well-consolidated

sandstones at both localities, but a range of lithologies from massive coarse
conglomerates to thinly-bedded mudstones occurs. The abundance and variety
of plant remains in both areas suggest lagoonal, deltaic or near-shore
deposits.
An attempt was made to obtain epidermal cuticles from leaf remains at
Port Waikato, where the youngest and presumably least metamorphosed
deposits of the New Zealand Geosyncline are located. Comparative study
of epidermal and stomatal characters impressed in the cuticle is very useful
in taxonomic study of both fossil and modern plants (Harris 1956; Stace
1965).
Direct examination of a few leaves in low angle reflected light showed
very faint outlines of epidermal cells (Fig. 2). However, a wide variety of
chemical methods were all unsuccessful in isolating cuticles (Blaschke 1974).
Some of these methods did successfully recover spores and pollen and should
have isolated cuticl~s had they been preserved. The most plausible cause of
loss of cuticle is subsurface weathering and oxidation in both upper Mesozoic
and post-Miocene times. There is, in fact, one published description of seed
cuticles of the fruit Carnoconites Srivastava emend. Sahni from Port Waikato
(Harris 1962), although seed cuticle may be more resistant than leaf cuticle
(Harris 1956). It is possible that small recoverable amounts of cuticle are
present in the finest siltstones but they are unlikely to be well enough
preserved for reliable taxonomic work.
In the absence of anatomical information, the study proceeded by measure-
ment and analysis of macro-morphological characters. These included length,
width and shape of the leaf, details of leaf apex, base, midrib and petiole,
and the frequency, angle, and type of furcation of the secondary veins. In
practice, only a few characters were found suitable for population analysis,
notably leaf and apex shape, and lamina width. Fewer than 5% of the
large number of leaves examined (c. 360) were preserved entire, and these
probably represented a variety of plant ages. For these two reasons, overal~
size was not a particularly useful character. Other characters such as details
of venation were not significantly variable between populations.
Simple statistical analysis of the measurements made-for example,
character correlations and frequency distributions of measurements for single
 No.6 BLASCHKE & GRANT-MACKIE - T aeniopteris 935
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PAKAWAU

CHRISTCHURCH

MT POTTS

o 100km
'-----'---..J

§ MURIHIKU SUPERGROUP

~ TORLESSE SUPERGROUP

FIG.I-Plant-fossil collection points mentioned in the text, and the distribution, both
exposed and subsurface, of Murihiku and Torlesse Supergroups in New Zealand.
 936 N.Z. JOURNAL OF GEOLOGY AND GEOPHYSICS VOL. 19

and combined populations-enabled species delineation to be based upon

population characteristics rather than individual specimens. This reduces the
uncertainty inherent in basing taxonomic conclusions exclusively upon rather
variable gross morphological data.

SYSTEMATICS

Port Waikato Population

Taeniopteris daintreei McCoy Figs 2, 3, 4

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1850 T. spatulata McClelland, Rep. Geol. Surv. India for 1848-9, p. 53.
1874 T. daintree; McCoy, Prodromus of the Paleontology of Victoria, p. 15.
1917 T. arctica Heer; Arber, N.z. Geol. Surv. Pal. Bull., 6, p. 44.
1934 T. spatulata McClelland; Edwards, Ann. Mag. Nat. Hist., Ser. 10, 13, p. 97.
1969 T. daintree; McCoy; Douglas, Mem. Geol. Surv. Victoria, 28, p. 53. (gives
additional synonomy)
HOLOTYPE: National Museum of Victoria, P12270 (McCoy 1874)
DESCRIPTION: Leaf simple, forate or linear oblong; size variable, length usually
greater than 60 mm, occasionally greater than 150 mm; width usually 3-13 mm,
occasionally up to 25 mm. Apex usually acute, very rarely acuminate, occasionally
obtuse; base acutely decurrent; margin entire. Prominent longitudinally ridged midrib,
usually 1;\;-%. width of lamina, less than lfl'O width of lamina when the latter more than
10 mm wide; narrowing in distal third of leaf. Secondary veins prominent, arising
from midrib at right angles or nearly so; sometimes simple, usually forking once, more
rarely twice, at variable distance between midrib and margin. Vein frequency O' 8-3' 2
(mean = 1· 98) per mm at midrib, 1· 6-3·4 (mean = 2·48) per mm at margin. Petiole
stout, up to 20 mm, occasionally 30 mmlong, sometimes faintly winged; base expanded,
occasionally rounded. Anatomicar details unknown.
DISCUSSION: Only one species of T aeniopteris is recognised from this
locality, although the population is variable. Arber (1917) designated the
Port Waikato species T. arctica Heer ( = Oleandra arctica Heer), a species
described from the Cretaceous of Greenland (Heer 1874). However, des-
cription and illustration of the Greenland species show greater leaf length
and greater frequency of furcation of secondary veins than do the Port
Waikato specimens; furthermore the possible presence of sori in several
illustrations of the former suggests that it may not belong to T aeniopteris
at all. It should be noted that Arber's (1917) allocation of the Port Waikato
population to T. arctica was based on a specimen subsequently found to
have come from upper Cretaceous strata of Pakawau, Nelson (McQueen
1955) .
The characteristics of the Port Waikato specimens conform very well to
those described for T. daintreei from Victoria (Douglas 1969), and close
similarity between floras of these two areas is more in keeping with known
geologic relations than is Arber's suggestion of identity with an Arctic
Cretaceous form. The New Zealand hypotype of T. arctica (from the upper
Cretaceous at Pakawau) also fits into the description of T. daintreei from
Victoria, but population studies allow separate recognition of the Pakawau
form as T. stipulata Hector (McQueen 1956).
 No.6 BLASCHKE & GRANT-MACKIE - Taeniopteris 937
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FIGS 2-4-Taeniopteris daintreei McCoy, from Puaroan of Port Waikato (N51/f659,

AU 1293). 2-Portion of leaf showing outlines of epidermal cells and possible
stoma (arrowed); approx. X 33. 3-Well-preserved incomplete leaves showing
typical form of apex and ba~e; note short petiole.4-Complete small leaf.
FIGS 5, 6-Taeniopteris thomsoniana Arber, from middle to upper Jurassic (Temaikan
to Puaroan) of the Clent Hills (S81/f569, AU 1296). 5-Leaf showing obovate
form.

Geology-I 6
 938 N.Z. JOURNAL OF GEOLOGY AND GEOPHYSICS VOL. 19

Clent Hills Population

Taeniopteris thomsoniana Arber Figs 5, 6

1913 T. daintreei McCoy; Arber, Proc. Roy. Soc. London, 86B. p. 340.
1917 T. thomsoniana Arber, NZ. GeoJ. Surv. Pal. Bull., 6, p. 47.
HOLOTYPE: British Museum (Natural History) (Arber 1917).
EMENDED DESCRIPTION: Leaf simple, narrow oblanceolate to oblanceolate; length of
mature leaves usually greater than 100 mm; width ranges from 5-25 mm, usually
10-22 mm. Apex obtuse; base acutely decurrent; margin entire. Midrib prominent,
usually less than lfro width of lamina. Secondary veins usually arising at more than
80° to midrib, slightly more acute near apex. Approximately 60% of the veins
dichotomise once, occasionally twice, the first dichotomy usually near midrib, second
dichotomy near margin; anastomoses very rarely present. Vein frequency 1·1-2·8
(mean = 1· 82) per mm at midrib; 1· 8-3·6 (mean = 2·67) per mm at margin.
Petiole stout, 0-40 mm long, hardly wider than midrib except at triangular base.
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Anatomical details unknown.

DISCUSSION: The Clent Hills population appears to be taxonomically
distinct from that at Port Waikato. Visible differences include overall size
and shape; moreover the frequency distribution of leaf-width measurements
of both populations combined and given equal weighting generates a
distinctly bimodal curve (Fig. 7).
It should be noted that the above description of T. thomsoniana is con-
siderably expanded from that of Arber (1917), taking into account a greater
range of variation than was described or figured by him.

STRATIGRAPHY

The age range for Gondwanan occurrences of T. daintreei is middle

Jurassic to Albian (Douglas 1969). The most recent microfloral dating of
the Port Waikato beds (middle to upper Tithonian, possibly lowest
Cretaceous, Waterhouse & Norris 1972) thus places T. daintreei from this
locality well within its established time range.
The Clent Hills deposits are thought to be Temaikan or younger. Although
extensive faulting complicates the stratigraphy, probable Puaroan bivalves
have been found in the vicinity of some plant-bearing localities in the area
(P. J. Oliver, pers. comm.). Arber also recorded T. thomsoniana from Tank
Gully near Mount Potts, some 25 km to the south-west of Clent Hills. The
presence of the species at both these localities has posed a problem in
accepting a middle to upper Jurassic age for the Clent Hills beds, as the
Mount Potts beds are overlain by Kaihikuan marine faunas (Campbell &
Forc.e 1972), and this would imply a surprisingly long time range for this
speCles.
However, recent revisits to Tank Gully and examination of New Zealand
Geological .Survey collections from Tank Gully have failed to turn up
T. thomsomana (G. Retallack, pers. comm.). It thus appears very likely that
Arber's material from Tank Gully and Clent Hills was mixed in transit, and
that T. thomsoniana is restricted to the middle to upper Jurassic beds of the
Clent Hills.
 No.6 BLASCHKE & GRANT-MACKIE - Taeniopteris 939

15

14

13 for Port Waikoto population

X = 6·13mm
12
0' = 3·03mm
11 for Clent Hills population
.!l! X =1399mm
0-10 d =3 86mm
E
ffi 9
aQ)
8
g> 7
E
13 6
~ 5
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20 22 24

FIG. 7-Combmed width analysis of Port Waikato and C1ent Hills T aenio pteris
246
populations. C1ent Hills (CH) class totals have been multiplied by - - to give
119
equal weighting with the Port Waikato (PW) sample. The width distribution
curve distinctly shows the presence of two different populations, with that from
C1ent Hills having a wider lamina, suggesting the existance of different taxa.
(The measured samples consisted of 246 specimens from Port Waikato, and 119
from C1ent Hills.)

Acceptance of the approximate contemporaneity of T. daintreei at Port

Waikato and T. thomsoniana at Clent Hills emphasises a provenance separa-
tion of the two species. The latter is confined to Torlesse rocks of the New
Zealand Geosyncline whilst T. daintreei is found predominantly, perhaps
only, in the Murihiku facies; and this provides another possible indication
of different provenances for Murihiku and Torlesse strata. (T. daintreei was
also recorded by Arber (1917) from the Malvern Hills; the specimen figured
is not clear enough for identification to be verified but the possibility that
T. daintreei occurs also in Torlesse rocks must for the time being be
accepted.)
Neither T. thomsoniana nor T. vittata have been recorded from Australia,
although in general there is a fairly marked similarity between New Zealand
and Australian middle Mesozoic floras (Jones & de Jersey 1947; Medwell
1954). Further analyses of Mesozoic floras may contribute towards the
solution of some of the outstanding problems of the New Zealand Geo-
syncline. Differences in taxonomic composition between Torlesse and Muri-
hiku floras, such as are suggested here, would lend significant support to
claims of derivation of the two sets of strata from different sources (Force
1974) ; whilst careful analysis of the affinities of these floras might indicate
more clearly than do marine biotas, where those sources lay.
 940 N.Z. JOURNAL OF GEOLOGY AND GEOPHYSICS VOL. 19

CONCLUSION

The botanical affinities of New Zealand T aeniopteris are of some

significance. On occasion over the last hundred years, several Australian and
New Zealand species, notably T. daintreei, have been identified as T.
spatldata, a common form in the middle to upper Jurassic Rajmahal Series of
India. Sahni (1948) relocated T. spatldata in Nipaniophyllum, the leaf form-
genus of a new order of gymnosperms, the Pentoxylales. This small group is
of considerable botanical interest. firstly because it is the only gymnosperm
order confined to the Southern Hemisphere, also uniquely confined to the
Jurassic, and secondly because its anatomy shows features common to two
important gymnosperm orders, the Bennettitales and the Cycadales (Vishnu-
Mittre 1957), as well as to the Palmales in the angiosperms. The Pentoxylales
have been implicated by some authors as a possible group from which the
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angiosperms arose (Meeuse 1961).

There are indeed some similarities between Nipaniophyllum from the
Rajmahal Hills in India and T. daintreei from Port Waikato. They share not
only a general morphological similarity, but a notably similar range of varia-
tion at the two localities. More significantly, a Pentoxylalean fruit, Carnocon-
ites cranwellii Harris, has been collected on one occasion from Port Waikato
(Harris 1962). This find constitutes the only substantiated Pentoxylalean
fossil outside India.
Unfortunately, except for the superficial morphological similarities which
are of the kind that has led to so many paleobotanical errors in the past, there is
no real evidence to conclude that T. daintreei and C. cranwellii came from the
same plant. One cannot ignore the fact that T aeniopteris is so abundant at
Port Waikato whilst only a single Carnoconites and no Pentoxylalean stems
have ever been found there, despite repeated searches by Lucy M. Cranwell
and others. Furthermore, Douglas (1969) has shown quite clearly that there
are several anatomical differences between T. spatulata in India and T.
daintreei in Victoria. Although evidence of this is not directly available in
New Zealand because of the poorer preservation of the fossils, it is believed
that unless clear Pentoxvlalean affinities for T. daintree; from Port Waikato
can be shown, Carnuconites and T. daintreei should not be aligned with each
other. Nevertheless, there remains the temptation to do so and the possibility
that such association was real, and it is suggested that this is an aspect of New
Zealand paleobotany which would be most rewarding from a botanical
standpoint.
ACKNOWLEDGMENTS
This study was done in 1974 as part of a B.Sc. Honours project by P. M. Blaschke
under the joint supervision of Drs]. A. Grant-Mackie and]. E. Braggins. We ar~
grateful to ]. F. Rigby, P. J. Oliver, and R. O. Gardner for comments on the manuscript.

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New Zealand. (Unpublished B.Se. Hons. thesis, lodged in the Library
University of Auckland.) 59 p. '
 No.6 BLASCHKE & GRANT-MACKIE - T aeniopteris 941

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