Palaeogeography, Palaeoclimatology, Palaeoecology 184 (2002) 321^345
www.elsevier.com/locate/palaeo
Quantitative palaeoclimate estimates from Late Cretaceous
and Paleocene leaf £oras in the northwest of the South Island,
New Zealand
Elizabeth M. Kennedy a; , Robert A. Spicer b , Peter M. Rees c
a
Institute of Geological and Nuclear Sciences, Ltd., P.O. Box 30-368, Lower Hutt, New Zealand
Department of Earth Sciences, The Open University, Walton Hall, Milton Keynes MK7 6AA, UK
Department of the Geophysical Sciences, University of Chicago, 5734 South Ellis Avenue, Chicago, IL 60637, USA
b
c
Received 2 July 2001; accepted 22 March 2002
Abstract
Three new plant macrofossil assemblages were collected from Late Cretaceous and Paleocene fluvio-lacustrine
sediments of the Pakawau and Kapuni groups in the northwest of the South Island, New Zealand.
Palaeoenvironmental interpretations were made from each locality and palaeoclimate was deduced from the
dicotyledonous angiosperm leaf component of each flora. A latest Cretaceous (Pakawau Bush Road locality) flora
yielded 58 different dicotyledonous leaf forms; the two Paleocene collections, Ian’s Tip and Pillar Point Track,
included 23 and 28 dicotyledonous leaf forms respectively. Quantitative palaeoclimate estimates were obtained using
both Leaf Margin Analysis (LMA) and the Climate Leaf Analysis Multivariate Program (CLAMP). Temperature
estimates suggest that there was a slight cooling from the latest Cretaceous into the early Paleocene in the northwest
Nelson region of New Zealand, supporting similar Southern Hemisphere palaeoclimate findings from Antarctic data.
Consistency in temperature estimates using different methods, including LMA, multivariate leaf morphological
analysis (CLAMP), oxygen isotope data, regional versus local studies and global palaeoclimate models, suggests that
the mean annual temperature for the Pakawau region in the latest Cretaceous was between 12 and 15‡C. LMA
produced temperature estimates between 6.5 and 8‡C for the two Paleocene assemblages whereas CLAMP-produced
estimates were slightly higher between 9 and 12.5‡C ; 2002 Elsevier Science B.V. All rights reserved.
Keywords: palaeoclimate; New Zealand; Late Cretaceous; leaf physiognomy; Paleocene
1. Introduction
Although New Zealand has signi¢cant leaf £o* Corresponding author. Tel.: +64 (4) 5704838;
Fax: +64 (4) 5704600.
E-mail addresses: e.kennedy@gns.cri.nz (E.M. Kennedy),
r.a.spicer@open.ac.uk (R.A. Spicer), rees@geosci.uchicago.edu
(P.M. Rees).
ras from non-marine sediments of Late Cretaceous^Tertiary age, relatively little time has been
devoted to their study. Predominantly non-marine
sequences in western South Island have yielded
well-preserved, although often fragmentary, leaf
fossils. These assemblages provide valuable information on New Zealand Cretaceous^Tertiary terrestrial palaeoenvironments, a subject which currently represents a considerable knowledge gap.
0031-0182 / 02 / $ ^ see front matter ; 2002 Elsevier Science B.V. All rights reserved.
PII: S 0 0 3 1 - 0 1 8 2 ( 0 2 ) 0 0 2 6 1 - 4
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Understanding the geological record of our terrestrial £oras, environments and climates is becoming increasingly important given the current focus
on present climate change, as well as questions
relating to the modern New Zealand biota and
changes in biodiversity.
By the end of the Cretaceous, the New Zealand
landmass had separated from both Australia and
Antarctica and was developing its own unique
biota (Stevens et al., 1988). Palaeogeographic reconstructions place the NW Nelson region at 50^
60‡S in the latest Cretaceous and early Paleocene
(Paleogeographic Atlas Project, University of Chicago), and the landscape was generally of low
relief (Fleming, 1979; LeMasurier and Landis,
1996) (Fig. 1).
This paper presents an outline of three localised
fossil £oras (de¢ned here as being collected from
three single localities) from the northwestern-most
part of the South Island (Fig. 2) and the palaeoclimate interpretations that were derived from
them. Sedimentary rocks of the Pakawau and Kapuni groups in northwest Nelson are the only
Late Cretaceous and Paleocene onshore representatives of a thick sequence of non-marine and
marginal marine strata at the southern end of
the hydrocarbon-producing Taranaki Basin. Leaf
Fig. 1. Paleogeographic reconstruction of the New Zealand
landmass at the end of the Cretaceous (ca. 65 Ma). Reconstruction by King et al. (1999). Coal measure deposition is
indicated by dashed line ¢ll and dotted ¢ll indicates shoreline
sandstone deposition.
fossils have been found in both of these groups
and provide insights into the ancient vegetation
and environments that produced the economically
important reserves of gas and oil.
We used leaf morphology approaches - Leaf
Margin Analysis (LMA; e.g. Wolfe, 1971) and
Climate Leaf Analysis Multivariate Program
(CLAMP; Wolfe, 1993) ^ as our primary sources
of climate parameter estimates as well as comparison with published and unpublished oxygen isotope data and general circulation models (e.g.
Valdes, 2000; Valdes et al., 1996).
2. Late Cretaceous £ora: Pakawau Bush Road
This is an ideal £ora for palaeoclimate analysis
because of its high diversity of dicotyledonous
angiosperms, which dominate the assemblage.
Based on palynological zonation, this site is dated
as the New Zealand Haumurian stage (Wellman,
1959; amended by Crampton et al., 2000), which
had been considered as broadly equivalent to the
international Maastrichtian stage. However, a recent re-evaluation of the New Zealand time scale
has revised the stratigraphic extent of the Haumurian to include the Campanian and the top
of the Santonian (Crampton et al., 2000). The
palynological zone to which this site was assigned
is PM2 (Raine, 1984). This restricts the age to the
mid to upper part of the Haumurian, from about
77^65 Ma (i.e. late Campanian to Maastrichtian).
Late Cretaceous dino£agellate stratigraphy of
New Zealand is more detailed (Roncaglia and
Schioler, 1997), but no dino£agellates were found
in the palynological samples used to date the Pakawau Bush Road macro£ora. Taxonomic studies
on other PM2 palynological zone leaf £oras in
New Zealand include those made by von Ettingshausen (1887), Edwards (1926), McQueen (1955,
1956), Mildenhall (1968) and Pole (1992).
Fossil specimens were collected from an outcrop of the thick, predominantly terrestrial, sediments of the Rakopi Formation (Thrasher, 1990).
This outcrop consists of approximately 20 m of
vertical sequence dipping at about 10‡ to the west
(Fig. 3). Useful fossil material was collected from
two distinct units (lower (PBL) and upper (PBU))
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323
Fig. 2. Locality map for the three study sites in the Pakawau area. The column (after Thrasher, 1992) shows inferred stratigraphic positions of the fossil sites within the regional geology. Pakawau and Kapuni group sediments in the collection area are Late
Cretaceous and Tertiary in age and range from the non-marine coal measures of the Rakopi Formation into marginal marine
sandstones of the North Cape Formation and back to the non-marine £uvial deposits of the Farewell Formation. Stars suggest
relative positions of the fossil localities within this general stratigraphy, however the column is not to scale and exact stratigraphic positions of the fossil localities, with respect to the formation boundaries, are unknown because of poor outcrop exposure.
The following are New Zealand Fossil Record File numbers and grid references for the three fossil localities: Pakawau Bush
Road (PBR) ^ FRF no. M25/f104, grid ref. NZMS 260 M25 818 689; IT ^ FRF no. M24/f52, grid ref. NZMS 260 M24 855
772; PPT ^ FRF no. M24/f24, grid ref. NZMS 260 M24 853 775.
separated by carbonaceous mudstone and thin
coals. At the base (eastern end) of the exposed
section is a coarse white sandstone with thin
bands of matrix-supported pebbly conglomerate.
Overlying this is the ¢rst of the fossil-bearing
units (PBL), a white siltstone to ¢ne sandstone
unit with infrequent visible bedding and sparse
plant fossil layers. Overlying PBL is at least 5 m
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of carbonaceous mudstone with thin coals. This
carbonaceous mudstone unit grades into the second plant fossil unit (PBU) which ranges from silt
to medium sandstone and, in contrast to PBL, has
numerous distinct bedding planes. Overall, there
is a coarsening upwards trend throughout PBU
from where the underlying coal and mudstone
unit grades into the carbonaceous clay-rich silt
and ¢ne sandstone of the base of PBU to a medium sandstone near the top of the exposed unit.
However, texture is variable within the unit. At
the top of the section is a second carbonaceous
mudstone and coal unit, but the contact between
this and the underlying PBU is not visible.
Interpretation of the depositional environment
is based on sediment texture, structure of the
units, coal analysis, quality of fossil preservation
and composition of the fossil assemblage. The depositional setting was most likely an ephemeral
lacustrine environment within a vegetated £oodplain. Lacustrine conditions dominated at the site
of deposition, but they alternated and were at
times contemporaneous with mire development
at the lake margins. There was also a £uctuating
£uvial in£uence that was more proximal during
deposition of PBL sediments.
The channel deposition of the coarse sandstone
unit below PBL was gradually replaced by overbank deposition resulting in the PBL plant-bearing unit. The £uvial source migrated further away
and the depositional environment developed as a
marginal lacustrine setting with carbonaceous
mudstone and peat swamp sedimentation. Increasing input of clastic sediments caused peat deposition to cease, with a gradual return to a more
£uvially in£uenced setting. The basal 2 m of the
upper fossil unit (PBU) are carbonaceous and relatively species-poor, containing relatively fewer
angiosperm leaf types, but with abundant remains
of a plant that may be an aquatic fern. There is
also a dominance of small-leafed species in the
lower part of this unit. We infer that conditions
were generally nutrient-poor and therefore detrimental to plant growth, consistent with a mire
environment. As mire in£uence decreased, species
diversity increased, and the arbitrary collecting
divisions of PBU show this pattern of change
well. The lowest two divisions, PBU1 and
PBU2, yielded six and seven dicotyledonous
forms respectively. Diversity increased to 12 forms
in PBU3 and 20 in PBU4. Divisions PBU5 and
PBU6 had lower diversity (11 and nine forms respectively), but the number of dicotyledonous leaf
forms increased again in the two highest collecting
divisions of PBU (PBU7 and PBU8, with 18 and
15 leaf forms respectively).
The leaf £ora is predominantly angiospermous
with minor components of podocarps, araucarians and ferns. With the exception of one probable monocotyledonous leaf form, the angiosperm
leaves are dicotyledonous. Leaf fossils collected
from the horizons in this sequence (Fig. 3) were
amalgamated for this palaeoclimate study, but
there is potential for subdividing the £ora for
Fig. 3. Generalised sequence at the Pakawau Bush Road locality. PBU1 to PBU8 are arbitrary 1 m divisions made for
collection purposes.
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more detailed palaeoclimate investigation. Fiftyeight di¡erent dicotyledonous leaf morphotypes
have been distinguished, with a total of 858 specimens assigned to these form groupings (Kennedy,
1993) (Figs. 4, 7a^i). Although these leaves have
not yet been formally described, a number of different dicotyledonous families appear to be represented including the Lauraceae, Proteaceae and
Fagaceae. Forty-three percent of these leaf morphotypes have entire margins. Other plant material includes a striking collection of over 100
specimens of a pentamerous £ower (Kennedy et
al., 1999) and several di¡erent types of seeds.
Faunal discoveries are restricted to two small
specimens of the freshwater bivalve Hyridella
(e.g. Winterbourn, 1973) and evidence of insect
damage on some of the leaves.
We conclude that the assemblage is predominantly locally derived. This is based on the depositional setting as a whole ; the presence of almost
complete specimens of some of the more delicate
leaf types; and the presence of £owers, many with
perianth still attached. Studies have found that
most leaves, particularly fragile ones, are unlikely
to be transported more than a few kilometres before disintegrating (Rich, 1989). It has also been
suggested that £uvio-lacustrine depositional environments are generally representative of the local
vegetation (Greenwood, 1991).
3. Early Paleocene £oras
3.1. Ian’s Tip (IT)
The preservation quality of the leaf fossil material is variable, and many of the specimens are
of a fragmentary nature. The leaves are mostly
preserved as impressions. Twenty-three morphotypes were determined from the dicotyledonous
leaf specimens (Figs. 5, 7j^m) and these were described for palaeoclimate analysis (Kennedy,
1998). Two of these leaf morphotypes probably
have Lauraceous a⁄nities (Fig. 5b,c). Another
form (Fig. 5e) is attributable to Banksiaeformis
Hill and Christophel (1988), a genus proposed
for specimens with the appearance of the modern
genera Banksia and Dryandra but which cannot
325
be assigned to the genus Banksiaephyllum due to
lack of cuticle. The IT leaf form closely resembles
a Proteaceous species described by von Ettingshausen (1887), Dryandra comptoniaefolia Ett.,
from the Paleocene of inland Canterbury, as
well as a leaf form (TARA-34) described by
Pole (1997) from Kakahu, a Paleocene locality
in South Canterbury. Pole suggested that the Kakahu leaf form was the same taxon as D. comptoniaefolia Ett. Other distinctive forms include a
compound leaf with narrow, serrated lea£ets (Fig.
5f), and a palmate leaf which has very variable
leaf shape (Fig. 5a).
Exposure is limited and sedimentary structures
are unclear at this locality. Many of the leaves
came from loose blocks within road-cut debris,
although further excavation revealed that plantbearing sediment (generally poorly bedded siltstone to ¢ne/medium sandstone) was also present
in situ. There is also some evidence of disrupted,
possibly rapid, deposition where leaves were
found at varied orientations within the sediment.
An erosional contact with a coarser white sandstone at the top of the outcrop indicates a change
to more proximal channel in£uence. This is all
supportive of deposition adjacent to a £uvial
channel, perhaps overbank deposition. A £uvial
setting is also consistent with palaeoenvironmental interpretations for the Farewell Formation as
a whole, with braided stream (Titheridge, 1977) or
coarse-grained meandering river (Bal, 1994a,b)
environments recognised.
In addition to the angiosperm leaves, which
make up most of the £ora, podocarp fragments
were collected, together with a possible fern fragment. Woody material is quite common, with
fragments ranging in size from small twigs to
small logs of up to V7 cm diameter. The collected assemblage also contains at least ¢ve unidenti¢ed di¡erent types of seeds.
The IT locality is situated in the vicinity of the
Cretaceous^Tertiary (K/T) boundary. However
the precise location of the K/T boundary has
not been located here due to poor outcrop exposure. Sedimentary rocks at the abandoned Puponga Coal Mine (a few hundred metres southeast along the road from IT) have been dated
as Haumurian (late Campanian^Maastrichtian).
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Fig. 4. Dicotyledonous leaf forms from the latest Cretaceous Pakawau Bush Road locality. Scale bars are 1 cm.
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Fig. 4 (Continued).
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328
E.M. Kennedy et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 184 (2002) 321^345
Fig. 4 (Continued).
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Fig. 4 (Continued).
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E.M. Kennedy et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 184 (2002) 321^345
Fig. 5. Dicotyledonous leaf forms from Ian’s Tip (IT), a Paleocene locality from Northwest Nelson, New Zealand. Scale bars are
1 cm.
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331
Fig. 5 (Continued).
Although IT itself has not yet been dated, a sample from a road-cutting between IT and Puponga
Mine, that is almost certainly stratigraphically below IT, has been dated as Paleocene (Ian Raine,
pers. comm., 2001). A sequence of samples along
the track to Pillar Point lighthouse, including the
Pillar Point Track (PPT) leaf locality which is
stratigraphically above IT, also produced Paleocene palynomorph assemblages (Raine, 1989).
3.2. Pillar Point Track PPT
It is likely that this locality produced the
youngest of the three £oras outlined in this paper.
Although the lack of continuous outcrop in the
area makes it di⁄cult to establish relative stratigraphic positions, this locality appears to lie
stratigraphically above IT. Palynological analysis
indicates PPT is Paleocene (Raine, 1989), falling
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Fig. 6. Dicotyledonous leaf morphotypes from the Paleocene Pillar Point Track (PPT) locality. Scale bars are 1 cm. Note that
PPT11 was not included in any analyses due to poor preservation of characters useful for palaeoclimate analysis.
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Fig. 6 (Continued).
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E.M. Kennedy et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 184 (2002) 321^345
Fig. 7. Selected leaves from Pakawau Bush Road (a^i), Ian’s Tip (IT) (j^m) and Pillar Point Track (PPT) (n^r). Scales are 1 cm2 .
(a) PB1, (b) PB9, (c) PB12, (d) PB13, (e) PB20, (f) PB26, (g) PB48, (h) PB 52, (i) PB38, (j) IT2, (k) IT4, (l) IT5, (m) IT11,
(n) PPT1, (o) PPT2, (p) PPT6, (q) PPT13, (r) PPT19.
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Fig. 7 (Continued).
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E.M. Kennedy et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 184 (2002) 321^345
into the PM3 palynological zone of Raine (1984)
and it is part of the Farewell Formation. The site
exposure has a lateral extent of approximately
5 m. These fossiliferous sediments are part of a
siltstone-to-¢ne-sandstone lens (or possibly a ¢ner
basal facies) within the predominantly conglomeratic sediments that make up the Farewell Formation in this area.
The generally ¢ne sediments, their laminated
nature, and the abundant, densely covered leaf
layers of the PPT locality suggest a lower energy
environment than that at IT. The presence of several examples of partially intact compound leaves
supports this interpretation, and a minor shallow
lacustrine situation is suggested, such as ponding
in a £oodplain environment. However, thin layers
of coarser sandstone within the ¢ne sediments
(siltstone and ¢ne sandstone) provide evidence
for a still proximal channel in£uence. This interpretation also concurs with the general £uvial depositional setting inferred for the Farewell Formation (e.g. Titheridge, 1977).
Twenty-eight dicotyledonous leaf morphotypes
have been described from the PPT site (Figs. 6,
7n^r). Only 27 of these were used for palaeoclimate analysis because PPT11 (Fig. 6o) does not
have enough morphological information preserved for inclusion in the analyses (although it
does have su⁄ciently distinctive venation to enable establishment of a separate form). The collection also includes podocarp shoot fragments, several di¡erent types of seeds, woody fragments and
a few specimens of a small £ower. The often
densely covered leaf layers meant that there was
some di⁄culty in distinguishing margins of individual leaves, nevertheless there were su⁄cient
specimens measured to enable meaningful palaeoclimate interpretation.
4. Quantitative leaf-based palaeoclimate analysis
Plants are ideal biological climate ‘recorders’.
Once they have germinated they are ¢xed in place
for their life cycle and therefore in order to survive they must be adapted to the environment,
including climate, in which they are growing.
The basis behind using leaf morphology to inter-
pret palaeoclimates is that many aspects of the
architecture of leaves represent an adaptation to
the particular climate under which they were
growing, and this architecture, preserved in the
fossil record, therefore re£ects ancient climate.
The quantitative palaeoclimate methods used
here were LMA (e.g. Wolfe, 1971) and the
CLAMP of Wolfe (1993). Both of these methods
utilise the relationships observed between leaf
morphology and climate in the modern environment. LMA is based on a positive correlation
observed in some modern £oras between mean
annual temperature (MAT) and the percentage
of leaf forms in a £ora that have entire margins.
This method originated from the observations of
two researchers early last century, Bailey and Sinnott (1915, 1916).
CLAMP is a more recent development on the
leaf margin/temperature correlation (Wolfe, 1993,
1994). It builds on the correlation used in LMA
by adding more leaf morphological characters to
the analysis and applying multivariate statistical
methods. CLAMP includes a number of di¡erent
character states (including aspects of the margin,
leaf size, lamina shape, apex and base shape) and
uses these to produce estimates for both temperature and precipitation. However, the temperature
estimates are more reliable than the precipitation
estimates, which tend to have high quanti¢able
uncertainties. CLAMP works by simultaneously
analysing a data set of modern leaf morphological
character states and a corresponding data set of
modern climate information using multivariate
analysis, in this case Canonical Correspondence
Analysis (CANOCO, ter Braak, 1986). In other
words, 31 characters are measured for a modern
leaf species, and this is repeated for all species
collected from a sample site. Sites with recorded
long-term meteorological data were chosen for
leaf collections and analyses.
Leaf morphological information from a fossil
£ora can then be included in the modern data
set of leaf £oras. By including it in the analysis,
the fossil £ora can be assigned a position relative
to those £oras in the modern, or predictor, data
set. The climate variables in the meteorological
data set are also assigned relative positions within
the cloud of points representing leaf £oras, and
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vectors for these climate parameters can be added.
Palaeoclimate estimates are then obtained by projecting the position of the fossil £ora on to each
of the climate variable vectors calibrated using the
modern data set. Although this method can produce acceptable results, as with LMA, the underlying plant physiology responsible for some of the
observed correlations between leaf morphology
and climate is still relatively little understood, particularly the function of teeth on the lamina (e.g.
Wolfe, 1993).
The advantage of leaf morphological methods
of palaeoclimate analysis over those that relate
modern systematic groupings and their climate
tolerances back to fossil £oras ^ Nearest Living
Relative methods (NLR) ^ is that they do not rely
on correct identi¢cation of fossil leaves to modern
taxa. Leaf morphotypes need to be distinguished
from one another but species relationships to extant taxa need not be determined. Moreover,
NLR methodology is fundamentally compromised by the evolutionary process. Leaf morphological methods do not rely on the assumption
made by NLR methodologies that a species’ environmental tolerances remain static over time. A
better approach is to exploit the repeated convergence of foliar physiognomy and climate that can
be demonstrated over geologically signi¢cant
spans of time.
A detailed discussion of the strengths and
weaknesses of using quantitative methods such
as LMA and CLAMP will not be presented
here. The purpose of this paper is simply to report
palaeoenvironmental, particularly palaeoclimate,
¢ndings from these three fossil leaf £oras and to
discuss them in the context of Late Cretaceous
and early Tertiary climate patterns on a New Zealand-wide, as well as global, scale. However, it is
not our intention to gloss over any uncertainty
typically associated with methods that attempt
to estimate climate data from ancient £oras.
Thus, the uncertainty measures stated on estimates should be considered minimum values.
LMA palaeoclimate estimates for the New Zealand £oras are shown in Fig. 8 and Table 1. These
were calculated from previously published regressions (Wolfe, 1979; Wing and Greenwood, 1993).
CLAMP percentage scores for the £oras are
337
Fig. 8. Schematic plot of Northern and Southern Hemisphere
gradients used in LMA based on Wolfe (1979) and Wolfe
and Upchurch (1987). The Northern Hemisphere gradient
(black line) was established by plotting the percentage of entire-margined species against MAT for east Asian £oras.
Wolfe inferred a Southern Hemisphere gradient (grey line)
from more limited data. The labelled symbols on the gradients show the positions of the three £oras from Pakawau ^
Pakawau Bush Road (PBR), IT and PPT.
shown in Table 2 and estimates are presented in
Table 1 from CLAMP analysis using three di¡erent data sets of modern £oras. Most existing
CLAMP predictor (modern) data sets are made
up of Northern Hemisphere £oras (Wolfe, 1993;
Spicer et al., 1996). In this paper, a 101-site predictor data set is used in the analyses. This data
set is based on the 103-site data set used by Herman and Spicer (1997) which includes signi¢cant
changes to the original published data set (Wolfe,
1993), such as the removal of sub-alpine sites and
the addition of samples from Mexico, collected
after the CLAMP method was published. Sub-alpine sites are de¢ned as those where the mean
temperature of the warmest month (WMM) is
less than 16‡C and the mean temperature of the
coldest month is less than 3‡C (Wolfe, 1993). The
inclusion of these sub-alpine samples can produce
erroneous results if they are not appropriate modern analogues for the fossil sites being analysed.
The 144- and 170-site data sets shown in Table
1 contain additional sites more recently collected
for use in the CLAMP method by Wolfe and
colleagues. Substantial additions include more recent data from Japan and the USA (Wolfe, 1997).
PALAEO 2871 25-7-02
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Table 1
Climate estimates from both CLAMP and LMA for the Pakawau £oras
Age (from palynology)
Analysis
Uncertainty
Pakawau Bush Road
Haumurian (PM2*)
% entire margins (as de¢ned in Wolfe,
1993)
MAT (‡C) (LMA)
MAT (‡C) (LMA)
MAT (‡C) (CLAMP)
WMM (‡C) (CLAMP)
Mean growing season precipitation
(CLAMP)
Growing season length (CLAMP)
44.8
IT
Teurian
(PM3*)
21.7
PPT
Teurian
(PM3*)
19.2
SH scale
NH scale
101 31
144 31
170 31
101 31
144 31
170 31
101 31
NA
S 0.8‡C (standard error)
S 1.8‡C
S 1.9‡C
S 2.8‡C
S 3.1‡C
S 3.0‡C
S 3.4‡C
S 234 mm
13.5
14.8
13.8
12.7
12.3
20.5
22.0
22.2
971
7.0
7.8
11.1
12.3
11.3
19.3
21.5
21.5
777
6.5
7.0
9.2
10.5
9.1
18.8
20.5
19.7
494
144
170
101
144
170
S 485 mm
S 452 mm
S 1.1 months
S 1.0 months
S 1.3 months
955
1032
8.0
7.4
7.4
1899
1795
6.8
7.4
7.0
1342
1223
5.8
6.5
5.9
31
31
31
31
31
The analysis column indicates which hemisphere scale was used in the case of LMA (after Wolfe, 1979) and which modern data
set was used in the case of CLAMP analysis. Data from three CLAMP analyses is included to provide some indication of the
variation in estimates and regression uncertainty values when di¡erent CLAMP data sets are used. All three data sets are almost
entirely comprised of Northern Hemisphere £oras. Modern CLAMP data from the Southern Hemisphere is still scarce. The 101site data set is similar to the original published 106-site data set (Wolfe, 1993) but with sub-alpine samples removed (Herman
and Spicer, 1997). Sub-alpine sites are de¢ned as those where the WMM is less than 16‡C and the mean temperature of the coldest month is less than 3‡C. These sites were outliers in the initial version of CLAMP (Wolfe, 1993). The 144-site data set includes
additional data from Japan, and the 170-site data set includes the additional Japanese data and the sub-alpine data (Wolfe, pers.
comm., 1998). The number 31 is the number of leaf character states included in the analysis. Precipitation estimates from
CLAMP have high regression uncertainty attached to them and should be interpreted with caution. *New Zealand palynological
zone PM2 extends from approximately 77 to 65 Ma, and PM3 from 65 to 54 Ma (Raine, 1984).
Although a small CLAMP data set of New Zealand sites does exist, no modern New Zealand
£oras were included in the predictor data sets
used for the analyses presented here. This is because adding New Zealand £oras into predominantly Northern Hemisphere data sets generally
produces estimates with higher uncertainty values.
In addition to this, when analyses were made for
New Zealand fossil sites with New Zealand predictor data sets, the fossil sites did not plot near
the cloud of modern New Zealand samples. This
suggests that there is no physiognomic similarity
between these Late Cretaceous and early Paleocene fossil £oras from Pakawau and the modern
New Zealand £oras in the existing data set, therefore making them inappropriate comparisons
(Kennedy, 1998).
The measure of uncertainty used here for
CLAMP-produced estimates is one standard deviation of the residuals. This is a measure of the
error about the regression line, and is an indication of minimum quanti¢able error only. Errors
such as taphonomic bias are more di⁄cult, if not
impossible, to quantify but must nevertheless also
be kept in mind when making interpretations.
Wilf (1997) calculated binomial sampling error
for temperature estimates from LMA. If this error
is calculated for the three £oras from Pakawau,
the values are as follows : Pakawau Bush
Road = S 2.0‡C, IT = S 2.6‡C, PPT = S 2.3‡C.
These values are signi¢cantly higher than the
0.8‡C of error used by Wing and Greenwood
(1993) for LMA estimates. Sampling error has
not been calculated for CLAMP-produced estimates here and, for consistency, one standard deviation of the residuals has been used throughout.
PALAEO 2871 25-7-02
PALAEO 2871 25-7-02
Each leaf form was assigned a score between 0 and 1 for each of 31 leaf character states. Percentages were then calculated from these scores for each character
state. The percentage score for each character is calculated from the sum of the raw scores (the numbers between 0 and 1 assigned to each character state for each
leaf form) divided by the number of leaf types included. The number of types is the number of types that had scores for each character state. Where a character
state cannot be scored for a leaf form because of lack of preservation, that form is not included in calculation of the percentage score, hence why the number of
forms included for each character state can di¡er from the total number of forms in the £ora. It is often the case that apical information is not preserved as this
tends to be an easily fragmented part of the lamina. For further details of the CLAMP scoring method refer to Wolfe (1993) and later re¢nements of the method
(e.g. Herman and Spicer, 1997).
E.M. Kennedy et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 184 (2002) 321^345
Table 2
CLAMP percentage scores for the Pakawau Bush Road, IT and PPT £oras, Northwest Nelson, New Zealand
339
340
E.M. Kennedy et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 184 (2002) 321^345
5. Latest Cretaceous and early Paleocene climates
of the northwest South Island, New Zealand
Palaeoclimate estimates from the three £oras
are summarised in Table 1. Data from both
LMA and CLAMP are displayed. Note that there
are three sets of CLAMP estimates because three
di¡erent modern (predictor) data sets were used.
The variation in CLAMP-produced estimates
within each leaf £ora shows that palaeoclimate
results from CLAMP are in£uenced by which predictor data set was used to produce the estimates.
It is therefore going to be essential for the con¢dent application of CLAMP to fossil £oras to
develop at least some understanding of how
data set changes a¡ect results and to determine
which predictor data sets are the most appropriate for speci¢c fossil £oras. This problem is still
being addressed and will no doubt require further
experimentation to re¢ne the CLAMP method.
For example, current research is showing that
although changing the predictor data set may
have the same general e¡ect on climate estimates
from fossil £oras, individual fossil £oras may be
a¡ected to di¡erent degrees. An example of this is
the introduction of modern data from New Zealand into a predominantly Northern Hemispherebased predictor data set which tends to result in
lower temperature estimates for the three fossil
£oras, but the estimates from each fossil £ora
are a¡ected by di¡erent amounts.
There is good agreement between all available
sources of quantitative and qualitative data of
relevance to the Pakawau Road locality (Table
3). An earlier LMA produced a MAT estimate
of 13^14‡C (Mildenhall, 1968). This estimate
was based on analysis of 16 leaf types from various localities within the Pakawau Group. Temperature estimates from oxygen isotope studies
using belemnites suggested minimum temperatures of about 14‡C for the coastal waters of
southern New Zealand in the Maastrichtian
(Clayton and Stevens, 1968; Stevens and Clayton,
1971). Palaeoclimate models developed by the Department of Meteorology at the University of
Reading indicated MATs of between 12 and
16‡C for New Zealand at a palaeolatitude of
50^60‡S around 70 Ma (Valdes et al., 1996).
CLAMP estimates suggest warm peak summer
average temperatures and growing season lengths
of 7^8 months (Table 1).
Temperature estimates for the Paleocene £oras
suggest that they grew under slightly cooler conditions than the Cretaceous Pakawau Bush Road
assemblage. In concurrence with the lower MATs,
growing season lengths were possibly slightly
shorter, but once regression uncertainties are considered, the di¡erence between the Late Cretaceous and Paleocene growing season estimates is
negligible. However temperature estimates are
also less consistent from the Paleocene £oras.
LMA of the IT £ora resulted in a MAT estimate
of 7^8‡C, whereas the CLAMP method (101 data
set) estimated V11‡C. Similarly, the PPT £ora
produced MAT estimates of 6^7‡C using LMA
and V9‡C using CLAMP (101 data set). When
a minimum error value (standard deviation of the
residuals here, or for LMA ^ binomial sampling
error or standard error) is taken into consideration however, there is little to separate estimates
Table 3
Palaeoclimate data for the latest Cretaceous Pakawau Bush Road locality from various sources, both quantitative and qualitative
MAT
LMA
CLAMP (101 data set)
CLAMP (144 data set)
LMA (regional Pakawau £ora; Mildenhall, 1968)
Oxygen isotope (minimum sea surface) (Clayton and Stevens, 1968)
Global palaeoclimate model (University of Reading)
14.8 S 0.8‡C
13.8 S 1.8‡C
12.7 S 1.9‡C
13^14‡C
V14‡C
12^16‡C
Precipitation
moderate (MGSP: 971 S 234 mm)
moderate (MGSP: 955 S 485 mm)
The general consistency in the temperature estimates promotes a greater degree of con¢dence in the results from this £ora than
could be expected from looking at any one predictor alone. The rainfall parameter is mean growing season precipitation
(MGSP).
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E.M. Kennedy et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 184 (2002) 321^345
from the two methods for each of the £oras.
WMM were similar to those from the Cretaceous
assemblage. Within each CLAMP analysis there
was a consistent slight WMM cooling from the
oldest to the youngest assemblage, albeit statistically insigni¢cant (Table 1).
CLAMP rainfall estimation is not yet very robust with high regression uncertainty values associated with the estimates. Rainfall estimates for
the Pakawau Bush Road £ora from the three
analyses were consistent and suggest moderately
high levels of precipitation. In the case of this site
there is also supporting evidence from maceral
analysis of coal within the fossil-bearing sequence
for abundant moisture levels (Kennedy, 1993).
The coal has a high tissue preservation index
which indicates a high water table (Newman,
1989). Rainfall estimates from the Paleocene £o-
341
ras were strongly a¡ected by the choice of
CLAMP predictor data set combination used in
the analysis, and we believe that little con¢dence
can be placed in the precision of rainfall estimates
from these two £oras at this stage. Of the three
analyses however, we suggest that more con¢dence can be placed in the rainfall estimates
from the 144 and 170 analyses than in those
from the 101 analysis because IT and PPT plot
in closer proximity to the modern cloud of sites in
the former two analyses. Estimates from the 144
and 170 analyses suggest high growing season
rainfall levels.
It was noted that in three-dimensional plots of
the three axes accounting for the greatest variation in the analysis, the PPT and IT localities
tended to plot in relatively close proximity to
each other, but away from the localities in the
Fig. 9. Plots showing the relative positions of the three New Zealand fossil localities after CLAMP analysis with modern (predictor) data sets. The small points are the modern £oras. The fossil sites are the labelled, larger, points. Plots from two analyses are
shown ^ one using 101 modern £oras (a, b) and the other using 144 modern £oras (c, d). The 101-site modern data set is similar
to the data set used in the published CLAMP manuscript (Wolfe, 1993). The biggest di¡erence between the 101- and 144-site
data sets is the addition of 36 £oras from Japan. This proved to be of signi¢cance to the plot positions of the New Zealand fossil £oras. When this large group of £oras from Japan is present in an analysis, the New Zealand fossil £oras tend to plot near,
or within, the cloud of sites from Japan. When the data from Japan is absent, however, the New Zealand sites plot further away
from the cloud of points representing the modern data. Compare the axis 2/axis 3 view of the 101-site analysis (b), with that of
the 144-site analysis (d). Scale values are relative coordinates in the regression space.
PALAEO 2871 25-7-02
342
E.M. Kennedy et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 184 (2002) 321^345
modern predictor data set (e.g. Fig. 9b). This suggests that they have a physiognomic signal that is
signi¢cantly di¡erent to that of the £oras in the
predictor data sets included here. The resulting
estimates are therefore less well constrained than
those from a fossil £ora that plots closer to the
cloud of predictor samples, such as the Pakawau
Bush Road £ora (Fig. 9). Although IT and PPT
plot away from the modern cloud of sites along
axis 3 in the 101 site analysis in particular, data
from this analysis was still included for comparison as this dataset is the one that is most similar
to the original published CLAMP dataset (Wolfe,
1993). The 144 analysis is an exception. In Figs.
9c,d, the IT and PPT fossil localities plot closer to
the modern vegetation sites than in any of the
other analyses made (most of which are not ¢gured here, but refer to Kennedy, 1998). The main
di¡erence between the 144 and 101 data sets is the
substantial addition of modern £oras from Japan
(Wolfe, 1997; Wolfe, pers. comm., 1998; Spicer
and Wolfe, 2001, CLAMP web page), and it is
these £oras that the New Zealand fossil sites
plot closest to. It is clear that the foliar physiognomic signatures of the IT and PPT fossil £oras
are more similar to those of the modern £oras
from Japan than those of the North American
£oras for example. The reason for this a⁄nity is
unknown but it emphasises the importance of collecting as wide a variety of modern analogue assemblages as possible for inclusion in CLAMP
data sets. The more representative CLAMP data
sets are of the physiognomy of the fossil £oras
being analysed, the stronger the climate estimates
are likely to be.
However, when IT and PPT were analysed with
a 38-site data set which only included £oras from
Japan, they again plotted in isolation. With £oras
such as IT and PPT, methods like the Nearest
Neighbour approach (Stranks, 1996; Stranks
and England, 1997) would be di⁄cult to apply
and the results obtained would need to be considered with caution. The Nearest Neighbour approach is based on the CLAMP data set. However, instead of estimating climate variables from
vectors based on the whole data set, this approach
uses a di¡erent method of calibration. The Nearest Neighbour method only takes climate varia-
bles from modern sites that plot in close proximity in physiognomic space to the fossil £ora under
consideration. It follows that if the fossil site plots
at some distance from its nearest modern neighbours, calibration is degraded.
The leaf forms in both the IT and PPT £oras
have particularly high length-to-width ratios
(L :W). The combined percentage of the L:W =
3^4:1 and L:W s 4:1 categories for IT is 77%,
and for PPT is 73%. This feature, in combination
with an over-representation of non-entire margined forms, is particularly characteristic of
stream-side plants (Wolfe, 1971). It has been argued that the over-representation of leaves from
stream-side plants in a fossil assemblage could
bias any palaeoclimate interpretation made from
that assemblage, because of the unusually high
percentage of toothed (i.e. non-entire margined)
forms (e.g. Wolfe, 1971, 1993). However, Wolfe
(1993) found from his studies of the CLAMP data
set that although over-representation of streamside plants could result in erroneous precipitation
estimates, there was little e¡ect on estimates of
temperature parameters.
The temperature estimates from the three £oras
therefore suggest that there was a slight cooling
between the Late Cretaceous sample and the Paleocene samples. Although temperature estimates
from LMA and CLAMP for the Paleocene £oras
are not in exact agreement, and the use of di¡erent CLAMP predictor data sets inevitably produces slightly di¡erent values, they nevertheless consistently suggest this minor cooling. There is
corroborating evidence from the Antarctic plant
record of a cooling from the latest Cretaceous
into the Paleocene (Truswell, 1990; Askin, 1992).
Askin (1992) reviewed both qualitative and quantitative information based on a variety of studies
covering research on wood, £oral composition
(NLR methods), foliar physiognomy, cuticle analysis and palynology. She concluded that Late Cretaceous to early Tertiary climates in the Antarctic
Peninsula region varied between cool and warm
temperate with high rainfall. The Antarctic Peninsula region was at palaeolatitudes of 59^62‡S in
the late Maastrichtian and early Tertiary (Francis,
1986), similar to the palaeolatitudes estimated for
the northwest Nelson region at that time.
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E.M. Kennedy et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 184 (2002) 321^345
6. Conclusions
The Pakawau Bush Road data suggests that
late Campanian or Maastrichtian lowland £oodplain vegetation in what is now the northwesternmost part of the South Island was dominated by
diverse angiosperms with minor components of
ferns and gymnosperms (including podocarps
and araucarians). The depositional environment
for this assemblage £uctuated, allowing intermittent peat deposition and burial which resulted in
thin coal seams interspersed with £uvial and lacustrine sediments depending on lake level £uctuations and the proximity of £uvial channels. Consistency in results from di¡erent methods ensures
a relatively strong palaeoclimate interpretation.
The local climate was temperate, with MATs of
between 12 and 15‡C, and there was signi¢cant
rainfall. We suggest that this local palaeoclimate
interpretation can be applied with reasonable con¢dence to a more regional setting, because of the
general agreement between estimates from the local leaf £ora, and methods such as oxygen isotope
analysis, and from a more regional (albeit limited
in the number of leaf forms) leaf morphology
study (Mildenhall, 1968).
The two Paleocene local £oras suggest that
early Paleocene angiosperm diversity was lower
than in the latest Cretaceous. However, it must
be noted that the relatively poor outcrop quality
of the Paleocene £oras as compared with that of
the Cretaceous locality did not allow for equally
large collections to be made. This may have disadvantaged the collection diversity of the Paleocene £oras. In addition, the restricted outcrop information meant that an equivalently detailed
interpretation of depositional setting was not possible, although this did not a¡ect the extraction of
palaeoclimate data.
There is perhaps less precision with the palaeoclimate interpretation from these two local £oras
because of the greater di¡erences in estimates
from the two leaf physiognomic methods and
the lack of information on New Zealand terrestrial early Paleocene climate from other methods
to compare with the leaf results. However, bearing this in mind, the estimates do suggest that the
IT £ora (the older of the two Paleocene £oras)
343
experienced slightly cooler temperatures than the
Cretaceous £ora, and that the PPT £ora grew in
still cooler conditions. Evidence from the physiognomic study of these Late Cretaceous and Paleocene leaves from New Zealand therefore supports
inferences made by other means that there was a
slight cooling from the latest Cretaceous into the
Paleocene at southern mid- to high latitudes.
The potential of New Zealand fossil leaf £oras
for palaeoclimate analysis is great. What is needed
is a broader study of coeval New Zealand fossil
£oras to provide a greater degree of con¢dence in
palaeoclimate estimates from individual localities
(particularly precipitation estimates, which have
relatively high levels of uncertainty). These New
Zealand data also need to be compared further on
a global scale, and this should be combined with
continued research into understanding the applications and limitations of the modern data sets
used in these analyses.
Acknowledgements
The valuable assistance and advice given by
John Lovis, Ian Daniel, Jane Newman, Ian Raine,
Judith Totman Parrish and Kirk Johnson is gratefully acknowledged. Ian Raine and Malcolm
Warnes provided palynological data and age determination. The manuscript was improved by
comments from Mike Pole and an anonymous
reviewer. This work was funded by an Open University, United Kingdom, PhD studentship and
the University of Canterbury, New Zealand. A
New Zealand Foundation for Research, Science
and Technology Post Doctoral Fellowship provided time for manuscript preparation.
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