THE LATE JURASSICAMMONITE FAUNA OF NEW ZEALAND_2 pptx

PART
6.
PALAEOECOLOGY
FAUNAL SPECTRA
The post-mortem drift of empty shells as observed in mod-
em
Nautilus
(e.g. Kennedy
&
Cobban 1976; Saunders
&
Landman 1987; Ward 1987, 1988) has, at least theoreti-
cally, the potential of distorting the relative proportions of
ammonite taxa at specific stratigraphic horizons. How-
ever, it is currently thought that in practice the effect of
post-mortem drift is minimal and that therefore the
faunal
spectra derived from study of ammonite assemblages can
provide a reasonably representative sample of ammonite
populations and habitats through space and time
(Westermann 1990,
1996b; Cecca 1992).
A number of authors (e.g. Ziegler 1967; Kennedy
&
Cohban 1976; Westermann 1990, 1996b; Cecca 1992;
Femandez-Lopez
&
Melendez 1996) have drawn atten-
tion to the ecological differences between the two major
groups of ammonites: the streamlined, smooth-shelled
Leiostraca of the early ammonite

literature (including no-
tably the phylloceratids and lytoceratids) and the strongly
ornamented Trachyostraca. Modem opinion envisages the
Lejostraca
as
having been pelagic types, capable of living
offshore and also of populating deep-shelf habitats
(250-
300 m depth; Westermann 1990, 1996b) (see also Cecca
et
al 1990; Mouterde
&
Elmi 1991). However, as noted
by Kennedy
&
Cobban (1976, p. 34) the local abundance
of Leiostraca in clearly shallow-water deposits indicates
that they were not entirely confined to deep-water situa-
tions. By contrast, the Trachyostraca are generally con-
sidered to have inhabited neritic shallow-water environ-
ments (ca. 30-100 m depth,
Westermam 1992).
Following Ziegler
(1967), the Late Jurassic ammonite as-
semblages of New Zealand can be represented by means
of spectra, illustrating the proportions of Leiostraca and
Trachyostraca (Fig. 58). Ziegler (1967, fig. 12) proposed
that the dominance of Leiostraca (in relation to
Trachyostraca) at a specific stratigraphic level was indica-
tive of deep water conditions (ca. 300-400 m depth). How-

ever, as cautioned by Kennedy
&
Cobhan (1976 p. 34)
"off-shore" rather than
"deep-water9'perse may he a more
appropriate appellation. Westermann
(1990, 1996b) and
Cecca
(1 992) regard Leiostraca as pelagic, with the capa-
bility of occupying deep water hahitats, in contrast to the

-
.
neriiic shallow water Trachyostraca. Fernandez-Lopez
&
Melendez (1 996) have correlated increases in abundance
of Leiostraca with changes accompanying eustatic rises in
sea level.
As may be seen from Fig. 58, in
the New Zealand Late
Jurassic sequence percentages of Leiostraca reach
55%,
56% and 50% in the Oraka Sandstone, Ohinerurn Forma-
tion and Waibarakeke Conglomerate, respectively. These
percentages may be interpreted as either indicating the
presence of offshore (pelagic) influences or local post-
mortem drift of otherwise pelagic ammonites (Tanabe
1979, p. 626; Cecca 1992, p. 265; Batt 1989, 1993).
Sedimentological studies by Meesook (1989) indicate a
generally deep water

(=
basin plain) origin for the Oraka
and Ohineruru, and that of a deep water fan for the
Waiharakeke. Thus the offshore explanation has perhaps
greater validity.
The remaining stratigraphic units, with percentages of
Leiostraca ranging from 30% (Kowhai Point Siltstone) to
5%
(Kinohaku Siltstone), are all clearly a reflection of shelf
or upper slope habitats, as also indicated by
Meesook's
(1989) sedimentological analyses. The Takatahi Forma-
tion, which lacks Leiostraca, is a major component of one
of Meesook's coarse large volume fan cycles that
prograded into the Kawhia basin during the Late Jurassic
(see also Meesook
&
Grant-Mackie 1995).
The changes in ammonite
faunal spectra are thus a reflec-
tion of a complex of changes: (i) changes in pelagic (off-
shore) influence; (ii) sedimentological changes relating to
the prograding development of fan systems; (iii) eustatic
changes in sea level. Similar fluctuations in the propor-
tions of Leiostraca and Trachyostraca, and presumably
reflecting a similar interplay of environmental factors, have
been documented by Sarti (1986) from Northern Italy,
Wendt (1963) and Christ (1960) from West Sicily, Vigh
(1 971) from Hungary, and Ceccaet al. (1990,1993), Cecca
(1992) and Fernandez-Lopez

&
Melendez (1 996) from the
Western Tethys (Fig. 59).
LITHOLOGICAL CHANGES ACCOMPANYING
LATE
JURASSIC EUSTASY
A lack of agreement on the precise overseas correlations
to be applied to the Late Jurassic sequence
o6New Zea-
land has been a significant impediment to the application
and testing of the recently proposed models of eustatic
cycles (e.g. Haq et al. 1988). Although a study of eustatic
influences was made by Stevens
(1 990a), this was done in
the absence of the refined correlations based on the am-
monite faunas
ofthis Bulletin. Nonetheless, Stevens noted
changes in
faunal diversity, first and last appearances,
turnover and influxes of Tethyan taxa that were probably
related to global eustatic changes.
In Fig. 60 the eustatic models of Haq et al. (1988; with
modifications by Strohmenger et al. 1991) and
Hallam
(1988, 1992) are matched with the standard lithological
column of the Late Jurassic (using the Kawhia and Port
Waikato sequences as the standards), and employing as a
means of calibration the age assessments derived from the
ammonite faunas,
as

documented in this Bulletin. As may
be seen from the diagram, there is some measure of agree-
ment between lithological expression and the postulated
eustatic highs and lows. Thus development of
coarse-
grained sequences (conglomerates, sandstones etc) tends
to correspond with falls
in sea level, when sea water is
SCALE
(m)
Y1
U
LITHOSTRAT. OSEAS
"?
COLU~~
CORREL.
p
UPPER
TITH,.

a
2
PUT1
SILTSTONE
W.
'
2
CONGLOM.
g
a

KINOHAKU
U
SILTSTONE
0
3
3
<
-
-
-
-
-
-
-
3
I
2
4
U
l'
2
t-
TAKATAHI
C
0
l
;
5
L
2

KOWHAI
$3
POINT
SILTSTONE
0
-
u
-
WAIKUTAKUTA
SILTST.
3
KIWI
SANOST.
OHINERURU
5
5
FORMATION
3
5
3
ORAKA
S,,,,,\
z
?
>
9
RENGARENGA
2
o
3

GROUP
g
2
Z
(part)
C
X
U
0
LEIOSTRACA
~
-
1
TRACHYOSTRACA
Percentage of total ammonite fauna
Figure
58
Faunal specha for the Late Jurassic ammonite faunas of New Zealand. The lithostratigraphic column is based on
Fleming
&
Kear (1960) and the overseas correlations are those adopted in the current study (Fig.
54).
The eustatic curve is
based on
tlaq et al. (1988). as modified by Strohmenger et al. (1991). The histograms illustrate the proportions expressed
as
a
percentage
of
Leiostraca and Trachyostraca in the ammonite faunas oftheNew Zealand Late Jurassic.

The
occurrences of giant
ammonites
in
the New Zealand sequence are also noted to the far right of the diagram (see p. 107 and Fig.
62).
shallower over the continental shelves. Conversely, the
example by Strohmenger et al. (1991) and
Li
&
Grant-
development of fine-grained sequences generally coincides
Mackie
(1
993), is reflected by development ofthe Takatahi
with periods of rises
in
sea level, when sea water deep-
Conglomerate. Another low in sea level
in
the later half
ened over the continental shelves.
ofthe Early Kimmeridgian (Marques et al. 1991) is prob-
ably reflected by development ofthe Kiwi Sandstone. The
The marked fall in sea level at the
boundary between Late
extensive siltstone sequences
ofthe Puti Siltstone and the
Kimmeridgian and Early Tithonian, as documented for
Kinohaku

Siltstone
appear
to
have
developed
during
Figure
59
Faunal spectra similar to those plotted in Fig.
58
have been published by authors such as Ziegler
(1967).
Recent
examples have been provided by Ceccaet al. (1990.
1993),
and the abovediagram
is
derived from these authors. The histograms
show the percentage proportions of Leiostraca and Trachyostraca for the Lower and Middle Tithonian of the Umbria-Marche
region of the Apennines of Northern Italy. A eustatic curve derived from
Haq
et al. (1988) and Strohmenger et al. (1991)
has
been plotted to the lefl of the figure. (Ph
=
Phylloceratina:
L
=
Lytoceratina).
2

z
0,
W
times of high sea level, which were presumably accompa-
nied by optimal conditions for the deposition of
fme-
grained sediments. However, a major exception is the
Waiharakeke Conglomerate, which appears to have been
deposited at a time when sea levels were probably at their
highest during the Jurassic (see Haq et al. 1988). There-
fore, although lithology to some degree matches the world-
wide sequences of eustatic fluctuations during the
Kimmerideian and Tithonian, the presence of mismatches,
Urnbria-Marche Apennines
(Italy)
by Strohmenger et al. (1991); Marques et al. (1991); Li
&
Grant-Mackie (1993) and Norris
&
Hallam (1995). None-
theless, considering that the New Zealand region in the
Late Jurassic was sited close to an active plate margin
(Sporli 1987; Sporli
&
Ballance 1989; Wilson et al. 1989),
and was being affected by movements associated with the
Rangitata Orogeny, the quality of the overall match with
the world-wide eustatic fluctuations is reasonably good.
-
such as that ofthe Waiharakeke conglomerate, is perhaps

The stratigraphic imprint of the eustatic fluctuations
ap-
evidence for contem~oraneous local tectonic activity
pears to continue into the lower part
ofthe Upper Tithonian,
V)
z
if the Coleman Conglomerate is interpreted as the record
Local tectonic activity, related to the early phases of the
of
a
marked regression, spanning the top of the
ponfi
zone
Rangitata Orogeny (Suggate
1978), was probably respon-
and the bottom of the
scruposus
zone (Haq et al. 1988;
sible for local shallowing and the deposition of
coarse-
grained sediment. The effects of similar local tectonism
Strohmenger et al. 1991).
Z
5
Z
0
I
k
L

-
on the Late Jurassic eustatic record have been documented The overlying formation, the Waikorea Siltstone, repre-
z
8
=
E
F:-
3%
2:
32
0
::
>
V]
z
K5
$6
01
-'k
2
2
;:
+-
LEIOSTRACA TRACHYOSTRACA
I
I
I
I I
l
INTER-

NATIONAL
STAGE
Z
5
Z
0
E
F
K
W
P
P
3
z
5
Z
0
I
2
W
-I
0
E
7
LOWEF
TITH.
z
5
X
K

W
I
z
X
K
W
P
P
a
Z
5
::
z
W
I
3
a
r
S
Figure
60
(opposite)
In this figure the eustatic curves of Haq et al. 1988 (modified by Strohmenger et al. 1991 and Hallam 1988)
have been fined to the New Zealand Late Jurassic
stratigraphic column (see Fig. 53). using the correlations derived from the
current ammonite study. The
litholo~ for the Kawhia column is from Fleming
&
Kear
(1

960). Kear
&
Fleming (1976), with
amendments by Meesook (1989) (see also Ballance
&
Campbell 1993, fig.
3).
The lithology for the Port Waikato column is
from Purser (1961) and Waterhouse (1978)). The dominant lithologies that
are
coarser than clay and silt (i.e. fine, medium,
coarse and very coarse sand and gravel) have been highlighted by the dark tone pattern.
sents a partial recovery of sea level. However, as indi-
cated by Ballance
(l988), much ofthe Waikorea Siltstone
appears to represent a marginally
marine environment,
unfavourable to most forms of marine life.
A
near-shore
environment is indicated, with exposure to frequent surges
of coarse-grained sediment flows. Although sparse
Buchio
and belemnites are present, no ammonites have been col-
lected to date and they appear to be genuinely absent.
The introduction
ofthe massive non-marine braidplain fan
deposits of the Huriwai Group (Rodgers
&
Grant-Mackie

1978; Waterhouse 1978; Ballance et al. 1980; Ballance
1988) terminated the Jurassic marine record in New Zea-
land and heralded the commencement of the main phase
of the Rangitata Orogeny (Suggate 1978) that continued
into the Early Cretaceous (Late Aptian) and was responsi-
ble for the development of an extensive ancestral land-
mass in
theNew ZealandiNew Caledonian region (Stevens
1989; Stevens
&
Fleming 1978).
RELATIONSHIF' OF AMMONITE BIOTA TO
LATE JURASSIC EUSTASY
As noted by Stevens
(l990a), biological changes that are
likely to accompany rising sea levels include: high diver-
sity, increases in speciation rates, influxes of off-shelf and
open oceanic faunas, and influxes of taxa with wide-rang-
ing geographic distribution (e.g. Cosmopolitan, Tethyan
etc.). On the other hand, falling sea levels are accompa-
nied by low diversity, extinctions, biostratigraphic gaps,
absence or severe restriction of shelf facies, and increases
in endemism. Such biological changes can be identified
in
theNew Zealand Jurassic fossil record (Stevens 1990a).
Hantzpergue (1995) and Hoedemaeker (1995) have made
similar studies in the European Kimmeridgian and lowest
Cretaceous, respectively. Although ammonite assemblages
form relatively minor constituents of the total fossil fauna
of the New Zealand Late Jurassic, they do nonetheless ex-

hibit the main features of the biological changes that are
probably related to sea level fluctuations.
The eustatic fluctuations
ofthe Kimmeridgian are reflected
by the tumover and diversity values for the ammonite as-
semblages of the Upper Heterian and Lower Ohauan.
Thus, the tumover and diversity values for the ammonite
assemblages
ofthe Ohiiernrn Formation and Waikutakuta
Siltstone can be interpreted as a probable response to the
rising sea levels of the time. Similarly, the influx of
Leiostraca in the Ohinerurn Formation (reaching 56.2%
ofthe total ammonite assemblage) is probably a reflection
of an influx of
offshore/oceanic taxa, moved towards the
shore by the rising sea level (Zeiss 1994; Geyssant 1994;
Hantzpergue 1995).
As already documented
by Stevens (1990a) in relation to
the total Jurassic fossil fauna, the rise in sea level to reach
a major peak in the Middle Tithonian is particularly well
marked in biotic
patterns. The ammonite assemblages of
the Kinohaku Siltstone show the highest values for
tumo-
ver and diversity, which as also has been demonstrated by
Hoedemaeker
(1995), may be interpreted as the fauna1
response to a major rise in sea level. However, unlike the
assemblages in the

Ohine~~ Formation, Trachyostraca
are dominant (95.03%) in the Kinohaku Siltstone and
Leiostraca constitute only 4.96% of the total ammonite
assemblage.
The continuing high stand of sea level in the Middle
Tithonian is also reflected in the ammonite assemblages
of the lower and middle parts of the Puti Siltstone. Values
for turnover, diversity and abundance are comparable with
those for the Kinohaku. After those for the Kinohaku,
these values are the second highest for the New Zealand
Late Jurassic. Also, like the assemblages of the Kinohaku,
Trachyostraca dominate in those
ofthe Puti, and Leiostraca
constitute only 10.82%
ofthe total.
As may be seen from Fig.
61, there is a substantial fall-off
in
turnover, diversity and abundance
in
the upper part of
the Puti Siltstone (i.e. in horizons above those at Puti Point
and Motutara Peninsula in the Kawhia Harbour sequence)
and this is undoubtedly a response to the falling sea level
of the time. The same trend is even more apparent
in
the
overlying
Coleman Conglomerate, which according to the
eustaticmodels

ofboth Haq et al. (1988) and Hallam (1988,
1992) (see also Strohmenger et al. 1991, fig. 8) coincides
with a marked low in sea level.
The poorly
fossiliferous Waikorea Siltstone represents a
minor recovery of sea level that was perhaps insufficient
to fully restore open-sea oceanic conditions (as noted by
Ballance 1988).
GIANT AMMONITES AND EUSTASY
The coincidence of occurrences of giant ammonites with
times of rising sea levels has been discussed by Stevens
(1988). A classic situation is that of the
Santoniani
Campanian of Westphalia involving the occurrences of
giant specimens of
Pnrnp~rzosia
and
Pachydiscus,
includ-
ing the largest ammonite known to science (Stevens 1988,
pp.
144-145). These occurrences are clearly related to the
shoreline of the
SantonianICampanian marine transgres-
sion and it is likely that the conditions associated with the
rising sea level (probably resulting in the generation of
regions of upwelling
and general tumover of the water
column) have served to lift on to inshore shelf areas repre-
sentative samples of populations that were normally resi-

dent in deeper waters some distance offshore. Situations
that are able to he interpreted as being comparable to those
in Westphalia can also be seen in the Kawhia Harbour se-
quence.
Figure
61
(opposite)
In this diagram the lithostratigraphical column and eustatic curve of Fig.
60
have been used as the basis for
plotting the following characteristics of the New Zealand Late Jurassic ammonite fauna:
(i)
number of taxa making a first
appearance;
(ii)
number of taxa making a last appearance; (iii) an expression of the extent of faunal turnover, obtained by
averaging the number of
first and last appearances, dividing by the total number oftaxa, and expressing the result as apercentage;
(iv) an expression of fauual diversity, as indicated by the number of taxa per stratigraphic interval; (v) abundance, expressed as
total number of specimens (e.g.
n
=
153), but also with separate histograms for Leiostraca and Trachyostraca. For a key to the
litholoeical
svmbols that have been used. refer to Fie.
53.
The occwences of giant specimens of Lytoceras in the
hybonotum zone) are probably absent from the New Zea-
Ohineruru Formation and Waikutakuta Siltstone, as

docu-
land record because the Early Tithonian is either not rep-
mented by Stevens
(1985a), appear to coincide with peri-
resented at all, or highly condensed.
~
~
ods of rising sea level at the top of the plaiynota zone
(Ohineruru) and at the top of the divisum zone
(Waikutakuta). The well-known occurrences of giant am-
monites at Puti Point and Motutara Peninsula, on the north
shore of Kawhia Harbour, appear to coincide with the pe-
riod of rising sea level that culminated in the major Mid-
dle Tithonian high stand in the
faNauxi and ponti zones
(Fig. 62).
A Late Kimmeridgian period of marine transgression and
high stand of sea level (beckeri and
eudoxus zones; see
Wignall
&
Ruffell 1990; Geyssant 1994; Zeiss 1994;
Hantzpergue
1995), although probably represented in the
Kawhia sequence by sandy intervals and conglomerates
in the Kowhai Point Siltstone and Takatahi Formation, does
not appear to be represented by influxes of offshore taxa,
comparable to those in the Early Kimmeridgian and Mid-
As may be seen
from Figs 61,62, and Strohrnenger et al.

dle
~ithonian. A possible explanation may lie with the
(1991, fig.
81, other episodes of rising sea level and high
fact that at this time New Zealand was very close to a
stands are recorded on the eustatic curve. However, these
tectonically active plate margin (Wilson et al. 1989, fig.
episodes do not appear to have a
faunal signature in the
6), and as a consequence, any eustatic effects may have
New Zealand sequence. The
high stands of the Early
been neutralised by tectonic activity.
Tithonian (base of the
damini zone and top of the
INTERNATIONAL TETHYAN
1
AMMONITEZONE
I
-
138
1
,
1
1 1
LOWER TlTHONlAN
HYBONOTUM
-
140
BECKERI

-
136-
EUSTATIC CURVE
z
a
g
4:
3
a
Z
5
U
E
r
GIANT AMMONITES
IN
NZ
Figure
h2
Giant ammonites occur at three stratigraphic levels in theNew Zealand Late Jurassic sequence. The relationship ofthese
occurrences to the New Zealand lithostratigraphic framework is shown in Fig. 58.
This diagram summarises the relationship of occurrences of giant ammonites in New Zealand to the Tethyan ammonite
zones and the international stages. based on the correlations derived from the present study (Fig. 54). The eustatic curve is based
on Haq et al. (1988) and Strohmenger et al. (1991).
109
UPPER TlTHONlAN
MIDDLE TlTHONlAN
UPPER
KlMMERlDGlAN
LOWER

KlMMERlDGlAN
DURANGITES
spp.
MICROCANTHUM
PONTl
FALLAUXI
SEMIFORME
EUDOXUS
ACANTHICUM
DIVISUM
HYPSELOCYCLUM
PLAWNOTA
CHAPTER
7.
PALAEOBIOGEOGRAPHY
AND
FAUNAL AFFIMTIES
TETHYAN
FAUNAL
LINKS
In the Southern Hemis~here. the coasts of Gondwana were
The Late
Mesozoic, and the MiddleLate Jurassic in par-
ticular were times when major differentiation became ap-
parent in the marine assemblages of the Northern Hemi-
sphere. For much of the time, a low-diversity northern
Boreal fauna can be clearly separated from a highly di-
verse southern Tethyan fauna (e.g. Stevens 1971,
1980a.
1989, 1990a,b,c, 199

1;
Stevens
&
Fleming 1978; Crame
1986, 1987). Such a
faunal configuration has been widely
viewed as a result of temperature zonation: with the Boreal
Realm representing cool or cold temperate assemblages
and the Tethyan representing
tropical/subtropical/warm
temperate assemblages (Stevens 1980a). However, as
Hallam (1969, 1971, 1975, 1977, 1984) has noted, other
environmental factors (e.g. salinity and eustatic variations)
may also have served to enhance the sharpness of the
BorealITethyan differentiation.
largely populated by wide-ranging assemblages of Tethyan
origin (Cecca et al. 1993; Fourcade et al. 1993). Although
some provincial groupings are apparent (e.g. Ethiopian,
Himalayan, Indo-Pacific, etc.), such provinces appear to
be best viewed as regional derivatives
ofthe wide-ranging
Tethyan Realm (Fig. 63).
Although at times the Northern Hemisphere's Boreal
Realm was counterbalanced by an "Austral Realm" in the
Southern Hemisphere, such a development only occurred
during those time periods when component southern
Gondwana continents were grouped around the South Pole,
such as in the Permian and Cretaceous (e.g. Stevens
1980~1,
1989, 1990c, 1991). At other times, notably in the Late

Jurassic, an Austral Realm cannot be recognised
-
pre-
sumably because the southern Gondwana continents were
sited some distance away from the South Pole. This
ab-
Figure
63
The Tethyan Realm, which includes the Late Jurassic ammonites of New Zealand, can be divided into various sub-
realms. Although other examples of nomenclature are available (e.g. Enay 1972b Challinor 1991; Hull 1995; Ross
et
al. 1992;
-
L
.
.
region occupied a nodal point between the East Pacific subrealm and the Indo-S.W. Pacific subrealm and at various times shared
components
with
both.
As shown
by
Stevens (1990) it is likely that faunal elements could disperse using routes via the Antarctic Peninsula and
South America or in the opposite direction geographically, via Indonesia and the Himalaya The
paleolatimde lines for Australasia
have been derived from
Howarth (l98I), Owen (1983b), Stevens (1985). Lawver
&
Scotese (1987), and Grunow et al. (1987).
Other reconstructions (e.g. Smith et al. 1981, 1994; Ross et al. 1992) place the New Zealand sector much closer to the South

Pole
(i.e. 80-85"s Lat.). The placement of the rift zones is after Fourcade et al. (1993); Cecca et al. (1993).
sence of land in the South Pole region probably encour-
aged the free interchange of
seawater between low and
high latitudes and resulted in the development of an equa-
ble marine climate in the
Southern Hemisphere.
It is likely that a
warm western boundary current, part of a
huge Tethyan-Pacific
,ve, flowed southwards from the
eastern Tethys to extend around the southern margins of
Gondwana (e.g. Stevens
198Oa). This oceanographic situ-
ation led to the development of wide-ranging circum-
Gondwana marine
populations of Tethyan origin.
Like contemporary bivalves and
belemnites (e.g. Crame
1986, 1987; Doyle 1987; Challinor
1991), the ammonite
assemblages of the Late Jurassic of New Zealand show
strong Tethyan affinities. However,
in
considering these
affinities in detail, it is important to make a clear distinc-
tion between the classical groupings of Leiostraca and
Trachyostraca (e.g. Kennedy and
Cobban 1976, p. 34).

The generally smooth-shelled and streamlined Leiostraca,
including the phylloceratids and lytoceratids, are gener-
ally accepted as being able to inhabit offshore or oceanic
environments, although their local abundance in what are
clearly shallow-water deposits indicates that they were not
entirely
confmed to deeper-water situations (Hallam 1969:
Kennedy and
Cobban 1976; Joly 1976; Westerrnann 1990;
Cecca
1992). On the other hand, the often coarsely sculp-
tured Trachyostraca are generally viewed as inhabiting
shallow inshore shelfenvironments
(Westemann 1990 Fig.
67).
As may be expected from such differences in habitat pref-
erences, the Leiostraca of the New Zealand Late Jurassic
show affinities that extend
inore widely geographically than
those
ofthe Trachyostraca. The affinities ofthe New Zea-
land Leiostraca, comprising taxa of
Phylloceras,
Partischeras,
Calliphylloceras, Holcoplzylloceras.
P~choph.vlloceras
and
Lytoceras
are essentially circum-
Gondwana, but with some northwards extensions into

Mexico. southern Europe, the Middle East and Asia Mi-
nor (i.e.
along the Tethyan seaway, cf. Cecca et al. 1993).
On the other hand, while still retaining a strong circum-
Gondwana flavour, the New Zealand Trachyostraca show
a more restricted set of affinities, apparently without any
direct links to southern Europe, the Middle East and Asia
Minor. However, linkages to Mexico remain very strong.
These differences may be interpreted to show that while
the Trachyostraca were largely dependent on migration
around reasonably continuous areas of continental shelf,
and were generally not able to undertake extensive
trans-
oceanic journeys, the Leiostraca, by contrast, were capa-
ble of crossing substantial areas of deep ocean.
In general, the circum-Gondwana links that are shown by
both Leiostraca and Trachyostraca of the New Zealand
Late Jurassic are a reflection of the various
fauna1 migra-
tions that occurred along shorelines and continental shelves
extending around the margin of Gondwana (Fig. 64). Link-
ages extended from the Himalayas and Malagasy, through
Indonesia,
PapuaNew Guinea and New Caledonia toNew
Zealand. Comparable linkages then continued on from
New Zealand through Western Antarctica to Chile and Ar-
gentina and northwards to Mexico and the southern USA.
Migrations occurred in various directions, depending on
the oceanic current systems, facies conditions and climatic
constraints of the time (e.g. Stevens

1980a).
As shown in Fig.
64
the Circum-Gondwana Links can be
achieved by using a conventional Gondwana assembly (e.g.
Lawver
&
Scotese 1987; De Wit et al. 1988; Grunow
Dalziel
&
Kent 1987; Grunow, Kent
&
Dalziel 1987;
Wilsnn et al. 1989; Gradstein et al. 1992). However. by
assuming an expanding earth hypothesis, other authors (e.g.
Owen 1976.
1983a,b; Shields 1979, 1983; Carey 1992)
have achieved an even tighter fit of the circum-Pacific
countries (Fig. 65). These alternative assemblies have,
for example, made direct linkages between New Zealand
and Central
&
South America (cf. Khudoley 1970, 1974;
Khudoley
&
Rzhonsnitskaya 1979) even more of aphysi-
cal possibility.
TRANS-PACIFIC FAUNAL LINKS: THE
MEXICAN CONNECTION
As discussed in

the previous section and as shown in Fig.
64, the New Zealand Late Jurassic ammonite fauna has
many close relationships that extend along the
Indo-Pa-
cific margin ofthe Tethys: i.e. Australia. New Caledonia,
New Guinea, Indonesia, India, Pakistan and Malagasy.
Other equally close relationships extend in the opposite
direction geographically
-
to West Antarctica and South
America.
All these relationships can be categorised as resulting from
population movements in the seaways around the periph-
ery of Gondwana (Perigondwanian distribution of Enay
1972b) (Fig. 64). However, a number of distinctive New
Zealand taxa appear to have very marked affinities with
Mexico, seemingly without linkages via West Antarctica
or South America. These taxa include
Epicephalites
maiwicki, Subneumayria
cf.
ordonezi, Idoceras heteriense,
I
speighti, Kossrnatia macnazrghti, K. pilicosfa,
K.
cf.
santarosano
and
Kawhiasphincfes antipodus.
It is ac-

cepted that collection failure may be responsible for the
absence of these taxa or close relatives from the interven-
ing areas around the Southeast Pacific rim.
Unconfomities
may introduce gaps in the record and appropriate facies
may be lacking at the specific time horizons.
For example, although there was probably some degree of
geographical continuity between Mexico and the Andean
region in
theLate Jurassic (e.g. Anderson
&
Schmidt 1983),
there is at the same time evidence of the widespread oc-
currence of hypersaline conditions that would be inimical
to
normal marine life (e.g. Legarreta
&
Uliana 1996).
Therefore, in practice, major environmental factors may
have placed restrictions, at various times. on the move-
ment of marine taxa
bktween Mexico and the Andes.
If it is
assumed that the linkage between New Zealand and
Mexico is valid and is ofprimary origin, then a number of
possibilities arise:
(i) Arkell (1953a p. 263; 1956 p. 599) and Khudoley
(1970, 1974; Khudoley
&
Rzhonsnitskaya 1979) in-

Figure
64
Although there is some uncertainty about the exact location of the South Pole in the late lurassic
(cf.
Struckmeyer,
Totterdell et al. 1990: Gradstein et al. 1992: Wilson et al. 1989). nonetheless there
is
consensus that the southern Gondwana
continents were grouped together at a distance away from the South Pole and that the pole position was situated some distance
off the Pacific
seahoard of the New Zealand region. The pole position. and the resultant latitude lines are those adopted by
Gradstein et al. (1992. fig.
38c).
and Metcalfe (1993.
fig.
8a).
The overseas relationships of New Zealand Late Jurassic Ammonoidea, excluding the presumably oceanic (and therefore
more widely distributed)
Leiostraca, that are summarised in this diagram show a broad distributional pattern around the southern
margin of Gondwana. As noted by
Crame&Howlen(1988,
p.
29). it is likely that such adistribution may have been accomplished
by
means of a circular migration path around the continental block comprising Australasia India
and
Antarctica. However, in
addition to this it is evident that it was also possible for at least some members of the same fauna to migrate along the Andean
Geosyncline to
extend at least as far north as Mexico and southern USA. Areas that are known to have been land in the Late

lurassic are bounded
by
the heavy black line (shoreline after l-lowarth 198
1:
Struckmeyer. Totterdell et al. 1990). Stars indicate
the occurrences of Late
lurassic ammonite taxa that have affinities with those of New Zealand.
terpreted the links between New Zealand and Mexico
in terms of direct trans-Pacific migration. However,
this view is now discounted, as is the related theory of
migration across the Pacific via a Pacifica continent
(Melville 1966, 1981; Nur& Ben Avraham 1977, 1978,
1981, 1982; Kamp 1980; Lovis 1989; Cooper 1989;
Hallam 1994b).
(ii) As commented on by Ross et al. (1992 p. 501), with
reference to ammonite biogeography, New Zealand is
tectonically complex and composed of a collage
oftec-
tonic blocks, some ofwhich probably originated in the
equatorial region or in
the low latitudes of the North-
em Hemisphere (e.g. Tozer 1982; Tozer et al. 1991).
In both New Zealand and Mexico, the occurrences of
the ammonite taxa with Mexican-New Zealand affini-
ties are sited on tectonic blocks that by common agree-
ment have been subject to movement in the past, al-
though by exactly how much and over what length of
time is subject to debate (cf. Bishop et al. 1985;
Coombs 1985;
MacKinnon 1983). The New Zealand

occurrences are sited on the Murihiku and Older
Torlesse (Rakaia) terranes (cf. Fig. 2), both with histo-
ries of lateral displacement (e.g. Cooper 1989). The
Mexican occurrences are sited on the fault-bounded
Tepehuano
Terrane, also with a history of lateral dis-
placement (Sedlock et al. 1993). If the relevant
Jurassic-bearing terranes in New Zealand and Mexico
have moved to the extent advocated by, for example,
Tozer (1982) and Tozer et al.
(1991), and particularly
ifthese terranes had undergone significant movements
in palaeolatitude in Late Jurassic times, then New Zea-
land and Mexico may have formerly been closely ad-
jacent and therefore able to share marine faunas.
(iii)
Anumber of continental assemblies involving assump-
tions of an expanding earth show New Zealand to have
been in close proximity to Mexico in Late Jurassic time
(e.g. Trurnit
1991a,b; Maxlow 1995a,b, 1996a,b) (Fig.
66).
Although options (ii) and (iii) have validity and need to be
considered in any palaeobiographical analysis, both op-
tions involve acceptance of what might be considered un-
orthodox geological theory (i.e. either long-distance travel
ofterrane units, extending into Late Jurassic times; or the
acceptance of an expanding earth hypothesis). However,
considering the vicissitudes endured by Wegener's Conti-
nental Drift theory (cf. Stevens

1980b), an open-minded
approach should be adopted and judgement deferred
pro
tern.
PALAEOCLIMATOLOGY
Most of the palaeogeographical reconstructions used in
discussions of Late Jurassic
palaeoclimatology show New
Zealand in a position very close to the South Pole (e.g.
Rowley 1992;
Moore, Hayashida et al. 1992; Moore, Sloan
et al. 1992; Ross et al. 1992; Valdes
&
Sellwood 1992;
Valdes et al. 1995). This near-polar position for New Zea-
land (i.e.
80-85"s Lat.; see Ross et al. 1992 for example),
has been disputed on general
palaeobiogeographical
grounds (Stevens 1977, 1980a. 1989. 1990a: Stevens
&
Fleming 1978; Broekhuizen 1984; Eagle 1993).
Oxygen isotope results support a mid-latitude position for
New Zealand in the Late Jurassic (Stevens 1971
:
Stevens
&
Clayton 1971; Price
&
Sellwood 1997). It is also ap-

parent that, as discussed above, alternative
palaeogeographical reconstructions, notably those that in-
corporate elements of earth expansion over time,
show the
New Zealand region to be sited in mid-latitudes in the Late
Jurassic (e.g. Maxlow
1996b) (Fig. 66). Also, there are
views that postulate that at least some segments
ofthe col-
lage of terranes that preserve the Mesozoic record
ofNew
Zealand may have been substantially displaced in Late
Jurassic times. For example, in the Late Jurassic some of
these displaced terranes may have had geographic posi-
tions that were significantly northwards
kom those they
occupy today (Tozer 1982; Tozer et al.
l99 1) (see discus-
sion in preceding section).
Although there is some evidence
that climates in the Early
and Middle Jurassic were generally cooler than those in
the Late Jurassic, and that polar ice may have existed in at
least the Early Jurassic (Chandler et al. 1992;
Parish 1993),
there is general recognition that Late Jurassic climates were
relatively more equable than those of today and that the
sub-tropical and warm-temperate climatic zones extended
further into mid-latitudes than they do so today
(Hallam

1994b). Nonetheless, despite the probableabsence ofpolar
ice caps in Late Jurassic times, warm conditions did not
extend as far as the Poles themselves and the polar regions
were typified by cool- or cold-temperate climates (Stevens
1977,
1980a).
As discussed in previous sections, the New Zealand Late
Jurassic ammonite faunas have a very strong Tethyan im-
print
-
with close affinities to regions such as Papua New
Guinea, Indonesia, Himalayas, Middle East, Malagasy and
the Eastern Pacific seaboard (Antarctic Peninsula, South
and
Cenhal America). Other invertebrate faunas mirror
the same affinities (Stevens
1990a; Damborenea
&
Mancenido 1992).
If the postulate is accepted that the Tethyan
faunas were
generally intolerant of cool or cold temperatures (Stevens
1980a), then the strong possibility exists that they were
stenothermal organisms that populated the tropical, sub-
tropical and warm-temperate climatic zones. It follows
then that even if some expansion of these climatic zones
had occurred, the Jurassic-bearing segments of New Zea-
land were probably sited no further southwards than
mid-
latitudes

-
either as part of the Gondwana landmass or as
component units of displaced terranes.
On the other band, if New Zealand was in fact situated in
a near-polar position, as favoured by some reconstructions
(e.g. Ross et al.
1992), then the implication is that warm-
temperate conditionskxtended very close to the South Pole.
However, a difficulty is that at the same time a
well-devel-
oped Boreal ammonite fauna was evident in the Northern
Hemisphere, presumably composed of cool- or
cold-tem-
perate organisms (Stevens 1977, 1980a; Ross et al. 1992).
But yet, no equivalent of the Boreal ammonite fauna has
been identified in the Southern Hemisphere.
I
i
EA
=
East Antarctica
A
=
Arabian Peninsula
WA =West Antarctica
I
=
India
G
=

Greenland M
=
Malagasy
Figure 65
(opposite)
Acceptance of the theory of an expanding earth (see Owen 1976. 1983a.b; Carey 1976, 1983) enables the
presentation of palaeogeographic reconshuctions that provide an alternative version, even if somewhat extreme, to the
reconstructions illustrated
in
Figs 63, 64. These alternative reconstructions show the Pacific Ocean to be completely closed in
the Jurassic, with the landmasses separated by complexes of anastomosing narrow seaways, basins and troughs. The upper
reconstruction has been modified from Shields (1979. 1983) and the lower reconstruction from Carey (1992). In the latter
diagram, the coarse stipple represents the Tethyan
orogenic belt. A= Arabian block:
EA
=
East Antarctica; G
=
Greenland;
I
=
Indian block;
M
=Malagasy; NZ
=
New Zealand; WA
=
West Antarctica.
As may be seen from the diagrams. closure
ofthe Pacific Ocean considerably simplifies the links that occurred between New

Zealand. West Antarctica and South America that have been brought into focus by the present study. Also, particularly in the
instance
ofthe Carey reconstruction, there is amarked simplification in the link that existed betweenNew Zealand and Mexico
for a large part of the Late Jurassic (see also Fig. 66).
Although the absence of Anti-Boreal or Austral elements
may be explained by postulating the presence ofenhanced
oceanic circulation in the Southern Hemisphere, as com-
pared with that of northern regions, there still remains the
proposition that New Zealand was sited substantially fur-
ther northwards than depicted in many
ofthe conventional
continental assemblies.
An adjunct to this discussion is that most of the
palaeoclimatic models (e.g. Moore, Hayashidaet al. 1992;
Moore, Sloan et al. 1992; Valdes
&
Sellwood 1992) show
cold temperatures, with winter ice, in southern Gondwana
in
regions that include the position for New Zealand, as
postulated by these authors. However, it is evident
from
the foregoing discussion that. if New Zealand was at this
time part of such a landmass that had temperatures
sufi-
ciently cold to form winter ice, such temperatures were
very much a feature of the continental interior, and the
coastline around New Zealand had more equable,
warm-
temperate, climates. This geographic situation is compa-

rable to that of the present-day South
Island of New Zea-
land
(40°460S lat.)
-
and reinforces the concept that
New Zealand probably occupied a mid-latitude position
in
the Jurassic (cf. Fig. 66).
Equator
1
/
\
\
~ustralid
Eastern Antarctica
/
South America
Tasmania
Western Antarctica
Figure
66
As a result of studies of global expansion tectonics. Maxlow (1996b) has produced a series of true to scale small earth
models (Maxlow
1996b, figs. 8, 10, 12, 14, 16,
19,
20, 21,24). This photograph is of one such model for the Late Jurassic (see
Maxlow
1996b. fig. 20), produced by Terrella Graphics, Western Aushalia.
The model depicts the progressively developing seaway of Panthalassa. the ancestor of the Pacific Ocean. However, at this

time
the
southern landmasses are
still
largely in contact and Aushalia. New Zealand, Antarctica. Central and South America
could have probably shared shelf faunas (including ammonites), if facies conditions were favourable and if migratory routes
were available. The particularly close geographic relationship
ofNew Zealand and Mexico, as depicted, is likely to be responsible
for the strong New
ZealandiMexican links shown by some of the New Zealand Late Jurassic ammonite taxa.
115
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