Jurassic evolution of Southern Hemisphere marine palaeobiogeographic
units based on benthonic bivalves

Susana E. Damborenea *

Departamento Paleontología Invertebrados, Museo de Ciencias Naturales, 1900 La Plata, Argentina

Received 28 September 2001; accepted 25 February 2002

Abstract

The distribution of benthonic Jurassic bivalve genera in the Southern Hemisphere is analysed here. For this region, palaeobiogeographic
units (biochoremas) are quantitatively characterized according to their biologic contents (mainly levels of endemism). Their evolution
through time is followed from the latest Triassic to the earliest Cretaceous. The Tethyan Realm is undoubtedly the most mature and
persistent through time, with three subordinate units: an Australian unit restricted to the Late Triassic, a North Andean unit, which appears
sporadically as an endemic centre, and an East African unit which is recognisable from Bajocian times onwards. From Late Triassic times,
a South Pacific Realm has been recognised, with a Maorian Province mostly based on the distribution of monotoid genera. A South Andean
unit is also recognisable through most of the Jurassic, and its reference either to the South Pacific unit or to the Tethyan Realm is a matter
of debate. Being a transitional biogeographic setting between Tethyan and South Pacific first-order units, it is included in the South Pacific
unit due to the common presence of antitropical (didemic) genera. The East African unit is included within the Tethyan Realm during the
Jurassic, but during Early Cretaceous times, it splits into two units, one of which was regarded as part of the “South Temperate Realm” by
Kauffman. The rank of all these units changed with time. Throughout the Jurassic, the ecotone between South Pacific and Tethyan
palaeobiogeographic units fluctuated in position with time. The approximate latitudinal location of the ecotonal boundary area and its shift
through time are recognised on the basis of faunal composition along the Andean region. © 2002 Éditions scientifiques et médicales
Elsevier SAS. All rights reserved.

Keywords: Palaeobiogeography; Bivalves; Jurassic; Southern Hemisphere; Endemism; Antitropical distribution; South Pacific; Tethyan

1. Introduction

The history of large-scale biogeographic patterns is
increasingly attracting the attention of biologists, since it is
evident that many biogeographic phenomena develop over
long periods of time. Thus, the origin, establishment and
evolution of palaeobiogeographic units of any rank or
biochoremas (biochores in the sense of Makridin, 1973, and
Westermann, 2000a) are of special interest. These units are
dynamic and change in range and rank through time. Those
based on benthonic faunas do not necessarily have a
coincident history with those based on pelagic taxa. These
aspects are clearly exemplified by the history and evolution

of Southern Hemisphere biogeographic units during Juras-
sic times.

Mainly as a consequence of the unevenness of land mass
distribution on the Earth, the northern–southern asymmetry
in some aspects of marine fauna distribution patterns is well
known at different times (see discussion in Crame, 1996b,
2000). Nevertheless, three major biogeographic units based
on benthonic invertebrates (one low-latitude and two high-
latitude) are recognised for Permian (see Shi and Grunt,
2000, and references therein), Triassic (e.g. Stevens, 1980),
Cretaceous (e.g. Fleming, 1963; Kauffman, 1973; Sohl,
1987; Stevens, 1980) and Cainozoic times (e.g. Fleming,
1963; Hayami, 1989). In contrast, the hemispherical asym-
metry of biogeographic units during most of the Mesozoic
has led many authors to recognise only two first order
palaeobiogeographic units for the Jurassic, mainly on the
basis of the distribution of ammonites: the Boreal and
Tethyan Realms (Hallam, 1969, 1971, 1977; Stevens, 1980,

* Corresponding author.
E-mail address: sdambore@museo.fcnym.unlp.edu.ar (S.E. Dambo-

renea).

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1990; Doyle, 1987; Challinor et al., 1992; Hillebrandt et al.,
1992; and many others, see discussion and references in
Westermann, 2000a,b). As Kauffmann (1973) aptly ex-
pressed, “poor knowledge of south temperate [bivalve]
faunas has led many authors to assume no “anti-Boreal”
Realm existed south of Tethys during the Mesozoic” and he
demonstrated that this is clearly not true for the Cretaceous,
although he admitted that “this was possibly true in the
Early Jurassic”. In fact, lack of data and proper analysis for
the Southern Hemisphere pervades also studies of other
fossil groups (see, for instance, Dommergues et al., 2001).

This twofold division hinders the determination of the
possible role that the austral regions may have had in the
origin and diversification of the biota (see Crame, 1997)
during the Jurassic. Widespread phenomena like austral
endemism or antitropicality are either ignored or their
relative importance is disregarded. Some authors who work
on Southern Hemisphere faunas (especially bivalves) have
challenged this twofold division (see Stevens, 1980, and
references therein; Crame, 1986, 1987, 1992, 1993; Dam-
borenea, 1993, 1996; Enay and Cariou, 1997, and references
therein), but their attempts were dismissed by some authors
(Hallam, 1994a, b; Westermann, 2000a, b) , while others
adhered to a threefold division, at least for some periods of
time within the Jurassic. In fact, several marine palaeobio-
geographic units of low rank have been proposed for the
Southern Hemisphere Jurassic on the basis of the known
distribution of different marine organisms. The relation-
ships, rank and history of these units were recently reviewed
by Enay and Cariou (1997) and Westermann (2000b).

This paper is an attempt to recognise the development
and evolution of biochoremas based on benthonic bivalve
endemism in the Southern Hemisphere. There is no space
here to discuss in detail either the processes involved or the
possible causes of the observed diversification of the rec-
ognised biogeographic units, or the complexities of nomen-
clature involved, and thus all these issues are deliberately
avoided or only mentioned without further discussion. The
paper is then mainly limited within the first rationale of
those listed by Rosen (1992), i.e. the determination of
palaeobiogeographical patterns with delimitation of prov-
inces and realms, as a starting point for further discussion.

Global studies on the distribution of Mesozoic bivalves
have largely been based on data from Cox et al. (1969) (for
instance, Kauffman, 1973; Hallam, 1977). Those databases
had a poor coverage of the Southern Hemisphere, but there
is now a wealth of recently published information from this
region which certainly adds substantial evidence and can be
used in this context. Palaeobiogeographic provinces based
on bivalves were recently analysed quantitatively for the
European Tethyan and Proto-Atlantic (Liu, 1995; Liu et al.,
1998). In contrast, there are no updated comprehensive
palaeobiogeographic analyses based on bivalves for the
Southern Hemisphere, although several contributions re-
lated to palaeobiogeographic issues using these organisms
are available for the South Pacific (Stevens, 1967, 1977,

1980, 1989, 1990; Hayami, 1984, 1987; Grant-Mackie et
al., 2000), Antarctica (Crame, 1987, 1992, 1996a, b) and the
South American margin of the Pacific (Damborenea and
Manceñido, 1979, 1988; Hillebrandt, 1981; Hallam, 1983;
Damborenea, 1993, 1996).

2. Database

Occurrences of Jurassic bivalve species were compiled
from various published sources as well as the author’s own
data, and plotted stage by stage. To frame the analysis, data
from latest Triassic and earliest Cretaceous were added. The
study area is restricted to the palaeo-Southern Hemisphere,
but the database was compiled on a global scale, not only to
provide the necessary framework for the detailed analysis of
southern regions, but also to adequately recognise patterns
of general distribution and endemism.

Data were then gathered within wide areas, each contain-
ing a large variety of habitats. Eleven such areas were
chosen for the present analysis (located in Fig. 1). On the
whole, they represent a wide coverage of the Southern
Hemisphere Jurassic seas, but some important gaps still
exist, limited by the availability of data (Table 1). A brief
description and list of main data sources are given in
Appendix A.

Both the amount and quality of data are, unsurprisingly,
very uneven, but although this precludes serious detailed
quantitative analysis at the moment, the database provides
enough information to obtain a broad framework. The
species’ distribution data compiled were systematically and
stratigraphically updated as far as possible and dubious
records were excluded. For this analysis, data were pro-
cessed at the genus-group level and according to the

Fig. 1. Location of the geographic units used in this study. 1, NW South
America; 2, Perú and northernmost Chile; 3, Central Argentina and Chile;
4, Southern Argentina and Chile; 5, Antarctica; 6, New Zealand–New
Caledonia; 7, Western Australia; 8, Western India; 9, Madagascar; 10,
Mozambique and eastern Africa; 11, Arabia and Iran. Base map for the
Early Jurassic, continent palaeopositions based on Smith and Briden (1977)
and Scotese (1991, 1997), palaeogeography compiled from many sources.

52 S.E. Damborenea / Geobios 35 (M.S. 24) (2002) 51–71



following seven time-intervals: Norian–Rhaetian, Hettan-
gian–Sinemurian, Pliensbachian–Toarcian, Aalenian–Bajo-
cian, Bathonian–Callovian, Oxfordian–Kimmeridgian and
Tithonian–Berriasian. Admittedly, this implies a loss of
detail in the information for some regions, but on the other
hand, it allows the use of some occurrences with uncertain
stratigraphic provenance. The age slice including Tithonian
and Berriasian has the extra drawback that in this way any
fact related to the Jurassic–Cretaceous boundary may be not
noticed in this study. Within the general aims of this paper,
though, this arrangement is satisfactory and gives enough
information to recognise biogeographic units and describe
their overall trends through time.

3. Methods

From the many methods that can be used to recognise
palaeobiogeographic units, a traditional approach based on
endemism has been followed, using the set of rules for their
definition, rank and nomenclature proposed by Westermann
(2000a). Bivalve genera were classified according to their
palaeobiogeographic affinities, following basically Kauff-
man (1973), Stevens (1980) and Damborenea (1993), as
follows (see examples in Fig. 2):

• Pandemic: Widespread bivalves, truly cosmopolitan
forms. It is interesting to note that many genera
traditionally referred to as “cosmopolitan” have not
been reported from the Southern Hemisphere during
the time-interval considered. These include, for in-
stance, Atrina, Cirtopinna, Ctenoides, Limopsis, Linot-
rigonia, Malletia, Martesia, Megalodon, Mytiloides,
Palaeomya, Paratancredia, Procardia, Septifer. Other
so-called “cosmopolitan” genera are present only in the
northern regions of the Southern Hemisphere (1, 2, 8, 9,
10, and 11 of this study) but are not known from the
southernmost areas. Examples are Hippopodium,

Neomegalodon, Pinguiastarte and Rollieria. All the
genera mentioned with such distributions were ex-
cluded from the list of “cosmopolitan” taxa for this
analysis.

• Endemic: These are the key elements to recognise and
characterise biogeographic units. For this study, only
true endemics (from each region compared at the global
scale) were counted as such.

• Low-latitude or Tethyan.
• High-latitude:

„ Austral (= Maorian or palaeoaustral).
„ Boreal.
„ Didemic, antitropical or bipolar, restricted to high-

latitudes and present in both hemispheres, being
absent from the low-latitude intervening areas
(Crame, 1993; Damborenea, 1993; Sha, 1996).

• Trans-temperate (Kauffman, 1973): This category in-
cludes taxa common in temperate regions, but with a
distribution not latitudinally limited, and at the same
time, not pandemic. For instance, bivalves with the
so-called “Pacific” distribution, i.e. along the margins
of the palaeo-Pacific ocean both in low and medium
(sometimes even high) palaeolatitudes, absent from
other areas.

Only pandemic and endemic taxa can be objectively
defined, reference of some genera to either of the other
categories is necessarily a matter of debate. This is not a
serious drawback in this context, however, since these last
categories are only accessory elements in this study. From
the nearly 500 bivalve genera known from this time-
interval, 52% are here regarded as low-latitude, 21% as
high-latitude, 6% as trans-temperate and 21% as pandemic
(Fig. 3).

Since geographic ranges of taxa change through time,
sometimes this results in a different categorization for the
same taxon. For instance, a genus may be endemic to a
certain region during a stage and then become widespread.

Table 1
Overview of the Norian to Berriasian bivalve data from the study areas. Total number of genus-level taxa are given for each time-interval and each region.

Regions
N

E
So

ut
h

A
m

er
ic

a
(1

)

Pe
rú

an
d

no
rt

he
rn

C
hi

le
(2

)

C
en

tr
al

A
rg

en
tin

a
an

d
C

hi
le

(3
)

So
ut

he
rn

A
rg

en
tin

a
an

d
C

hi
le

(4
)

A
nt

ar
ct

ic
a

(5
)

N
ew

Z
ea

la
nd

–N
ew

C
al

ed
on

ia
(6

)

W
A

us
tr

al
ia

–N
ew

G
ui

ne
a

(7
)

W
es

te
rn

In
di

a
(8

)

M
ad

ag
as

ca
r

(9
)

M
oz

am
bi

qu
e

an
d

E
A

fr
ic

a
(1

0)

A
ra

bi
a

an
d

Ir
an

(1
1)

Tithonian–Berriasian 16 74 27 27 11 14
Oxfordian–Kimmeridgian 5 9 31 24 6 31 9 64 6
Bathonian–Callovian 31 23 120 35 57 10
Aalenian–Bajocian 33 69 16 44 24 8 4 26 10
Pliensbachian–Toarcian 5 66 97 33 25 5 16 17 27
Hettangian–Sinemurian 15 54 70 5 6
Norian–Rhaetian 12 47 11 23 17 11

S.E. Damborenea / Geobios 35 (M.S. 24) (2002) 51–71 53



Fig. 2. Some examples of Early Jurassic bivalves from the Central Argentina region (3) according to generic palaeobiogeographic affinities. Specimens
housed at La Plata Natural Sciences Museum (MLP) and National Geological Survey, Argentina (DNGM). (A)–(C) Low-latitude; (A) Ctenostreon
raricostatum (BAYLE AND COQUAND), MLP 28428, Pliensbachian, Piedra Pintada, ×0.44; (B) Lycettia hypertrigona DAMBORENEA, MLP 16308,
Pliensbachian, Piedra Pintada, ×0.7; (C) Gervillaria pallas (LEANZA), MLP 19079, Pliensbachian, Nueva Lubecka, ×0.8. (D)–(F) High-latitude; (D)
Kolymonectes weaveri DAMBORENEA, MLP 23804a, Pliensbachian, Chacay Melehue, ×1; (E) Kalentera nov. sp., MLP 24294, Early Pliensbachian, Atuel
river, ×1; (F) Agerchlamys wunschae (MARWICK) and epizoic Harpax rapa (BAYLE AND COQUAND), MLP 23658, Pliensbachian, arroyo Ñireco, ×1.
(G)–(I) Trans-temperate (Pacific); (G), (H) Weyla alata angustecostata (PHILIPPI), internal and external views of right valve, MLP 19076, Early Toarcian?
Sierra de Agnia, ×0.34; (I) Otapiria neuquensis DAMBORENEA, MLP 16480, Pliensbachian, Piedra Pintada, ×2. (J)–(K) Endemic, J, Groeberella
neuquensis (GROEBER), DNGM 7340, Pliensbachian, Nueva Lubecka, ×1; (K) Gervilletia turgida (LEANZA), MLP 6260, Pliensbachian, Piedra
Pintada, ×0.6.

54 S.E. Damborenea / Geobios 35 (M.S. 24) (2002) 51–71



An example is Kalentera, which was endemic to the
Maorian Province in Late Triassic times but became anti-
tropical during the Early Jurassic. Similarly, Gryphaea was
restricted to high-latitudes during the Triassic but later
became pandemic. Conversely, a previously widespread
taxon may have a temporarily restricted distribution. The
recognition of these subtle changes is highly dependent on
a very detailed knowledge of recorded distributions, and is
thus not always easy to establish in the fossil record.
Nevertheless, these changes were herein recognised and
taken into account as far as possible.

4. Biogeographic units and their evolution

The percentage of cosmopolitan bivalve genera, even
restricted as mentioned above, was high worldwide during
most of the Jurassic (about 21% for the whole Jurassic),
leaving only a few endemic taxa (16%) as diagnostic of
palaeobiogeographic units. It is interesting to note here that
for the whole Jurassic endemic genera are proportionately
equally abundant in high-latitude and in low-latitude re-
gions.

For the following analysis, percentage of endemism was
calculated excluding cosmopolitan forms (see Kauffman,
1973) and including only strictly endemic taxa. In the
following discussion, all the percentages quoted are calcu-
lated over total minus cosmopolitan genera. At different
times through the Jurassic, endemism within the different
areas of the Southern Hemisphere varied, and was used here
to recognise palaeobiogeographic units.

A summary of the results regarding the percentage of
endemic genera in each of the units is presented in Table 2.
These percentages are low according to normalised scales
based on species’ distribution (see discussion in Wester-
mann, 2000a), but are of the same order as those used by
other authors to define palaeobiogeographic units on the
basis of bivalve genera (e.g. Kauffman, 1973), and they
allow the recognition of units of different rank. Neverthe-
less, taking into account the nature of the database, no
attempt is made here to establish a table of minimum values
for each rank. It is interesting to note that the distinction
between Tethyan and Boreal Realms based on bivalves in
the Northern Hemisphere rests on only a few taxa. Accord-
ing to Liu (1995), for instance, for the Pliensbachian, only
the presence of Hippopodium and Meleagrinella is charac-
teristic of the Boreal Realm and Weyla and Lithiotis of the
Tethyan Realm in Europe (note that these taxa were not
even endemic).

Five basic biochoremas and two of higher rank can be
recognised according to the distribution and percentage of
endemic taxa (Fig. 4). Some of the basic units are here
regarded as belonging to the Tethyan Realm, while others
can be grouped in another high rank unit which, following
Westermann’s (2000b) recommendations, is called South
Pacific (Challinor, 1991 = Austral Realm in Damborenea,
1993). Overall endemism for this high rank unit is over 60%
during the latest Triassic, between 15% and 30% for the
Early Jurassic, and over 30% for Middle and Late Jurassic.
These fluctuations suggest its change in rank through time.

All units are further characterised by other aspects, such
as the presence and relative abundance of high-latitude or
strictly low-latitude taxa (Table 2), overall diversity, and the
presence/absence of certain higher rank taxonomic groups.
Antitropical (or didemic) taxa were also common within
monotoids, pectinoids, inoceramoids and other bivalve
groups and add character to some of these units.

On a worldwide scale, this implies the presence of three
first order units during the Jurassic based on bivalve data:

Fig. 3. Percentage of genera with different palaeobiogeographic affinities in
selected bivalve families. Average composition for the whole time-interval
at the top, broken lines indicate average percentage of low-latitude (long
dashes) and high-latitude (short dashes) genera.

S.E. Damborenea / Geobios 35 (M.S. 24) (2002) 51–71 55



Boreal, Tethyan and South Pacific. The distribution of
bivalves and the corresponding proposed palaeobiogeo-
graphic zonation is evidently not symmetrical to the palae-
oequator. In fact, in view of the unbalanced distribution of
land/water masses and the related uneven oceanic current
patterns (which are mostly hypothetic at this stage), such a
palaeobiogeographic asymmetry is only to be expected. The
data analysed here do not support the alternative of over-
stressing this asymmetry to the point of reducing the
palaeobiogeographic zonation for the Jurassic to only two
first order units.

At the family level, all the three first order palaeobiogeo-
graphic units are well characterised. During the Jurassic and
Early Cretaceous, most genera of Anomiidae, Burmesiidae,
Ceratomyopsidae, Cuspidariidae [Dattidae], Diceratidae,
Dicerocardiidae, Isoarcidae, Lithiotidae, Mactromyidae,

Malleidae, Megalodontidae, Myalinidae, Myopholadidae,
Myophoricardiidae, Mysidiellidae, Ostreidae, Pergamidi-
idae, Protocardiidae, Ptychomyidae, Pulvinitidae, Requieni-
idae, Sowerbyidae and Unicardiopsidae are restricted to
low-latitudes and characterise the Tethyan Realm. To these,
a group of families with more than average low-latitude taxa
(Fig. 3) should be added: Laternulidae, Pinnidae, Pros-
pondylidae, Tancrediidae, Astartidae, Arcticidae, Myophori-
idae and Gryphaeidae. The first four of these have no
high-latitude taxa, and the others contain less than average
high-latitude genera.

Families strictly restricted to high-latitudes in the Juras-
sic are fewer; among them Asoellidae, Minetrigoniidae and
Palaeopharidae are present in both Boreal and South Pacific
Realms whereas Sportellidae and Yoldiidae are known only
from the Boreal Realm. There is a consistent group of
families which have more than average high-latitude taxa
and at the same time less than average low-latitude genera
(Fig. 3), all of them diverse and abundant in both Boreal and
South Pacific Realms: Monotidae, Inoceramidae, Oxytomi-
dae, Trigoniidae, Retroceramidae and Buchidae. It should
be pointed out here that updated systematic knowledge of
bivalve faunas is still very uneven in the Southern Hemi-
sphere and that future revisions are likely to alter this
picture slightly.

As these basic units change with time in rank, geographic
spread and even realm to which they are assigned, the
discussion below will follow a stratigraphic order (Figs. 5
and 6) and will be restricted to the Southern Hemisphere.

4.1. Late Triassic

Three basic units can be recognised: the South Pacific
Realm contains the Maorian Province, whilst all the other
regions belong to the Tethyan Realm, with an Australian
Province and a North Andean endemic centre (Fig. 5(A)).

Table 2
Percentages of endemic, low-latitude, high-latitude and trans-temperate genera and subgenera (calculated excluding cosmopolitan taxa) for the
palaeobiogeographic units recognised here for each time-slice. Units not recognizable at a given time are indicated by 0 in the endemic portion of the table
(lack of endemic taxa) or no figures (insufficient data).

Taxa Endemic Low-latitude High-latitude Trans-temperate

First order units: Tethyan South
Pacific

Tethyan South
Pacific

Tethyan South
Pacific

Tethyan South
Pacific

Subordinate
palaeobiogeographic units

E
as

tA
fr

ic
an

A
us

tr
al

ia
n

N
or

th
A

nd
ea

n

So
ut

h
A

nd
ea

n

M
ao

ri
an

E
as

tA
fr

ic
an

A
us

tr
al

ia
n

N
or

th
A

nd
ea

n

So
ut

h
A

nd
ea

n

M
ao

ri
an

E
as

tA
fr

ic
an

A
us

tr
al

ia
n

N
or

th
A

nd
ea

n

So
ut

h
A

nd
ea

n

M
ao

ri
an

E
as

tA
fr

ic
an

A
us

tr
al

ia
n

N
or

th
A

nd
ea

n

So
ut

h
A

nd
ea

n

M
ao

ri
an

Tithonian–Berriasian 14 0 14 8 72 72 50 14 3 17 0 11 25
Oxfordian–Kimmeridgian 13 0 0 50 13 73 50 43 7 0 31 7 0 13
Bathonian–Callovian 8 18 20 88 55 40 0 27 20 4 0 20
Aalenian–Bajocian 17 0 9 15 17 66 82 56 44 0 9 19 28 17 0 11 11
Pliensbachian–Toarcian 0 0 9 7 22 53 46 0 19 27 44 19 20 33
Hettangian–Sinemurian 0 14 0 52 24 10
Norian–Rhaetian 50 14 0 47 50 52 6 0 24 29 0 10 18

Fig. 4. Number of endemic genera through time in the recognised palaeo-
biogeographical units.

56 S.E. Damborenea / Geobios 35 (M.S. 24) (2002) 51–71



The Maorian Province (Diener, 1916) presents very
high endemism: the genera Caledogonia, Heslingtonia,
Hokonuia, Kalentera, Maorimonotis, Ouamouia, Torastarte
and Triaphorus are endemic to New Zealand–New Cale-
donia (region 6 here) during this time. This province is very

well characterised during the Middle and Late Triassic, not
only by the percentage of endemics (47%), but also by the
abundance (29%) of genera with antitropical or austral
distribution, such as Eomonotis, Inflatomonotis, Maoritrigo-
nia, Minetrigonia and Phaenodesmia. Strictly low-latitude
genera are very few (6%).

An Australian Province is also well characterised by
several endemic genera (50%), such as Gervillancea,
Guineana, Krumbeckiella, Prosogyrotrigonia and Somar-
eoides. Low-latitude genera are equally numerous, but
high-latitude and trans-temperate forms are absent.

A North Andean endemic centre extends along the Andes
from Venezuela to northern Chile. The endemic genera
Isopristes, Perugonia and Schizocardita are abundant ele-
ments of the Late Triassic fauna from Peru and northern
Chile (totalling 14%), and some genera with high-latitude
distribution are also present (24%).

The central Argentina–Chile area represents a transitional
zone, with no endemic taxa but with a high percentage
(43%) of high-latitude genera, including Gryphaea, Liostrea
and Phaenodesmia. The abundance of trans-temperate taxa
(28%) is mainly due to genera distributed along the eastern
margins of the palaeo-Pacific in both the hemispheres.
Reference of this transitional area to the Tethyan or the
South Pacific Realm is still doubtful.

Fig. 5. Palaeobiogeographic maps for four time-slices: (A) latest Triassic; (B) Toarcian; (C) Bajocian and (D) Oxfordian. Thick broken lines show the
approximate position of ecotone between Tethyan and South Pacific Realms for each time-slice. Pie diagrams represent percentage of genera according to
palaeobiogeographic affinities. Base maps for the Late Triassic, Early, Middle and Late Jurassic, continent palaeopositions based on Smith and Briden (1977)
and Scotese (1991, 1997), palaeogeography compiled from many sources.

Fig. 6. A summary of the evolution of the biogeographic units here
recognised according to their percentage of endemism.

S.E. Damborenea / Geobios 35 (M.S. 24) (2002) 51–71 57



Extensive areas without Late Triassic outcrops or reliable
palaeontologic data in the key regions prevent adequate
discussion of the boundary between realms at this time.

4.2. Early Jurassic

At the beginning of the Jurassic, the percentage of
endemism is low for all the regions. This is not restricted to
the Southern Hemisphere, but is also true at a global scale:
Liu (1995) concluded that provinciality in the Northern
Hemisphere during Hettangian, Sinemurian and Toarcian
was unrecognisable. Although it is difficult to point a
definite cause, this situation is probably a consequence of
the end-Triassic extinction. Some authors have indicated
that endemic bivalve taxa were relatively more vulnerable
to mass extinction events (Hallam and Miller, 1988). Our
data clearly show this pattern for the end Triassic extinction,
and it is also evident that the biota and endemic centres took
some time to recover (see discussion in Hallam, 1996). For
instance, the lack of endemic genera makes the Maorian unit
difficult to recognise during Hettangian and Sinemurian
times, when, possibly apart from Torastarte, no endemic
bivalves are present in New Zealand–New Caledonia. It is
interesting to note that, based on the available data, the
low-latitude regions of the Southern Hemisphere also con-
tain no endemic genera. On the other hand, in transitional
zones, such as the South Andean areas (2 and 3 here), an
endemic centre appeared during Sinemurian times, with a
few endemic genera (14%), such as Gervilletia (Fig. 2(K)),
Groeberella, Lywea and Quadratojaworskiella. This en-
demic centre is regarded here as part of the South Pacific
Realm despite the relatively high proportion of low-latitude
taxa, due to the common occurrence of genera with anti-
tropical or austral distributions (24%), such as Agerchlamys
(Fig. 2(F)), Asoella, Harpax, Kalentera (Fig. 2(E)),
Palaeopharus and Kolymonectes (Damborenea, 1998,
2002).

By Pliensbachian–Toarcian times (Fig. 5(B)), the Mao-
rian unit (= South West Pacific Province in Hallam, 1977)
had recovered, having 22% of endemic genera, including
Torastarte and Pseudaucella (Grant-Mackie et al., 2000)
and 44% of high-latitude genera. A South Andean endemic
centre persists (see Fig. 6) with a low percentage of
endemics (7%), including Groeberella (Fig. 2(J)) but with a
high percentage of high-latitude taxa (27%), which include
Asoella, Harpax, Kolymonectes (Fig. 2(D)), Palaeopharus,
Ochotochlamys, Praebuchia? and Radulonectites (Dambo-
renea and Manceñido, 1992; Damborenea, 1993, 2002).
This interval is characterised by the highest percentages of
high-latitude taxa for the whole Jurassic in this region and
also the highest levels of trans-temperate (mainly east
Pacific) taxa (see Table 2). A North Andean endemic centre
within the Tethyan Realm is again discernible (9% of
endemic genera), with its southern limit extending farther
south than earlier in the Lower Jurassic (see discussion
below and Fig. 7). The peculiar genera Lithiotis and

Opisoma, characteristic of the Jurassic tropics during
Pliensbachian times (see discussions in Geyer, 1977; Hill-
ebrandt, 1981; Hayami, 1984; Nauss and Smith, 1988;
Aberhan and Hillebrandt, 1999), extended to northern Chile
during the Toarcian, but are absent further south in the
South Andean unit.

Cox (1965) mentioned that the presence of the species
Weyla ambongoensis (THEVENIN) in Toarcian beds of
eastern Africa: “affords a somewhat meagre evidence that a
faunal subprovince comprising the western part of the
present Indian Ocean region and extending over northern
Africa had come into existence”. Nevertheless, data from
this area are scarce, and no endemic bivalve genera have
been reported so far for the Early Jurassic.

4.3. Middle Jurassic

During Middle Jurassic (Fig. 5(C)) and early Late Juras-
sic times, the Southern Hemisphere bivalve benthonic
faunas maintain their high proportion of cosmopolitan
genera, always above 60% of all the taxa, but still some
endemic and high-latitude genera are present.

The two South Pacific subunits maintain their level of
endemism between 15% and 20% for the whole Middle
Jurassic (see Table 2), although at the same time they
experienced the proportional increase of low-latitude taxa.
The influx of low-latitude genera reaches 44% for the

Fig. 7. Shift of the ecotone between the Tethyan and high-latitude units
based on bivalves during the Early Jurassic (Pliensbachian–Toarcian) in a
global context. Data from Smith and Tipper (1986), Smith (1989) and
Aberhan (1998, 1999) for western North America; Hayami (1990) for
northeastern Asia; Liu (1995) and Lui et al. (1998) for Europe; Dambore-
nea (1996) for South America. Base map for the Early Jurassic, continent
palaeopositions based on Smith and Briden (1977) and Scotese (1991,
1997), palaeogeography compiled from many sources.

58 S.E. Damborenea / Geobios 35 (M.S. 24) (2002) 51–71



Maorian and 56% for the South Andean units, and is only
exceeded during the Tithonian–Berriasian. Based on this
evidence, many authors (see Grant-Mackie et al., 2000)
concluded that the Maorian Province ceased to exist in the
early Middle Jurassic. Nevertheless, a palaeogeographic
unit is here still recognised (though probably with lower
rank) throughout the Middle Jurassic. There are two alter-
natives to name this unit for Middle and Late Jurassic times:
it can either be regarded as an extension of the Maorian
Province, or it can be called Austral as the origin of the
Austral Province recognized by Kauffman (1973) for Early
Cretaceous times. The first alternative is followed here, but
regardless of how we choose to call it, the continuity of an
austral biogeographic unit from Triassic (Maorian Province)
to Cretaceous (Austral Province) is evident from the results
of this study. This unit is recognised on the basis of the
following endemic genera: Haastina (Bajocian), Mala-
gasitrigonia (Aalenian–Bajocian), Kanakimya (Aalenian-
–Bajocian?) and Moewakamya (Bathonian–Oxfordian). En-
demic genera include Anditrigonia (Bajocian–Tithonian),
Andivaugonia (Bajocian–Callovian), Eoanditrigonia (Bajo-
cian–Callovian), Neuquenitrigonia (Bajocian), Lamber-
trigonia (Callovian) for the South Andean unit. High-
latitude taxa in these two units are proportionately fewer;
they include Retroceramus (Bajocian–Oxfordian) and
Scaphogonia (Bajocian–Tithonian) for both units, Praebu-
chia? (Aalenian) for the South Andean, and Fractoceramus
(Bajocian) and Hijitrigonia (Bajocian) for the Maorian unit.
Again, probably as a consequence of the end Pliensbachian
extinction event, endemism in the Maorian unit drops from
22% at the late Early Jurassic to 17% at the early Middle
Jurassic. In this connection, Aberhan and Fürsich (2000)
have pointed out that the Pliensbachian–Toarcian extinction
event affected the endemic bivalve species more than others
in the Andean region.

A new East African unit (= Provincia Etiopico-Indo-
Malgascia in Ficcarelli, 1968; Ethiopian Province in Hal-
lam, 1977) appears to be recognisable from Bajocian times
on the basis of its endemic genera: Indolucina (Bajocian-
–Oxfordian), Agrawalimya (Bathonian–Callovian), In-
docorbula (Bathonian–Callovian), Indomya (Bathonian-
–Callovian), Indoweyla (Bathonian–Callovian), and
Venilicyprina (Bathonian–Callovian). This unit contains,
however, a high percentage of strictly low-latitude genera
(66% for Aalenian–Bajocian and 88% for Bathonian–Call-
ovian times) and lacks high-latitude genera. For these
reasons, it is here regarded as part of the Tethyan Realm at
this time.

The North Andean endemic centre is still faintly discern-
ible during the Aalenian, with the presence of the endemic
Gervilleiognoma. Species of Heterostrea, a genus strictly
restricted to low-latitude areas in both hemispheres, extend
not only to southern Perú but also to northern Chile, pushing
further south the boundary between Tethyan and South
Pacific units. Published data from younger times are very
scarce and incomplete, not allowing their use in this context.

4.4. Late Jurassic

The general situation observed for Bathonian–Callovian
times is generally maintained for the Late Jurassic (Fig.
5(D)). The Maorian unit is extended geographically to
include the Antarctic and South Atlantic Plateau regions,
and even some western Pacific localities (Timor, Sula, Buru,
Ceram; see Hayami, 1984), but is somewhat less strongly
recognisable, and its rank should be lowered, since its
overall endemism at the generic level diminishes to 13%
during Oxfordian–Kimmeridgian and to only 8% during
Tithonian–Berriasian times. Endemic genera include
Moewakamya (Oxfordian), Jeletzkiella (Oxfordian–Kim-
meridgian) and Praeaucellina (Tithonian–Berriasian),
whilst Malayomaorica (Oxfordian–Kimmeridgian) has an
austral distribution reaching Australia–New Guinea. Retro-
ceramus, Scaphogonia, Lyapinella? and Anopaea are high-
latitude genera, mostly didemic in distribution.

On the other hand, the South Andean unit maintains its
levels of endemism. The apparently very high percentage of
endemism for the Oxfordian–Kimmeridgian is due to the
small number of taxa reported so far and this figure is not
statistically significant. Endemic genera include the trigo-
nioideans Anditrigonia (Bajocian–Tithonian), Antutrigonia
(Tithonian–Berriasian), Notoscabrotrigonia (Tithonian) and
Splenditrigonia (Tithonian–Berriasian), and Retroceramus
and Scaphogonia represent the high-latitude taxa.

The East African unit maintains a percentage of endemic
taxa between 13% and 14%, including Africomiodon and
Eoseebachia (Oxfordian–Kimmeridgian), with a very high
percentage (over 70%) of low-latitude taxa. It is interesting
to note here that the influence of high-latitude taxa in this
area steadily increases towards the end of the Jurassic.

4.5. Early Cretaceous

By the end of the Jurassic and beginning of the Creta-
ceous, the situation changes and antitropical bivalve genera
are again well established at least within inoceramoids and
monotoids (Crame, 1986, 1993). According to Kauffman
(1973), recognition of a relatively mature “South Temperate
Realm” is clear, with well-defined “Austral” and “East
African” Provinces as subordinate biogeographic units by
the beginning of the Cretaceous. The change of the East
African unit from the Tethyan (during most of the Jurassic)
to the South Pacific Realm is indicated by the relatively high
percentage (14%) of high-latitude taxa during Tithonian-
–Berriasian times, possibly as a direct consequence of the
opening of the Mozambique channel. Kauffman (1973)
includes the East African Province within the Indo-Pacific
Region in the Early Cretaceous, according to its content of
southern Pacific bivalve lineages, but with a geographic
range now restricted to southern Africa, Madagascar and
Mozambique. To the north, a transition zone developed
(India, Arabia, NE Africa) with strong Tethyan influence
(the north Indian Ocean Subprovince in Kauffman, 1973).

S.E. Damborenea / Geobios 35 (M.S. 24) (2002) 51–71 59



During the Early Cretaceous (Berriasian), the South
Andean unit contains a few endemic trigoniodeans, such as
Antutrigonia, Splenditrigonia, and Transitrigonia. Anopaea
is a genus with didemic distribution, and during the Berria-
sian, it is present in New Zealand and Antarctica, whilst the
endemic Praeaucellina lingers from Tithonian times in the
Maorian unit. According to Kauffman (1973), the Austral
Province of the Indo-Pacific Region (South Temperate
Realm) was strongly developed at the beginning of the
Cretaceous, including Australia, New Zealand and New
Guinea.

From Middle Cretaceous time onwards, the North An-
dean unit is clearly recognisable as a Caribbean
Subprovince/Province (Kauffman, 1973), and an Austral
Realm was proposed by Fleming (1963) on the basis of
bivalve and gastropod distribution.

5. Transitional zones

Transitional zones occurred between the Tethyan and
South Pacific first order units. To study the evolution of
these ecotones requires a very detailed set of data and is
outside the scope of this paper. Nevertheless, some broad
features will be pointed out, since it is interesting to
compare them with the evolution of Boreal/Tethyan eco-
tones. The evolution of the boundary between Tethyan and
Boreal Realms has been the subject of many contributions,
most of them based on the distribution of ammonite taxa
(e.g. Imlay, 1965; Hallam, 1969, 1981; Fürsich and Sykes,
1977; Hayami, 1990; Dommergues and Meister, 1991;
Hillebrandt et al., 1992; Hallam, 1994b), and on the basis of
bivalve distribution was recently followed in Europe by Liu
(1995). All the data indicate that a northward migration
from Pliensbachian to Bathonian times took place, and that
the Bathonian can be recognised as the time of the greatest
Tethyan spread during the Jurassic. The boundary then
shifted southwards during the Callovian, when the distinc-
tiveness of the Boreal bivalve fauna strongly increased and
was maximum at the Oxfordian (Liu, 1995). The Callovian
and Early Oxfordian marked the widest spread of the Boreal
Realm according to Hallam (1971).

From the possible transitional zones located in
palaeotemperate regions of the Southern Hemisphere, the
relationship between Maorian-type and other faunas in the
western Pacific was studied by Hayami (1984). In the
eastern Pacific, the South Andean ecotone has been recog-
nised for Early Jurassic times (Fig. 7) and described in detail
(Damborenea, 1996). Based on species’ distribution along
the eastern palaeo-Pacific margin, a southward shift of this
boundary of about 8–10° latitude has been proposed for the
Hettangian–Toarcian interval. It is interesting to compare
these observations with the evolution of the boundary areas
between Tethyan and Boreal Realms in the Jurassic. To
make this comparison, the boundary areas proposed on the
basis of bivalve distribution (Hayami, 1984; Smith and

Tipper, 1986; Smith, 1989; Hayami, 1990; Liu, 1995;
Damborenea, 1996; Liu et al., 1998; Aberhan, 1998, 1999),
which are not always coincident with the limits proposed on
the basis of ammonites, have been plotted in Fig. 7 for the
Pliensbachian and Toarcian. The results show a high con-
gruence between the behaviour of the boundary in both
hemispheres during the Early Jurassic: the southward shift
in the Southern Hemisphere is matched by an equivalent
northward shift in the Northern Hemisphere (Fig. 8). There
are not yet detailed enough data from the Southern Hemi-
sphere to follow this comparison for later Jurassic times. It
is interesting to point out, though, that the greatest expan-
sion of low-latitude bivalve faunas in the Southern Hemi-
sphere occurred during Middle Jurassic times (see also
Grant-Mackie, 2000), and that the northward shift of the
Maorian/Tethyan boundary during Late Jurassic times in
western Pacific regions, albeit somewhat complicated by the
tectonic history of the area (Hayami, 1984, 1987), roughly
corresponds with the widest expansion of the Boreal Realm
in the Northern Hemisphere. These coincidences suggest
that the main faunal shifts are related in their origin.

6. Brief comparison with data from other organisms

Since this analysis is only based on one group of
organisms, it is interesting to compare the results with those
obtained using other groups, with the aim of meaningfully
evaluating the evolution of palaeobiogeographic patterns
during the Jurassic. Nevertheless, most other benthonic
invertebrate groups (gastropods, corals and echinoderms)
are still insufficiently known to be used in palaeobiogeog-
raphy for the Jurassic at a global scale. From the scattered
available data, it is evident that some groups of restricted

Fig. 8. Approximate palaeolatitude variation of the Tethyan/Boreal and
Tethyan/South Pacific ecotones through the Jurassic, indicating possibly
symmetrical expansion of Tethyan Realm during Middle Jurassic. Data
compiled from various sources (see discussion in text). Solid lines: average
positions proposed by different authors on the basis of bivalve distribution.

60 S.E. Damborenea / Geobios 35 (M.S. 24) (2002) 51–71



tropical affinities, such as reef building corals and certain
sponges like Stylothalamia, are completely absent from the
Southern Hemisphere low-latitude areas belonging to the
South Pacific palaeobiogeographic unit as understood in this
paper (Hillebrandt, 1981; Crame, 1987; Beauvais, 1992).
Gastropods have been used to recognise palaeobiogeo-
graphic units for the Cretaceous (Sohl, 1987), but are poorly
known on a global scale from Jurassic and Triassic. Within
these limitations, Tong and Erwin (2001) noted that the
relationships between Triassic Tethyan and American gas-
tropods are weak, even at the generic level. Brachiopods
from austral regions do show a considerable degree of
endemism which supports the recognition of a Maorian
Province during the Late Triassic and Early Jurassic (Mac-
Farlan, 1992; Manceñido and Dagys, 1992); the endemism
extends during Middle and Late Jurassic in New Zealand-
New Caledonia (MacFarlan in Grant-Mackie et al., 2000).

As expected (see discussion in Masse, 1992), there are
important differences between the results obtained in this
study and the palaeobiogeographic units recognised on the
basis of pelagic organisms. One of the most interesting
questions that needs further discussion is the observed
differences in the first order palaeobiogeographic units for
the Southern Hemisphere between those based on bivalves
as here described and those based on ammonites. According
to the pattern of ammonite distribution, most authors
consistently recognise only two realms (Boreal and
Tethyan) for the Jurassic (see references in Cecca, 1999;
Westermann, 2000b; Grant-Mackie et al., 2000). Hill-
ebrandt (1981) and Hillebrandt et al. (1992) distinguished
an Eastern Pacific and an Indo-SW Pacific Subrealm during
some lapses of time within the Early and Middle Jurassic,
but they admit that for the Late Jurassic, lack of data
severely constrains the discussion of ammonite provincial-
ism in circum-Pacific regions. Dommergues et al. (2001)
recently reviewed the distribution of Early Jurassic am-
monoids trying to match different morphologic sets previ-
ously recognised with analytical distribution patterns, de-
fined by considering the distribution and abundance of
species, but they did not use data from the southern
high-palaeolatitude regions. In fact, a different picture
seems to be emerging as new data are known. Riccardi
(1991) found a nearly continuous presence of endemic
ammonites from Pliensbachian to Oxfordian in South
America, with a sharp increase during Aalenian, and almost
no endemics during Kimmeridgian–Berriasian. Westermann
(1996) recognised a high proportion of endemic taxa for the
Bajocian of New Zealand, with a marked increase in
Andean affinities for the Bathonian–Callovian (see Grant-
Mackie et al., 2000, for references). According to Enay and
Cariou (1997), an austral ammonite fauna of low diversity
became progressively better established around east and
south Gondwana from Oxfordian times.

Knowledge of Jurassic radiolarians from the Southern
Hemisphere is just emerging (see Kiessling and Scasso,
1996; Aita et al. in Grant-Mackie et al., 2000), but clearly

points to non-Tethyan affinities for the Late Jurassic faunas
from Antarctica and the Waipapa Terrane (New Zealand).
These data support a palaeolatitudinal zonation for Late
Jurassic radiolarian faunas in the Southern Hemisphere,
with a distinct austral palaeobiogeographic unit (which is
not symmetrical with the boreal one), characterised by low
endemism and dominated by Parvicingula–Praeparvicun-
gula (Kiessling and Scasso, 1996).

Only Boreal and Tethyan Realms are also recognised on
the basis of the belemnite distribution (Stevens, 1973) but
according to Challinor et al. (1992) a South Pacific Province
(referred to the Tethyan Realm) may be distinguished in the
Jurassic. Belemnitina (which define the Boreal Realm) are
dominant in Boreal regions throughout the Jurassic but are
also known in South America–New Zealand–New Cale-
donia up to the Middle Jurassic (Challinor et al., 1992;
Doyle et al., 1997).

Brief references to palaeoclimate and palaeogeography
can be made to explain these differences. According to
Golonka and Ford (2000), greenhouse conditions prevailed
during the Sinemurian–Toarcian, with a warm, humid envi-
ronment, and moderate temperatures into high-latitudes
with no evidence of significant continental glaciation. Based
on several evidences, Price (1999) concluded that the extent
of polar ice during the Mesozoic was probably only one-
third the size of the present-day. Kiessling and Scasso
(1996) suggested that Antarctic surface waters may have
been warmer on average than those in equivalent northern
high-latitudes, according to the distribution of Pantanelli-
idae radiolarians. During the Jurassic, climates are said to
have been milder, temperatures more equable across lati-
tude, but with some seasonality due to the different day
lengths through the year. This may have been a particularly
important factor affecting marine benthonic biota at high-
latitudes. Pelagic taxa are probably less affected by season-
ality than benthonic inhabitants of littoral environments,
such as bivalves. But apart from climate, purely geographic
factors, affecting the ocean circulation patterns, may have
had considerable influence. For instance, the distribution of
land and sea during the Jurassic in high-latitude regions;
whilst in boreal regions there was a polar ocean, nearly
completely surrounded by land or shallow seas, the austral
regions were part of the proto-Pacific ocean, which was
continuous with the eastern Tethys. Enay and Cariou (1997)
emphasized that the lack of such geographical trap in austral
regions explains why the austral ammonite faunas were
never as distinct from the Tethyan as the Boreal ones. In
addition, an important migration route for benthonic organ-
isms since the Early Jurassic, the Hispanic Corridor (Dam-
borenea and Manceñido, 1979; Smith, 1989; Boomer and
Ballent, 1996; etc.), remained as a barrier for most pelagic
animals until Late Middle or even Late Jurassic times (Elmi,
1993; Damborenea, 2000; Aberhan, 2001).

Regarding the evidence from terrestrial organisms,
Balme (in Grant-Mackie et al., 2000) indicated that there is
general agreement that latitudinal zonations based on

S.E. Damborenea / Geobios 35 (M.S. 24) (2002) 51–71 61



megafossil plant distribution existed in the Jurassic, but
these do not support the existence of substantial climatic
barriers.

7. Conclusions

A preliminary, hierarchical, palaeobiogeographic frame-
work is proposed for the Southern Hemisphere Jurassic
based on the distribution of bivalve genera. Two first order
palaeobiogeographic units (Realms or Subrealms) are rec-
ognised for this region:

• Tethyan Realm
• South Pacific Realm/Subrealm: This unit, though not

symmetrical with the Boreal Realm of the Northern
Hemisphere, is clearly recognisable according to bi-
valve data.

The following five basic units are characterised accord-
ing to the content of endemic genera and subgenera:

• North Andean [Endemic centre]: Though data are very
few, it is distinctive sporadically as an endemic centre
during the Late Triassic and Pliensbachian–Bajocian.
The geographic range varies also with time, but
roughly includes areas 1 and 2 of the present study
during the Norian–Rhaetian and Pliensbachian–Toar-
cian, but only area 2 (due to lack of data) for Aalen-
ian–Bajocian.

• East African [Endemic centre → Subpro-
vince → Province]: Includes Madagascar, Mozam-
bique and western India (areas 8, 9 and 10 here) during
the Jurassic. On the basis of bivalve endemic genera, it
begins to be recognisable as an endemic centre by
Middle Jurassic times (Bathonian–Callovian), forming
the origin of the East African Province, well established
at the beginning of the Cretaceous.

• Australian [Province → vanishes]: Includes Australia,
New Guinea and surrounding areas (area 7 here). Well
characterised during the Late Triassic, it is not recog-
nisable during the Jurassic.

• Maorian [Province → Subprovince]: Very well charac-
terised during the Late Triassic, it is almost not recog-
nisable at the beginning of the Jurassic, but recovers
quickly (though never attains the same levels of ende-
mism) and from Pliensbachian times maintains an
almost steady amount of endemic taxa, declining again
at the end of the Jurassic. In the Early Cretaceous, it is
known as Austral Province. The geographic range
varies, apart from New Zealand–New Caledonia (area
6) it included Antarctica (area 5) at some intervals.

• South Andean [Endemic centre → Subprovince]: From
the Hettangian it is recognisable as an endemic centre,
from Middle Jurassic (Bajocian) onwards, it maintains
a moderate level of endemism and can be regarded as a
subprovince. It includes areas 3 and 4.

The North Andean and Australian units are included
within the Tethyan Realm, whilst the Maorian and South

Andean units are grouped together in the South Pacific first
order unit. The East African units begins to develop as part
of the Tethyan Realm, but near the end of the Jurassic, it
begins to experience the increasing influence of South
Pacific faunas, especially in its southernmost areas (south-
ern Africa and Madagascar)

Although significant progress has been made on under-
standing the distribution of Southern Hemisphere bivalves
during the Jurassic, many unsolved questions still remain.
This paper is a contribution to Southern Hemisphere bivalve
palaeobiogeography, to be complemented (and hopefully
integrated) with Northern Hemisphere schemes.

Acknowledgements

I am grateful to Dr. G. Westermann, Dr. F. Cecca and the
organizing committee of the International Conference Pa-
leobiogeography and Paleoecology 2001 for their invitation
to present this paper, and to the La Plata University for a
travel grant to attend. Drs. J. Grant-Mackie and A. Crame
made constructive reviews and provided helpful suggestions
for the improvement of the manuscript. Research was
funded by the Consejo Nacional de Investigaciones Cientí-
ficas y Técnicas (CONICET), Argentina. This is a contribu-
tion to IGCP Project 458 “Triassic–Jurassic boundary
events”.

Appendix A

A.1Brief description of regions and data sources

1 NW South America (Venezuela, Colombia, northern-
most Perú) (Geyer, 1973, 1979; Guzmán, 1984, and refer-
ences therein): Though rich faunas are known to occur, the
published information from this region is very scarce and
cannot be compared with the other data sets, being confined
to Norian to Toarcian and Tithonian–Berriasian.

2 Perú and Northern Chile: Comprises most of Perú and
the northernmost regions of Chile (up to about 26° S
present-day latitude) References: listed in Damborenea
(1996); and Jaworski (1915, 1922), Körner (1937), Cox
(1949, 1956), Harrington (1961), Pérez and Reyes (1977,
1983, 1985, 1986, 1991), Hayami et al. (1977), Westermann
et al. (1980), Chong and Hillebrandt (1985), Riccardi et al.
(1990a, b), Pérez et al. (1987), Romero et al. (1995),
Aberhan and Hillebrandt (1996, 1999), Rubilar (1998).

3 Central Argentina and Chile: Comprises what is
regionally known as the Neuquén basin. References: listed
in Damborenea (1996); and Burckhardt (1900a, b, 1903),
Haupt (1907), Jaworski (1914, 1915), Stehn (1923), Fuen-
zalida Villegas (1937), Leanza (1941), Lambert (1944),
Sokolov (1946), Levy (1967), Thiele Cartagena (1967),
Cecioni and Westermann (1968), Hallam et al. (1986),
Damborenea (1990), Leanza and Garate-Zubillaga (1987),

62 S.E. Damborenea / Geobios 35 (M.S. 24) (2002) 51–71



Riccardi (1988), Riccardi et al. (1990a, b, c, 1997), Lo Forte
(1988), Damborenea et al. (1992), Leanza (1993), Malchus
and Aberhan (1998), Rubilar (1998), Damborena (2002) and
unpublished data. Well documented for the whole time-
interval of the present study.

4 Southern Argentina and Chile: References: listed in
Damborenea (1996); and Feruglio (1936), Leanza (1968),
Riccardi (1977), Olivero (1988) and unpublished data. Very
few data known, only from Pliensbachian–Toarcian and
Tithonian–Berriasian, Damborena (2002).

5 Western Antarctica: References: Stevens (1967), Th-
omson and Willey (1972), Thomson (1975a, b, 1981, 1982),
Willey (1975a, b, c), Jones and Plafker (1976), Quilty
(1978, 1982, 1983), Edwards (1980), Crame (1981, 1982a,
b, 1983, 1984, 1985, 1996a), Jeletzky (1983), Medina and
Ramos (1983), Thomson and Tranter (1986), Riccardi et al.
(1990c), Doyle et al. (1990), Thomson and Damborenea
(1993), Crame et al. (1993), Crame and Kelly (1995), Kelly
(1995). This region comprises mainly the Antarctic Penin-
sula and adjacent areas. The few data available from the
Falkland Plateau have been included here.

6 New Zealand–New Caledonia: References: Trechmann
(1918, 1923), Marwick (1935, 1953, 1956), Avias (1953),
Fleming (1959, 1962, 1964, 1987), Grant-Mackie (1960,
1976a, b, 1978a, b, c, d, 1980a, b), Speden (1970), Stevens
(1978), Freneix et al. (1974), Speden and Keyes (1981),
Begg and Campbell (1985), Crampton (1988), Grant-
Mackie and Silberling (1990), Damborenea and Manceñido
(1992), Damborenea (1993) and unpublished data (in Hud-
son, 1999). Very well documented for the whole time-
interval of the present study.

7 Western Australia, New Guinea and Sula Islands:
References: Etheridge (1910), Whitehouse (1924), Teichert
(1940), Skwarko (1967, 1973, 1974, 1981a, b, 1983),
Coleman and Skwarko (1967), Skwarko et al. (1976), Sato
et al. (1978), Grant-Mackie (1994).

8 Western India: References: Kitchin (1903), Cox
(1935b, 1937a, 1940, 1952), Agrawal (1956a, b), Kanjilal
and Singh (1973, 1980), Singh and Kanjilal (1974, 1977,
1982), Agrawal and Rai (1978), Kanjilal (1979a, b, 1980a,
b, 1981), Singh and Rai (1980), Singh et al. (1982), Jaitly
and Singh (1983), Pandey and Agrawal (1984), Jaitly
(1986a, b, c, 1988, 1989, 1992), Jaitly et al. (1995), Pandey
et al. (1996), Fürsich and Heinze (1998), Fürsich et al.
(2000). No data for Late Triassic and Early Jurassic are
available.

9 Madagascar: References: Newton (1889, 1895), Dou-
villé (1904), Thevenin (1908), Barrabé (1929), Besairie
(1930), Nicolaï (1950–1951). Reliable data only from
Pliensbachian times onwards.

10 Mozambique and eastern Africa: References: Weir
(1930, 1938), Cox (1935a), and references therein, (1937a,
b), Venzo (1949), Jaboli (1952), Desio et al. (1960), Cox
(1965), Ficcarelli (1968), Jordan (1971). Reliable data only
from Pliensbachian times onwards.

11 Arabia and Iran: References: Fischer (1915), Cox
(1936), Arkell (1956), Fantini-Sestini (1966), Kalantari
(1981), Lewy (1982), Hautmann (2001). Data are few and
very uneven, bivalves from this region are in need of
revision.

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