See

discussions, stats, and author profiles for this publication at:

https://www.researchgate.net/publication/223375824

Bivalve palaeobiogeography and the

Hispanic Corridor: Time of opening and
effectiveness of a proto-Atlantic seaway
ARTICLE in PALAEOGEOGRAPHY PALAEOCLIMATOLOGY PALAEOECOLOGY JANUARY 2001
Impact Factor: 2.34 DOI: 10.1016/S0031-0182(00)00172-3

CITATIONS

READS

63

103

1 AUTHOR:
Martin Aberhan
Museum fr Naturkunde - Leibniz Insti
73 PUBLICATIONS 1,786 CITATIONS
SEE PROFILE

Available from: Martin Aberhan

Retrieved on: 01 March 2016

Palaeogeography, Palaeoclimatology, Palaeoecology 165 (2001) 375394

www.elsevier.nl/locate/palaeo

Bivalve palaeobiogeography and the Hispanic Corridor:

time of opening and effectiveness of a proto-Atlantic seaway
Martin Aberhan *
Museum fur Naturkunde, Institut fur Palaontologie, Invalidenstr. 43, D-10115 Berlin, Germany
Received 8 December 1999; accepted for publication 19 July 2000

Abstract
The Hispanic Corridor is an embryonic seaway between the eastern Pacific and western Tethyan oceans that has
been postulated to have preceded Middle Jurassic drifting and the birth of the Atlantic Ocean by many millions of
years. In this study the distribution of pectinoid bivalve morphotypes is analysed to examine the origin of the Hispanic
Corridor and its effectiveness during Early Jurassic times. A comparison of pectinoid faunal similarities, based on
similarity coefficients, between various regions at opposite ends of the Corridor suggests that it opened progressively
during Early Jurassic times. However, with this approach it is difficult to exclude alternative dispersal routes, and to
determine when faunal interchange began. Analysis of the percentage of pectinoid morphotypes that were (1) present
at opposite sides of the Corridor, (2) simultaneously absent in the western Pacific/eastern Tethys and (3) confined to
relatively low palaeolatitudes provides evidence that the Hispanic Corridor developed from an effective barrier in
earlier Early Jurassic (Hettangian to Sinemurian) times into a filter, allowing the passage of a few morphotypes,
during later Early Jurassic (Pliensbachian and Toarcian) times. The apparently two-way faunal exchange through the
Hispanic Corridor is consistent with the establishment of a megamonsoonal circulation for Pangaea, which may have
caused seasonally changing directions of oceanic surface currents through the Corridor. 2001 Elsevier Science B.V.
All rights reserved.
Keywords: bivalves; Hispanic Corridor; Jurassic; palaeobiogeography; palaeoceanography

1. Introduction
One of the most prominent palaeogeographic
changes in earth history was initiated by the Early
Mesozoic breakup of the supercontinent Pangaea.
Although there is no direct sedimentological or
geophysical evidence, palaeontological data suggest that prior to drifting and the birth of the

* Fax: +49 30 20938868.

E-mail address: martin.aberhan@rz.hu-berlin.de (M. Aberhan)

Atlantic Ocean a shallow marine connection was

established across rifting continental crust linking
the western end of the Tethyan Ocean with the
eastern Pacific ( Fig. 1). The time of formation
of this narrow passageway, called the Hispanic
Corridor (Smith, 1983), and its subsequent effectiveness for the equatorial dispersal of marine
organisms is the subject of continuous debate and
was certainly controlled by both plate-tectonic
activity and eustatic sea-level changes (e.g. Hallam,
1983; Smith and Tipper, 1986; Smith et al., 1990;
Westermann, 1993; Stanley, 1994).

0031-0182/01/$ - see front matter 2001 Elsevier Science B.V. All rights reserved.
PII: S0 0 3 1 -0 1 8 2 ( 0 0 ) 0 0 17 2 - 3

376

M. Aberhan / Palaeogeography, Palaeoclimatology, Palaeoecology 165 (2001) 375394

Fig. 1. Early Jurassic (Pliensbachian) palaeogeography (modified from Barron et al., 1981; Damborenea, 1987) and possible dispersal
routes for marine organisms (modified from Hallam, 1983, 1994). C1=Hispanic Corridor; C2=Viking Corridor; P1, P2=routes
around Pangaea margins; T=trans-Pacific route.

The Early Jurassic, divided into four stages

(Hettangian,
Sinemurian,
Pliensbachian,
Toarcian), and estimated to span about 26 million
years (Gradstein et al., 1994), is a critical time
interval in the geological history of the Hispanic
Corridor. Based on the biogeography of the Early
Jurassic bivalve genus Weyla, Damborenea and
Mancenido (1979) postulated the presence of a
shallow seaway connecting the western Tethyan
and eastern Pacific oceans as early as Pliensbachian
times. Biogeographical evidence from a variety of
fossil groups offers support for this hypothesis (see
below). In recent years, our knowledge of Early
Jurassic bivalves along the eastern palaeo-Pacific
margin has increased substantially for both western
South America (e.g. Damborenea, 1987, 1996;
Aberhan, 1994) and western North America
(Aberhan, 1998a). In the present paper, new data
on the distribution of a particular group of
bivalves, the pectinoids, are used to: (1) analyse
the permeability of the Hispanic Corridor during
the various stages of the Early Jurassic; (2) investigate if dispersal was preferentially one-way (west
east or eastwest) or if a two-way faunal exchange
prevailed; and (3) consider Early Jurassic palaeoceanography in the light of the results obtained
from (2).

2. Early Jurassic palaeogeography and the

dispersal of marine organisms
During the Early Jurassic, the supercontinent
Pangaea, surrounded by the superocean
Panthalassa or palaeo-Pacific, was nearly symmetrical about the equator. About 56% of the land
was in the northern hemisphere (Parrish, 1993;
Fig. 1). As the supercontinent began to break up
and sea level rose during the Jurassic, several
epeiric seaways became established, separating the
areas of exposed land (e.g. Hallam, 1994). Apart
from the Hispanic Corridor, another seaway,
termed the Viking Corridor by Westermann
(1993), opened in Late Pliensbachian time between
Greenland and Norway and connected the Arctic
and Tethys oceans.
Possible migration routes of shelf faunas
between the eastern Pacific and western Tethys
are indicated in Fig. 1. Apart from utilizing the
Hispanic or Viking corridors, organisms could
have migrated around the periphery of Pangaea,
either by the north or the south, or could have
crossed the palaeo-Pacific, either as long-range
dispersal of larvae or via island hopping. Finally,
it has been proposed that fossil biotas could have
been transported across the palaeo-Pacific on

M. Aberhan / Palaeogeography, Palaeoclimatology, Palaeoecology 165 (2001) 375394

various displaced terranes, which now form large

parts of the North American Cordillera, but originated in more remote positions in the ancient
Pacific. However, for the bivalve bearing Early
Jurassic terranes a location close to the western
margin of Pangaea is most plausible (e.g. Aberhan,
1998b and references cited therein).

3. Materials and methods

3.1. Early Jurassic pectinoid bivalves
The present analysis is based on the distribution
of bivalves of the order Pectinida. Representatives
of this sessile group of organisms are fairly
common and well preserved in the studied areas
where they occur in a wide range of environments.
Their diversity is well documented in the literature,
and a uniform taxonomic concept can be applied.
What little is known about the larval development
of Jurassic pectinoids (Palmer, 1989) and their
modern counterparts ( Waller, 1991) suggests that
they possessed planktotrophic larvae, and therefore depended on oceanic currents for their dispersal (but see Waller, 1993 for an example of a
lecithotrophic-type larval shell in a reef-adapted
pectinid from the Caribbean). Recently, distributional patterns of pectinoid bivalves proved useful
in constraining the Early Jurassic tectonic evolution of western Canadian terranes (Aberhan,
1998b, 1999), thus testifying to their high potential
in geologically orientated palaeobiogeographic
analyses.
3.2. The morphotype concept
Studies on Jurassic bivalve biogeography are
usually carried out at the generic/subgeneric level.
Compared with species-based data sets, this clearly
results in the loss of potentially useful information.
On the other hand, a sound taxonomic database
at species level is not feasible in most cases, especially if large data sets are involved. The morphotype
concept, as applied in the present study, forms a
compromise. A morphotype groups together
species that exhibit very similar morphological

377

adaptations and are obviously closely related.

Although a classification of taxa in morphotypes
does not reach the acuity of a species-level data
set, a much finer taxonomic (and biogeographic)
resolution is obtained than from genera alone. It
avoids taxonomic biases introduced by misidentifications and contrasting species concepts of
different authors, and yields a uniform taxonomic
concept in a manageable amount of time without
having to revise a whole fauna at the species level.
A definition of pectinoid morphotypes, which are
named after a characteristic representative, is presented in Appendix A.

3.3. The studied areas and temporal framework

The geographic areas selected for the study were
lumped into five major geographic units. These
are: (1) northwestern Europe (Great Britain,
Sweden,
Denmark,
Belgium,
Luxemburg,
Germany, Switzerland, northern France); (2) western Tethys (SW-France, Italy, Spain, Portugal,
Morocco, Algeria); (3) western South America
(Colombia, Peru, Chile, Argentina); (4) western
North American craton (Huayacocotla trough of
eastern Mexico, western Nevada, northern
California, cratonal areas of western Canada); and
(5) western North American terranes (Antimonio
terrane of NW-Mexico, Wrangellia, Stikinia,
Quesnellia, Cadwallader terrane). The areas of the
last category are crustal fragments that are allochthonous relative to the autochthonous craton, but
palaeogeographic evidence suggests proximity to
the North American craton throughout the Early
Jurassic (e.g. Smith and Tipper, 1986; Aberhan,
1998b). Data for comparison with the western
Pacific were included where deemed necessary.
The data (see Appendix B and references cited
therein) were analysed separately for four time
intervals corresponding to the four stages of
the Early Jurassic: Hettangian, Sinemurian,
Pliensbachian and Toarcian. Only records with
illustrations entered the database, provided that
the specimens were well preserved enough to allow
assignment to a morphotype and the age was
determinable at the stage level. Faunal affinities of

378

M. Aberhan / Palaeogeography, Palaeoclimatology, Palaeoecology 165 (2001) 375394

the various areas to each other are established by

calculating Dice (D) and Jaccard (J ) similarity
coefficients. These coefficients are defined as
D=[2C/(N +N )]100
1
2
and
J=[C/(N +N C )]100
1
2
where N is the number of taxa in sample 1, N is
1
2
the number of taxa in sample 2, and C is the
number of taxa common to both samples.

4. Results
4.1. Comparison of faunal similarities
Dice and Jaccard similarity coefficients show
exactly the same temporal trends (e.g. Fig. 2A),
and therefore only Dice coefficients are reproduced
in Fig. 2B and C.
In order to gain an overall impression of the
faunal similarities between the eastern Pacific and
the western Tethys and adjacent epicontinental
seas of NW-Europe, data for the three eastern
Pacific areas and those for the two European/NWAfrican areas were combined (Fig. 2A). The
similarity of pectinoid bivalves increases steadily
from comparatively low values in the Hettangian
and Sinemurian to intermediate values in the
Pliensbachian and highest values in the Toarcian.
If the data from the various eastern Pacific areas
are compared with the western Tethyan Ocean
(Fig. 2B), pectinoids from the North American
craton and those from the terranes show a trend
of increasing similarity to the western Tethys with
time. Dice coefficients between South America and
the western Tethys are higher than those for the
other two Pacific areas. Values fluctuate around a
level of 45 from Hettangian to Pliensbachian times,
and show a marked increase in the Toarcian. In a
very similar way, similarity values between both
the North American craton and the terranes and
those from NW-Europe (Fig. 2C ) increase through
time. Similarity between the South American
craton and NW-Europe decreases from the

Fig. 2. Early Jurassic similarities of pectinoid bivalve morphotypes between western Tethys, NW-Europe and various eastern
Pacific regions. (A) Increasing Dice and Jaccard coefficients
between eastern Pacific and Europe/NW-Africa suggest progressive opening of the Hispanic Corridor. (B) Dice coefficients
between western Tethys and various eastern Pacific areas. (C )
Dice coefficients between NW-Europe and various eastern
Pacific areas.

Hettangian to lower values in the Sinemurian and

Pliensbachian and reaches a peak in the Toarcian.
4.2. Dispersal of selected morphotypes
As a next step we analyse how the distribution
of morphotypes can contribute to the reconstruction of the early development of the Hispanic

379

M. Aberhan / Palaeogeography, Palaeoclimatology, Palaeoecology 165 (2001) 375394

Corridor. In order to preclude alternative dispersal

routes, a taxon must fulfil three criteria, at least
during the early phases of its duration: (1) presence
on either end of the Hispanic Corridor; (2) simultaneous absence in western Pacific regions; and
(3) confinement to relatively low palaeolatitudes.
Of the 48 morphotypes considered in this study,
15 meet criterion (1) (see Appendix A). In order
to evaluate criterion (2), the possible presence of
these morphotypes in the North-East and Far East
of Russia, in Japan and in New Zealand was
explored. A survey of the relevant literature reveals
that seven morphotypes appear to be absent from
western Pacific regions, at least throughout Early
Jurassic times, and therefore meet criterion (2):
Placunopsis striatula, Pseudopecten equivalvis,
Camptonectes auritus, Camptonectes subulatus,
Eopecten abjectus, Eopecten velatus and Weyla
meeki. To these may be added, Oxytoma inequivalvis, Propeamussium pumilum and Chlamys textoria. Although known from western Pacific areas
during Early Jurassic times (see Efimova et al.,
1968; Milova, 1976, 1985, 1988; Hayami, 1975;
Sey, 1984; Sey and Polubotko in Damborenea
et al., 1992), all these records are younger than
their first occurrences at opposite ends of the
Hispanic Corridor. The difference in age is smallest
for Propeamussium pumilum. Its first appearance
in the Pliensbachian of Europe (Johnson, 1984) is
followed by apparently contemporaneous occurrences in the Toarcian of South America and
North-East Russia. However, as records from
South America are from the Early Toarcian (e.g.
Hillebrandt and Schmidt-Effing, 1981; Aberhan,
1994), they predate the mid-Toarcian occurrence
in eastern Russia (Polubotko and Repin, 1988),
and consequently this morphotype is potentially
useful in constraining the early history of the
Hispanic Corridor.
From the five morphotypes that do not meet
criterion (2), one (Meleagrinella substriata)
appears to have occurred first during the
Hettangian in North-East Russia where it is represented by the species Meleagrinella subolifex
Polubotko ( Efimova et al., 1968; Sey and
Polubotko in Damborenea et al., 1992). As this is
earlier than oldest records from eastern Pacific and

European areas, this morphotype does not provide

any clues concerning the opening of the Hispanic
Corridor. Western Pacific records of the other four
morphotypes are of the same ages as first occurrences on opposite sides of the Hispanic Corridor:
Oxytoma cygnipes (Hettangian of New Zealand,
Damborenea and Mancenido, 1992); Entolium
corneolum ( Toarcian of North-East Russia,
Milova, 1988); Entolium lunare (Sinemurian of
Japan, Hayami, 1975); and Chlamys valoniensis
( Hettangian of North-East Russia, Milova, 1976;
Hettangian/Early Sinemurian of New Zealand,
Damborenea and Mancenido, 1992). Because a
dispersal route other than the Hispanic Corridor
cannot be excluded, these morphotypes are not
considered further here.
For the 10 morphotypes that fulfil criteria (1)
and (2), the known Early Jurassic palaeolatitudinal ranges are presented in Table 1. Three morphotypes (Oxytoma inequivalvis, Placunopsis striatula
and Chlamys textoria) were spread over a relatively
broad palaeolatitudinal range during the Early
Jurassic, and may have used a variety of alternative
migration routes. The remaining seven morphotypes, however, seem to be confined to a belt
extending from about 40 north to 45 south of

Table 1
Selected morphotypes of pectinoid bivalves and their Early
Jurassic palaeolatitudinal ranges. N=northern palaeolatitude;
S=southern palaeolatitude. Records from the North-East and
Far East of Russia are not considered as they are commonly
from terranes, the Early Jurassic palaeolatitudinal positions of
which are poorly constrained. For discussion see text. Data
based on Aberhan (1994, 1998a,b), Damborenea (1996),
Damborenea and Mancenido (1979), Johnson (1984),
Trechmann (1923) and unpublished information
Morphotype

Palaeolatitudinal range

Oxytoma inequivalvis
Placunopsis striatula
Propeamussium pumilum
Pseudopecten equivalvis
Camptonectes auritus
Camptonectes subulatus
Chlamys textoria
Eopecten abjectus
Eopecten velatus
Weyla meeki

60N75S
70N42S
35N45S
40N40S
30N35S
40N35S
70N45S
35N38S
35N35S
40N30S

380

M. Aberhan / Palaeogeography, Palaeoclimatology, Palaeoecology 165 (2001) 375394

the palaeoequator. Their restriction to relatively

low palaeolatitudes, simultaneous presence in
Europe and along the eastern Pacific margin, and
absence, or at least stratigraphical restriction to
younger stages, from western Pacific and eastern
Tethys suggest dispersal via the Hispanic Corridor
at various times during the Early Jurassic.

5. Discussion
5.1. Interpretation of faunal similarity patterns
The general trend of pectinoid bivalves is a
progressive increase in similarity between the western Americas and Europe during the Early Jurassic
(Fig. 2). The most marked exceptions are
Sinemurian and Pliensbachian similarity values
between South America and NW-Europe, which
are relatively low in comparison with the
Hettangian value (Fig. 2C ). A diversity analysis
of bivalves from the Andean basins showed that

origination rates of endemics were very high from

Late Sinemurian to Late Pliensbachian times
(Aberhan and Fursich, 1997, 2000). However, a
generally high percentage of Andean endemics
cannot explain the relatively low similarity of
pectinoid morphotypes in mid-Early Jurassic times
between South America and NW-Europe. In fact,
the percentages of pectinoid morphotypes that are
endemic to either NW-Europe or South America
are by no means significantly higher during the
Sinemurian and Pliensbachian than during the
Hettangian and Toarcian (Fig. 3).
This suggests that it is high Hettangian rather
than low Sinemurian and Pliensbachian similarity
that deviates from the normal trend. Examination
of the six Hettangian morphotypes common to
South America and NW-Europe reveals that only
two, Camptonectes subulatus and Eopecten velatus,
are considered as candidates for dispersal through
the Hispanic Corridor (see Section 4.2). Since they
are first recorded from the pre-planorbis beds
and the planorbis Zone respectively, their origins

Fig. 3. Degree of endemism of pectinoid bivalve morphotypes in Early Jurassic time. Data based on Appendix B; absolute age dates
from Gradstein et al. (1994).

M. Aberhan / Palaeogeography, Palaeoclimatology, Palaeoecology 165 (2001) 375394

probably lie in the Triassic (Johnson, 1984). Their

occurrence in South America and Europe may
be the result of dispersal around southern
Gondwanaland or along a circum-Laurasian route
during the Late Triassic hothouse climate system.
For these reasons, relatively high Hettangian Dice
coefficients between South America and European
areas are unlikely to reflect migration via the
Hispanic Corridor in earliest Jurassic time.
Applying a similar approach, Hallam (1983)
has analysed the similarity of Jurassic bivalve
genera between Europe, North America and South
America. Rising Simpson similarity coefficients
between Europe and the western Americas ( Fig. 4)
are in good agreement with similarity curves of
pectinoid morphotypes (Fig. 2). Hallams data
were interpreted to indicate that some degree of
isolation persisted between the Americas and
Europe as at first only a few organisms filtered
through the Hispanic Corridor, and that relatively
free intermigration was established by the Middle
Jurassic (Hallam, 1983; Smith and Tipper, 1986;
Smith, 1988). These conclusions were essentially
confirmed in a recent analysis by Damborenea
(2000), who used the same method as Hallam,
although within a temporally and spatially more
finely resolved framework and with an updated

Fig. 4. Simpson similarity coefficients of bivalve genera between

the western Americas and Europe through time as calculated
by Hallam (1983). Absolute age dates from Gradstein et al.
(1994).

381

data set. By analogy, the pectinoid data presented

herein are compatible with the concept of the
Hispanic Corridor opening progressively during
Early Jurassic time. The major shortcomings of
Hallams approach, however, are that the latitudinal range of taxa as well as data from the western
Pacific and eastern Tethys are not taken into
account. As a consequence, it is impossible to
disentangle the relative importance of high versus
low palaeolatitude migration routes, and to determine when the interchange of organisms began.

5.2. Excluding alternative dispersal routes

Whilst a similarity study with only a few regions
fails to test if and to what degree the Hispanic
Corridor was utilized for migration by organisms,
analysis of distributional patterns holds much
more promise. Because of ample data, establishment of the presence/absence of pectionoid
morphotypes on opposite sides of the Hispanic
Corridor and in the western Pacific was fairly
straightforward. Less information is available from
the eastern Tethys, the pre-Toarcian bivalve record
of which is rather poor or lacking, and from high
palaeolatitude areas.
Data from high northern and southern areas
are considered as particularly important for evaluating high latitude routes for dispersal. Although
the concept of climatic equability in the Jurassic
has been challenged recently (Crowley and North,
1991), there is no doubt that temperature gradients
from the equator to the poles were distinctly lower
compared with the present (e.g. Hallam, 1998 and
references cited therein). Still, the geographic
differentiation of Early Jurassic bivalves into
boreal, austral, bipolar and low latitude forms
(Damborenea, 1993, 1996; Aberhan, 1998b, Liu
et al., 1998), and the well-known TethyanBoreal
provinciality among the ammonites (e.g. Hallam,
1994), suggest that environmental conditions
varied enough to produce latitudinally distinct
faunas. Although documented less intensely, the
absence of a pectinoid morphotype from such
relatively high palaeolatitude areas as the southern
Andes and New Zealand in the southern hemisphere, and northwestern Canada, Arctic Canada

382

M. Aberhan / Palaeogeography, Palaeoclimatology, Palaeoecology 165 (2001) 375394

and East Greenland in the northern hemisphere,

is therefore regarded as adequate evidence to
render unlikely its dispersal around southern
Gondwanaland, northern Laurasia or via the
Viking Corridor.
The palaeolatitudinal range from 40N to 45S,
inferred for those five morphotypes that have
apparently utilized the Hispanic Corridor (see
Sections 4.2 and 5.1, Table 1), seems to be fairly
large to be called low latitude, but is compatible
with palaeoclimatic data. In the Early Jurassic
terrestrial realm, phytogeographic data suggest
that the transition from warm to cool-temperate
conditions in the northern hemisphere took place
at about 45 (Ziegler et al., 1993), and growth
ring studies of fossil wood indicate a broad equatorial zone ranging in latitude from approximately
30N to 30S (Creber and Chaloner, 1984).

With respect to low palaeolatitude routes, an

alternative to the Hispanic Corridor is long-range
larval dispersal across the ancestral Pacific. For
Pangaean times, tropical trans-oceanic faunal dispersal was hypothesized both from east to west
(e.g. Kristan-Tollmann and Tollmann, 1982), and
from west to east (e.g. the pantropical model of
Newton, 1988; see comments by Smith et al.,
1990). For the five pectinoid morphotypes under
discussion, the apparent absence in the western
Pacific and eastern Tethys renders a trans-Pacific
dispersal more unlikely than migration through
the Hispanic Corridor. In addition, it is known
that open oceans act as effective barriers to dispersal of many marine invertebrates, and during
Early Jurassic times the central palaeo-Pacific
ocean was at its widest (Smith and Tipper, 1986;
Smith et al., 1990).

Fig. 5. Postulated origination and subsequent dispersal of five pectinoid bivalve morphotypes during Early Jurassic time. Geographic
distribution and timing of occurrences suggest that dispersal through the Hispanic Corridor occurred during Pliensbachian and
Toarcian times.

M. Aberhan / Palaeogeography, Palaeoclimatology, Palaeoecology 165 (2001) 375394

5.3. Origin of the Hispanic Corridor

The European and eastern Pacific stratigraphic
ranges of the five morphotypes considered to have
used the Hispanic Corridor indicate that three of
them migrated during Pliensbachian and two
during Toarcian times ( Fig. 5, Table 2). Even if
it cannot be excluded from the available data
that a few morphotypes migrated prior to the
Pliensbachian, there is no unequivocal support for
this hypothesis. This argues for an opening of the
Corridor in the Pliensbachian, and suggests that it
was not a viable dispersal route for pectinoid
bivalves during earliest Jurassic times. The similarity curve (Fig. 2A) is consistent with a
Pliensbachian origin of the Hispanic Corridor, but
of course does not prove it (see discussion above).

383

To put these results in perspective, literature

data on time and direction of dispersal of various
fossil groups via the Hispanic Corridor, including
calcareous sponges, corals, bivalves, ammonites,
belemnites and brachiopods, are summarized in
Table 2. These data are consistent with the results
of the present study, and point at an initial opening
in early Pliensbachian time. Only in one case
has an earlier origin been inferred (Stanley and
Beauvais, 1994). Based on the distribution of
corals, these authors conclude that the Hispanic
Corridor may have played an important role
during Sinemurian time. Their data, however,
imply eastwest dispersal of two coral species in
late Hettangian time and westeast dispersal of
two other species later in the early Pliensbachian
(Stanley and Beauvais, 1994, fig. 7). Whilst the

Table 2
List of marine invertebrates bearing on the Early Jurassic origin of the Hispanic Corridor. Het=Hettangian; Plb=Pliensbachian;
Toa=Toarcian; EW=from east to west; WE=from west to east
Taxon

Group

Presumed time
of dispersal

Direction

Reference

Stylothalamia
Phacelostylophyllum rugosum,
Actinastrea plana
Stylophyllopsis victoriae,
Actinastrea minima
Camptonectes auritus,
Pseudopecten equivalvis
Eopecten abjectus,
Propeamussium pumilum
Weyla meeki
Weyla

calcareous sponge
corals

early Plb
late Het

EW
EW

Hillebrandt, 1981
Stanley and Beauvais, 1994

corals

early Plb

WE

Stanley and Beauvais, 1994

bivalve morphotypes

Plb

EW

this study

bivalve morphotypes

Toa

EW

this study

bivalve morphotype
bivalve

Plb
early Plb

WE
WE

Lithiotis (=Plicatostylus)

bivalve

Plb

Opisoma

bivalve

mid Toa

EW

Bouleiceras
Dayiceras/Dubariceras
Frechiella
Leukadiella, Paroniceras

ammonite
ammonites
ammonite
ammonites

early Toa
early Plb
mid Toa
mid Toa

Oregonites
hastitid ancestors of Dicoelitidae

ammonite
belemnites

EW

Thecideaceans

brachiopods

Plb
latest Plb or
earliest Toa
late Plb

this study
Damborenea and Mancenido, 1979, 1988;
Hallam, 1983
Loriga and Neri, 1976; Hillebrandt, 1981;
Hallam, 1983; Nauss and Smith, 1988
Hillebrandt, 1981; Hallam, 1983;
Aberhan and Hillebrandt, 1999
Hillebrandt, 1973
Smith, 1983; Smith and Tipper, 1986
Hillebrandt, 1973
Hillebrandt and Schmidt-Effing, 1981;
Jakobs, 1995
Smith and Tipper, 1986
Jeletzky, 1980

Squamiplana (Cuersithyris)

brachiopod

early Plb

EW

Mancenido and Damborenea, 1990;

Baker and Mancenido, 1997
Mancenido and Dagys, 1992

384

M. Aberhan / Palaeogeography, Palaeoclimatology, Palaeoecology 165 (2001) 375394

latter event is consistent with all the other data

presented in Table 2, a late Hettangian age seems
surprisingly old. Even more extreme is the hypothesis of a corridor being in existence possibly as
early as the Late Triassic, which was based on the
palaeobiogeography of brachiopods and corals
(Sandy and Stanley, 1993; Stanley, 1994 and references cited therein). Fluctuations in eustatic sealevel and tectonic activity along the corridor are
regarded as the main control of its effectiveness.
During Hettangian time, Jurassic eustatic sea-level
was at its lowest stand, and Late Triassic sea-level
was characterized by a major regression (e.g.
Hallam, 1992). This is in accord with the Late
Triassic to earliest Jurassic stratigraphic record of
the numerous rift basins along the postulated
corridor, which consists of continental sequences
comprising lacustrine and fluvial clastic rocks,
evaporites, coals and basalts (Olsen, 1997). The
salt deposits apparently formed in evaporitic lakes
and continental playa or sabka environments fed
by non-marine waters. Marine sediments, including salt deposits of a probably marine source, are
limited to the opposite ends of the rift in Spain,
west of Morocco and in Mexico (Smith and
Tipper, 1986; Stanley, 1994 and references cited
therein). For these reasons, the idea of a preJurassic to earliest Jurassic marine connection
seems to be very speculative, and needs to be
tested rigorously against other models such as
trans-Pacific dispersal.

5.4. Effectiveness of the Hispanic Corridor

Reflecting its effectiveness a migration route
may be classified as a corridor if it allows relatively
free migration, or as a filter if it allows the passage
of some organisms but not others. A very effective
barrier that could only be overcome by chance
events is called a sweepstake route (Simpson, 1940;
McKenna, 1973). During Pliensbachian times, 35
pectinoid morphotypes are recognised, 8% of
which apparently passed through the Hispanic
Corridor ( Fig. 6). In the Toarcian (28 morphotypes) this number is slightly less (7%). Due to the
rigorous criteria applied to identify corridor voyagers, these values may be too low. Widespread taxa

Fig. 6. Percentage of pectinoid morphotypes that apparently

utilized the Hispanic Corridor for the first time during the four
stages of the Early Jurassic. Numbers in the diagram indicate
the total number of morphotypes per time interval.

that also extend into high palaeolatitudes and

western Pacific/eastern Tethys areas obviously
could tolerate a broad range of environmental
conditions. As eurytopic taxa they are also the
most likely to spread through a filter, in which
strong fluctuations in environmental parameters
such as temperature and salinity can be expected.
Restriction to low palaeolatitude taxa excludes
these potential travellers, and the number of those
that actually passed through the corridor may be
somewhat higher. Nevertheless, such a low percentage of pectinoid morphotypes travelling along the
Corridor suggests that it acted as a filter during
Pliensbachian and Toarcian times. This view is
supported by the observation that, in contrast to
some shallow water bivalves, Early Jurassic offshore genera, such as Posidonotis and Otapiria,
were unable to pass the Corridor (Damborenea,
2000, but see discussion on the distribution pattern
of Posidonotis in Aberhan and Palfy, 1996).
Studies comparable with the present one, which
determine the percentage of a fauna that migrated
via the Hispanic Corridor, have not been carried
out so far. But again, faunal similarities of bivalves
( Hallam, 1983; Damborenea, 2000; Figs. 2 and 4)
are compatible with the hypothesis of a progressively opening marine connection from Early
to Middle Jurassic times. Both palaeobiogeographic and geological evidence indicate that a
relatively open corridor was in operation, at least
intermittently, in Middle and Late Jurassic times
(e.g. Hallam, 1977, 1983; Westermann, 1977, 1993;
Riccardi, 1991; Boomer and Ballent, 1996).

M. Aberhan / Palaeogeography, Palaeoclimatology, Palaeoecology 165 (2001) 375394

Development from a filter into a corridor is paralleled by an overall rise in Hallams (1988) eustatic
sea-level curve of the Jurassic. Obviously, this
increase in sea-level was not compensated by tectonic activity along the rift, and probably both
processes have reinforced each other.
The ecological diversity of bivalves, characterized by various life habits and feeding modes,
makes this group a potential source of information
on the physical conditions existing along the route.
Pectinoid bivalves are an epifaunal suspensionfeeding group, and individual taxa belong to the
free-lying, byssate and cemented guilds. Whilst
Eopecten abjectus shows a strong preference
for condensed ferruginous oolites, the species
Propeamussium pumilum, Camptonectes auritus,
Pseudopecten equivalvis and Weyla meeki were
remarkably eurytopic with respect to the substrate
(Johnson, 1984; Aberhan, 1998b). P. pumilum was
also able to live under variable conditions of
oxygen tension and turbulence, and probably had
an opportunistic adaptive strategy (Johnson, 1984;
Aberhan, 1993); C. auritus could cope with high
physical stress, oxygen deficiency and various temperatures (Johnson, 1984), and was also reported
from the upper brachyhaline salinity regime, close
to fully marine conditions (e.g. Fursich, 1993);
and P. equivalvis was able to tolerate high levels
of water energy (Johnson, 1984). Such broad
environmental tolerances are additional evidence
that the Corridor was an ecological barrier that
could only be overcome by eurytopic taxa.
Considering the large amounts of evaporites that
have been deposited in the rifts, it seems likely
that extreme salinities or strongly fluctuating salinity values were limiting biotic exchange. This may
explain the low percentage of pectinoids, which
are at best subordinate faunal components in
salinity-controlled Mesozoic environments. On
the other hand, ammonites apparently used the
Corridor from Pliensbachian times onwards
( Table 2), although they could not cope with
palaeosalinities which deviated from normal
marine values. In this respect it would be interesting to analyse the dispersal of other bivalve groups
such as the relatively euryhaline oysters. Anyway,
the ecology of bivalve larvae and, in the case of

385

ammonites, transport of eggs may be more important factors than the ecological requirements of
adults.
5.5. A two-way migration route
Stratigraphic age relationships provide clues to
the direction of migrations. From the five morphotypes considered to have used the Hispanic
Corridor for dispersal, four originated in Europe
( Fig. 5; Table 2). Only Weyla meeki occurred first
in the eastern palaeo-Pacific. This indicates that
among pectinoids eastwest migration prevailed.
In contrast, for late ToarcianAalenian times,
Hallam (1983) concluded that bivalve traffic was
largely one-way, as eastern Pacific taxa moved
eastward to occupy habitat space which had been
vacated during the Early Toarcian mass extinction
of bivalves in Europe. Examination of Table 2,
however, shows that Early Jurassic migration
from east to west was at least as common as in
the opposite direction. This view of a two-way
migration route is corroborated by several bivalve
genera (including oysters, bakevelliids and
trigoniids), which, according to their known
distribution through time, probably migrated
along the Corridor (Damborenea, 2000). Similarly,
Westermann and Riccardi (1985) did not recognise
a preferred direction in the migration of Middle
Jurassic ammonites through the Corridor.

6. Biotic dispersal through the Hispanic Corridor

and Early Jurassic palaeoceanography
On an evolutionary time scale, organisms with
a holopelagic life cycle have the potential to occur
virtually everywhere, provided their ecological
requirements are met. Sessile, benthic organisms
such as pectinoid bivalves depend more strongly
on oceanic currents for their dispersal during the
larval stages. Given the apparently two-way faunal
exchange through the Hispanic Corridor, oceanic
circulation must have been favourable for larval
transport in both directions. In principal, oceanic
circulation is driven by wind stress and by density
variations of seawater. In the following I

386

M. Aberhan / Palaeogeography, Palaeoclimatology, Palaeoecology 165 (2001) 375394

speculate on the possible influence of both factors

on oceanic circulation through the Hispanic
Corridor.

6.1. Wind-driven circulation

Present-day equatorial current systems in the
open oceans are a direct result of the trade winds.
These blow at about 45 to the equator and induce
westward flows in the surface water (the north and
south equatorial currents). An eastward flowing
equatorial countercurrent is driven by the horizontal pressure gradient that is set up when westward flowing surface water is blocked by a
landmass and the sea-surface slopes up to the
west.
Predicted equatorial ocean currents of the Early
Jurassic palaeo-Pacific are similar to those encountered in the modern Pacific (Parrish, 1992). With
the opening of the Hispanic Corridor some
throughflow of the equatorial current may have
introduced warm Tethyan surface water into the
Pacific. However, it is unlikely that during this
embryonic stage of the Atlantic Ocean a circumglobal, westward flowing equatorial current was
fully established.
An Early Jurassic equatorial countercurrent,
flowing eastward through the Hispanic Corridor,
is not a viable hypothesis. In the case of westward
throughflow of the equatorial current a countercurrent would not have existed at all. In contrast,
with a barrier in place across western Tethys, the
equatorial countercurrent may have been strong
(Parrish, 1992). However, at the western end of
the Hispanic Corridor its driving force, the west
east difference in sea level across the palaeo-Pacific,
was no longer effective.
Because of its size and the distribution of land
in both hemispheres, a strong monsoonal circulation should have existed for Pangaea (e.g.
Kutzbach and Gallimore, 1989; Parrish, 1992,
1993). During the northern winter, northeast trade
winds would have generated a westward flowing
current through the Hispanic Corridor. In analogy
to the present-day summer monsoonal circulation
in Asia, the intertropical convergence zone would
have shifted considerably northward during the

northern summer. As a consequence, the southeast

trade winds would have blown across the palaeoequator and, having been diverted to the right by
the Coriolis force, would give rise to a current
flowing from the eastern Pacific to western Tethys.
Current flow through the Corridor in opposite
directions would have happened semi-annually.
Such a reversal of the normal trade wind pattern
across the continent is in agreement with changes
of surface wind vectors obtained in numerical
model simulations of Pangaean climates (see
models of Kutzbach and Gallimore, 1989;
Chandler et al., 1992).

6.2. Density-driven circulation

The density of seawater is controlled by variations in temperature and salinity. The Mesozoic is
generally conceived as a time interval when the
temperature distribution was more even than
today, so that salinity becomes an important factor
in the density of seawater. It is therefore possible
that vertical circulation in the Mesozoic oceans
was driven largely by salinity differences rather
than primarily by temperature as today (e.g. van
Andel, 1994). In the case of the Hispanic Corridor,
it is conceivable that a thermohaline circulation
was established similar to that between the
presentday Mediterranean and Atlantic Ocean.
Evaporation in the Mediterranean Sea in summer
exceeds the influx of freshwater, and high-salinity
surface water is formed. When this surface water
cools in winter it sinks, and flows out at depth
into the Atlantic Ocean, and relatively lowsalinity water flows into the Mediterranean at
the surface.
While the western end of the Hispanic Corridor
was close to the equator, its eastern end was
located in the subtropics at about 25N during the
Pliensbachian (Fig. 1). Considering this difference
in palaeolatitudes and the extension of the
Corridor from one margin of a supercontinent to
the other, it can be expected that palaeoclimatological parameters differed at opposite ends of the
seaway. Information on palaeoclimate and on
the kind of water masses can be obtained from

M. Aberhan / Palaeogeography, Palaeoclimatology, Palaeoecology 165 (2001) 375394

geological data and from conceptual and numerical

climate models. During the Late Triassic and earliest Jurassic, salt deposits in the eastern and central
part of the rift suggest extreme arid conditions,
which contrast with synchronous lake, playa and
coal swamp deposits in basins in the western part
of the rift complex (Hay et al., 1982). The
only Lower Jurassic marine sediments known at
the western end of the Corridor are in the
Huayacocotla trough of eastern Mexico, which
Schmidt-Effing (1980) has interpreted as an aulaocogen associated with the opening of the Gulf of
Mexico. The marine sedimentary fill consists of
shales, siltstones and sandstones of Sinemurian
age, which suggest relatively humid conditions. In
contrast, the Early Jurassic carbonate platform
deposits of western Tethys and the abundant Early
Jurassic evaporites in northern Africa and northeastern North America apparently formed under
more arid conditions.
For Pliensbachian time, conceptual models of
rainfall patterns predict relatively high rainfall in
the equatorial eastern Pacific and moderately low
rainfall in the western Tethys (Parrish et al., 1982;
Parrish, 1992). Numerical modelling of Pangaean
climate during the Early Jurassic yielded very high
evaporation rates in western Tethys, particularly
during the northern winter (Chandler et al., 1992).
In this model, however, the annually averaged
precipitation minus evaporation values are negative at both ends of the Corridor, and it is difficult
to assess their seasonal variability. Nevertheless, it
is conceivable that, during the Early Jurassic,
supersaline surface water formed by evaporation
in the western Tethys and sank into deep basins,
thereby causing a flow of eastern Pacific surface
water through the Hispanic Corridor into western
Tethys. The Corridor was relatively shallow and
the floor of the rift system probably had a marked
relief. Because vertical barriers could hinder the
movement of saline bottom currents, it is unclear
to what extent a bottom return flow of relatively
high salinity carried water from the western Tethys
back to the eastern Pacific.
In conclusion, the establishment of a megamonsoonal circulation for Pangaea (cf. Kutzbach and
Gallimore, 1989; Parrish, 1993) offers a plausible
mechanism for seasonally changing directions of

387

currents flowing through the Hispanic Corridor,

and deserves further testing and modelling. The
densities of water masses in the equatorial eastern
Pacific and subtropical western Tethys, and their
seasonal variability, are insufficiently known at
present, and it is difficult to evaluate how they
affected oceanic currents within the Corridor.
From the available evidence it seems possible that
a density-driven current reinforced a wind-driven,
eastward flowing current that was in operation
during the northern summer.

7. Conclusions
Palaeobiogeographic patterns of pectinoid
bivalve morphotypes provide further evidence that
the Hispanic Corridor was in existence during
Early Jurassic times. In order to determine its time
of formation and subsequent effectiveness, the
current method of calculating similarity coefficients
alone seems to be insufficient. The presence of
common taxa in a few areas at opposite sides of
the Corridor does not, in itself, imply a protoAtlantic marine connection because several alternative dispersal routes cannot be ruled out. A
more rigorous approach is to identify the percentage of taxa per time interval that were (1) present
at opposite sides of the Corridor, (2) simultaneously absent in the western Pacific/eastern
Tethys and (3) confined to relatively low palaeolatitudes. The results suggest that during Hettangian
and Sinemurian times the future site of the
Hispanic Corridor most likely was an effective
barrier to the spread of marine organisms. The
Hispanic Corridor first opened in the
Pliensbachian and acted as a filter until Toarcian
times, providing a restricted level of faunal
exchange between the eastern Pacific and western
Tethys oceans. Apparently, development into a
marine corridor was not before the Middle
Jurassic. Bidirectional faunal exchange through
the Corridor is in agreement with the establishment
of a megamonsoonal circulation, which may have
caused seasonal alternation of flow directions
within the Corridor.

388

M. Aberhan / Palaeogeography, Palaeoclimatology, Palaeoecology 165 (2001) 375394

Acknowledgements
I wish to thank Bill Hay, Axel von Hillebrandt,
Wolfgang Kiessling and Dave Lazarus for critically
reading the manuscript, and Gerd Westermann
and an anonymous reviewer for their constructive
reviews. This study was financially supported by a
grant from the Deutsche Forschungsgemeinschaft
(Ab 109/1-1), which is acknowledged with
gratitude.

Appendix A: Definition of morphotypes of Early

Jurassic pectinoid bivalves occurring in NW-Europe,
the western Tethys and the eastern palaeo-Pacific
margin
axa marked with an asterisk are known from both
ends of the Hispanic Corridor. Note that the
definition of morphotypes of European Jurassic
pectinids and propeamussiids corresponds to the
species concept of Johnson (1984).
Otapiria inaequicostata Geyer: shell sub-orbicular, left valve ornamented with ribs of different
orders of strength.
Otapiria neuquensis Damborenea: shell sub-ovate,
radial ornament similar on both valves.
Otapiria pacifica Covacevich and Escobar: shell
elongated, ornament stronger on right valve than
on left valve.
Otapiria tailleuri Imlay: shell elongated, radial
ornament more or less equally developed on
both valves.
Lupherella boechiformis (Hyatt): includes all
material referable to this genus.
Anningella carixensis (Cox): includes all material
referable to this genus.
Oxytoma calva (Schlonbach): shell smooth or
very weakly ribbed.
1Oxytoma inequivalvis (J. Sowerby): left valve
with moderately high number of well-developed
primary ribs and intercalated secondary ribs.
1Oxytoma cygnipes ( Young and Bird ): left valve
with small number of spinose ribs.
Arctotis? frenguellii Damborenea: includes this
species and Arctotis sp. A in Aberhan (1998a).

Meleagrinella
papyria
(Quenstedt):
shell
orthocline, ribs relatively widely spaced.
1Meleagrinella substriata (Munster): shell prosocline, with numerous strong radial ribs.
Terquemia arietis (Quenstedt): attachment area
of right valve relatively large.
Terquemia pectiniformis ( Eudes-Deslongchamps):
attachment area of right valve relatively small.
1Placunopsis striatula (Oppel ): includes all radially ribbed nominal species of Early Jurassic
Placunopsis.
Propeamussium laeviradiatum ( Waagen): dorsal
margins of right valve extended into horn-like
processes, left valve coarsely ribbed.
1Propeamussium pumilum (de Lamarck): dorsal
margins of right valve extended slightly beyond
hinge line, left valve finely ribbed.
Propeamussium patriciae Poulton: shell large, with
radial ornament present on both valves.
Kolymonectes carlottensis ( Whiteaves): right
valve with relatively broad ribs.
Kolymonectes coloradoensis ( Weaver): right valve
with very fine radial striae.
1Entolium lunare (Roemer): small byssal notch
present in juveniles.
1Entolium corneolum ( Young and Bird ): byssal
notch absent.
Posidonotis semiplicata (Hyatt): includes all material referable to this genus.
Agerchlamys wunschae (Marwick): ornament of
radial riblets and commarginal lamellae, which
give rise to a reticulate pattern.
1Camptonectes auritus (Schlotheim): sub-orbicular disc, fine divaricate striae on all parts of disc.
1Camptonectes subulatus (Munster): sub-orbicular disc, fine divaricate striae restricted to anterior
and posterior margins of disc.
Camptonectes obscurus (J. Sowerby): ornament
consisting of radial striae and commarginal lamellae, striae restricted to within a few centimetres
of the umbo.
Camptonectes (Costicamptonectes): refers to
Camptonectes (Costicamptonectes) sp. A in
Aberhan (1998a).
Canadonectites paucicostatus Aberhan: right and
left valve smooth except for fine radial riblets on
antero-dorsal part of right valve.

M. Aberhan / Palaeogeography, Palaeoclimatology, Palaeoecology 165 (2001) 375394

1Chlamys textoria (Schlotheim): imbricate lamellae present on the plicae.

1Chlamys valoniensis (Defrance): ornament lacking on the plicae.
Chlamys pollux (dOrbigny): presence of long
tubular spines.
1Eopecten abjectus (Phillips): intercalary costae
rapidly gain the same height as original costae,
two median costae greatly enlarged and bearing
tubercles.
Eopecten hartzi (Rosenkrantz): ornament on left
valve differentiated into costae and striae, a few
costae are enlarged.
Eopecten spondyloides ( Roemer): original costae
similar in height, intercalary costae rapidly gain
the same size as original costae.
1Eopecten velatus (Goldfuss): ornament on left
valve differentiated into costae and striae, no
enlarged costae.
Ochotochlamys aequistriata Aberhan: ornament
of radial riblets and commarginal lamellae and
folds equally developed on right and left valves.
Ochotochlamys bureiensis Sey: style of ribbing on
left and right valves different.
Pseudopecten barbatus (J. Sowerby): long spines
present on right valve.
Pseudopecten dentatus (J. de C. Sowerby): disc
flanks high and vertically striated, down-sulcal
tongueing of commarginal striae, modal number
of plicae 17/18.
1Pseudopecten equivalvis (J. Sowerby): disc flanks
low, commarginal striae curvilinear.
Pseudopecten veyrasensis (Dumortier): vertically
striated disc flanks, down-sulcal tongueing of
commarginal striae, modal number of plicae 14.
Radulonectites sosneadoensis ( Weaver): includes
all eastern Pacific forms referable to this genus.
Weyla alata (von Buch): shell concaveconvex to
plano-convex, ribs of right valve with almost flat
crests and concave interspaces, ribs of left valve
narrow and with concave interspaces.
Weyla ayarti (Dubar): right valve small, with
only up to six subtriangular main ribs.
Weyla bodenbenderi (Behrendsen): shell concave
convex, ribs of right and left valve relatively
broad, number of ribs increasing by intercalation
and branching.

389

1Weyla meeki Damborenea: shell plano-convex to

feebly biconvex, ribs and interspaces of right and
left valve triangular in cross-section.
Weyla unca (Philippi): shell clearly biconvex, ribs
and interspaces of right and left valve triangular
in cross-section.
Weyla yukonensis Aberhan: shell biconvex, ribs
of right valve with more or less flat crests and
triangular interspaces, ribs of left valve narrow,
triangular in cross-section and with concave
interspaces.

Appendix B: Stratigraphic and geographic distribution of Early Jurassic pectinioid bivalve

morphotypes
1=NW-Europe; 2=western Tethys; 3=South
America; 4=North American craton; 5=western
North American terranes. [ ]=occurrence
assumed. Data based on Aberhan (1994 and references cited therein), Aberhan (1998a,b and references cited therein), Arkell (1933), Behmel and
Geyer (1966), Benecke (1905), Berini (1957),
Bertuletti (1962), Brauns (1871), Bronn (1836),
Chapuis and Dewalque (1853), Cossmann (1903),
Cox (1928, 1936), Damborenea (1987 and
references cited therein), Damborenea (1996),
Damborenea and Gonzalez-Leon (1998), Dareste
de la Chavanne (1920), Dechaseaux (1936), Dubar
(1948), Fucini (1920), Gardet and Gerard (1946),
Gemmellaro (18721882), Geyer (1973), Goldfuss
(1833, 1835), Hallam (1987), Holder (1978, 1990),
Johnson (1984), Joly (1936), Kuhn (1935, 1936,
1938), de Lamarck (1819), Lanquine (1929),
Lentini (1973), Leymerie (1881), Lissajous (1907
1912), Oppel (1853, 1856), Pedersen (1986),
Phillips (1829, 1871), Quenstedt (18511852,
18561857, 18651866, 18821885), Riccardi et al.
(1990), Rollier (1915), Schafle (1929), Schlonbach
(1863), Staesche (1926), Terquem (1855),
Terquem and Piette (1865), Troedsson (1951),
Vacek (1886), de Verneuil and Collomb (1853),
Zieten (1833) and unpublished information from
the Huayacocotla trough in eastern Mexico and
the Antimonio terrane of NW-Mexico.

390

M. Aberhan / Palaeogeography, Palaeoclimatology, Palaeoecology 165 (2001) 375394

Morphotype
Otapiria inaequicostata
Otapiria neuquensis
Otapiria pacifica
Otapiria tailleuri
Lupherella boechiformis
Anningella carixensis
Oxytoma calva
Oxytoma inequivalvis
Oxytoma cygnipes
Arctotis? frenguellii
Meleagrinella papyria
Meleagrinella substriata
Terquemia arietis
Terquemia pectiniformis
Placunopsis striatula
Propeamussium laeviradiatum
Propeamussium pumilum
Propeamussium patriciae
Kolymonectes carlottensis
Kolymonectes coloradoensis
Entolium lunare
Entolium corneolum
Posidonotis semiplicata
Agerchlamys wunschae
Camptonectes auritus
Camptonectes subulatus
Camptonectes obscurus
Camptonectes (Costicamptonectes)
Canadonectites paucicostatus
Chlamys textoria
Chlamys valoniensis
Chlamys pollux
Eopecten abjectus
Eopecten hartzi
Eopecten spondyloides
Eopecten velatus
Ochotochlamys aequistriata
Ochotochlamys bureiensis
Pseudopecten barbatus
Pseudopecten dentatus
Pseudopecten equivalvis
Pseudopecten veyrasensis
Radulonectites sosneadoensis
Weyla alata
Weyla ayarti
Weyla bodenbenderi
Weyla meeki
Weyla unca
Weyla yukonensis

Hettangian

Sinemurian
3
3
3
4

Pliensbachian

Toarcian

4, 5

1, 2, 3, 4, 5
1, 3
1
1
1, 2

1
1
1, 2, 3, 4, 5
1, 4
1
1, 5
1
1
1, [2], 3, 5

3, 4, 5
1
1, 3

4
1,
3,
4,
4,
1,
1,

1, 2, 3
1, 2, 3
1, 2

5
1, 2, 3, 4, 5
1, 3, 4, 5
1, 2

4, 5

4, 5

1, 2, 3
5

1, 2, [3]
4, 5

1, 2
3, 5

2, 3
4, 5
5
5
2
[2], 3, 4

1, 2
1, 2
1, 2

[1], 2
1, 2
[1], [2]

3, 4, 5

4
4
5

3,
4,
3,
4,

5
5
4, 5
5

1
1, [2], 3, [4], 5
1, 4, 5
4

1, 2, 3, 4, 5
5
3

1, 5
2

1, [2], 3, 5
1
1, 2

1,
1,
1,
4,

2, 3, 4
2
2, 3
5

1,
1,
3,
5
1,
1,
1
5

5
2, 3, 4, 5
5

5
3
1, 2
3, [4], 5
3, 5
3, 5
[1], 2, 3
1, 2, [4]

5
1, 2, 3, 4, 5

1
3, 4
1, 2, 3
4, 5
4
1,
1,
1,
3,
3,
2
3,
2,
3,

2
2, 3
2
4, 5
4, 5
5
5
5

2, 3, 5
2, 4

1, 2, 3, 5

1, 3
1
1, 2, 3
4
1, 2
1, [2]
1, 2, 3

3
3, 5
2
3

M. Aberhan / Palaeogeography, Palaeoclimatology, Palaeoecology 165 (2001) 375394

References
Aberhan, M., 1993. Benthic macroinvertebrate associations on
a carbonate-clastic ramp in segments of the Early Jurassic
back-arc basin of northern Chile (2629S). Rev. Geol.
Chile 20, 105136.
Aberhan, M., 1994. Early Jurassic Bivalvia of northern Chile.
Part I. Subclasses Palaeotaxodonta, Pteriomorphia and
Isofilibranchia. Beringeria 13, 3115.
Aberhan, M., 1998a. Early Jurassic Bivalvia of western Canada.
Part I. Subclasses Palaeotaxodonta, Pteriomorphia and
Isofilibranchia. Beringeria 21, 57150.
Aberhan, M., 1998b. Paleobiogeographic patterns of pectinoid
bivalves and the Early Jurassic tectonic evolution of western
Canadian terranes. Palaios 13, 129148.
Aberhan, M., 1999. Terrane history of the Canadian Cordillera:
estimating amounts of latitudinal displacement and rotation
of Wrangellia and Stikinia. Geol. Mag. 136, 481492.
Aberhan, M., Fursich, F.T., 1997. Diversity analysis of Lower
Jurassic bivalves of the Andean Basin and the PliensbachianToarcian mass extinction. Lethaia 29, 181195.
Aberhan, M., Fursich, F.T., 2000. Mass origination versus mass
extinction: the biological contribution to the PliensbachianToarcian extinction event. J. Geol. Soc. London 157,
5560.
Aberhan, M., Hillebrandt, A.v., 1999. The bivalve Opisoma in
the Lower Jurassic of northern Chile. Profil 16, 149164.
Aberhan, M., Palfy, J., 1996. A low oxygen tolerant East Pacific
flat clam (Posidonotis semiplicata) from the Lower Jurassic
of the Canadian Cordillera. Can. J. Earth Sci. 33, 9931006.
Arkell, W.J., 1933. A monograph of British Corallian Lamellibranchia. Monogr. Palaeontogr. Soc. London 5, 181228.
Baker, P.G., Mancenido, M.O., 1997. The morphology and
shell microstructure of the thecideidine brachiopod Ancorellina ageri from the Lower Jurassic of Argentina. Palaeontology 40, 191200.
Barron, E.J., Harrison, C.G.A., Sloan II, J.L., Hay, W.W.,
1981. Paleogeography, 180 million years ago to present.
Eclog. Geol. Helv. 74, 443470.
Behmel, H., Geyer, O.F., 1966. Beitrage zur Stratigraphie und
Palaontologie des Juras von Ostspanien. III. Stratigraphie
und Fossilfuhrung im Unterjura von Albarracin (Provinz
Teruel ). N. Jb. Geol. Palaont. Abh. 124, 152.
Benecke, E.W., 1905. Die Versteinerungen der Eisenerzformation von Deutsch-Lothringen und Luxemburg. Abh. Geol.
Spez.-Karte von Elsa-Lothringen, N.F. 6, 1598.
Berini, L., 1957. Studi paleontologici sul Lias del Monte
Albenza (Bergamo). Lamellibranchi e gastropodi del Lias
inferiore. Riv. Ital. Paleont. Stratigr. 63, 3164.
Bertuletti, C., 1962. Studi paleontologici sul Lias del Monte
Albenza (Bergamo). I lamellibranchi dell Hettangiano. Riv.
Ital. Paleont. Stratigr. 68, 169192.
Boomer, I., Ballent, S., 1996. EarlyMiddle Jurassic ostracod
migration between the northern and southern hemispheres:
further evidence for a proto AtlanticCentral America connection. Palaeogeogr., Palaeoclimatol., Palaeoecol. 121,
5364.

391

Brauns, D., 1871. Der untere Jura im nordwestlichen Deutschland von der Grenze der Trias bis zu den Amaltheenthonen
mit besonderer Berucksichtigung seiner Molluskenfauna.
Vieweg & Sohn Braunschweig. 493 pp.
Bronn, H.G., 1836. In: Lethaea geognostica. Schweizerbart,
Stuttgart pp. 225480.
Chandler, M.A., Rind, D., Ruedy, R., 1992. Pangaean climate
during the Early Jurassic: GCM simulations and the sedimentary record of paleoclimate. Geol. Soc. Am. Bull. 104,
543559.
Chapuis, F., Dewalque, C., 1853. Description des fossiles des
terraines secondaires de la province de Luxembourg. Mem.
Acad. R. Belge 35 303 pp.
Cossmann, M., 1903. Note sur LInfralias de la Vendee et des
Deux Se`rres. II. Pelecypodes. Bull. Soc. Geol. Fr., Ser. 4
3, 497545.
Cox, L.R., 1928. The Belemnite Marls of Charmouth, a series
in the Lias of the Dorset Coast. VI. Gastropod and Lamellibranch Molluscs. Q. J. Geol. Soc. London 84, 233245.
Cox, L.R., 1936. The Gastropoda and Lamellibranchia of the
Green Ammonite Beds of Dorset. Q. J. Geol. Soc. London
92, 456471.
Creber, G.T., Chaloner, W.G., 1984. Influence of environmental factors on the wood structure of living and fossil trees.
Bot. Rev. 50, 357448.
Crowley, T.J., North, G.R., 1991. Paleoclimatology. Oxford
Monogr. Geol. Geophys. 18, 1349.
Damborenea, S.E., 1987. Early Jurassic Bivalvia of Argentina.
Part 2. Superfamilies Pteriacea, Buchiacea and part of Pectinacea. Palaeontographica (A) 199, 113216.
Damborenea, S.E., 1993. Early Jurassic South American pectinaceans and circum-Pacific palaeobiogeography. Palaeogeogr., Palaeoclimatol., Palaeoecol. 100, 109123.
Damborenea, S.E., 1996. Palaeobiogeography of Early Jurassic
bivalves along the southeastern Pacific margin. In: XIII
Congreso Geologico Argentino and III Congreso de Exploracion de Hidrocarburos Actas 5, 151167.
Damborenea, S.E., 2000. Hispanic Corridor: its evolution and
the biogeography of bivalve molluscs. In: Hall, R.L., Smith,
P.L. (Eds.), Advances in Jurassic Research 2000, GeoRes.
Forum 6, 369380.
Damborenea, S.E., Gonzalaz-Leon, C.M., 1998. Late Triassic
and Early Jurassic bivalves from Sonora, Mexico. Rev. Mex.
Cienc. Geol. 14, 178201.
Damborenea, S.E., Mancenido, M.O., 1979. On the palaeogeographical distribution of the pectinid genus Weyla (Bivalvia,
Lower Jurassic). Palaeogeogr., Palaeoclimatol., Palaeoecol.
27, 85102.
Damborenea, S.E., Mancenido, M.O., 1988. Weyla: Semblanza
de un bivalvo Jurasico Andino. In: V Congreso Geologico
Chileno Tomo 2., C13C15.
Damborenea, S.E., Mancenido, M.O., 1992. A comparison of
Jurassic marine benthonic faunas from South America and
New Zealand. J. R. Soc. N.Z. 22, 131152.
Damborenea, S.E., Polubotko, I.V., Sey, I.I., Paraketsov, K.V.,
1992. Bivalve zones and assemblages of the circum-Pacific
region. In: Westermann, G.E.G. (Ed.), The Jurassic of the

392

M. Aberhan / Palaeogeography, Palaeoclimatology, Palaeoecology 165 (2001) 375394

Circum-Pacific. Cambridge University Press, Cambridge,

pp. 300307.
Dareste de la Chavanne, J., 1920. Fossiles liasiques de la region
de Guelma. Mater. Carte Geol. Alger., Ser. 1 (Paleont.)
5 72 pp.
Dechaseaux, C., 1936. Pectinides jurassiques de lest du Bassin
de Paris. Revision et biogeographie. Ann. Pale`ont. 25,
1148.
de Lamarck, J.B, 1819. Histoire naturelle des animaux sans
vertebres Tome 6. Verdie`re. 343 pp.
de Verneuil, M., Collomb, E., 1853. Coup doeil sur la constitution geologique de quelques provinces de lEspagne. Bull.
Soc. Geol. Fr. Paris, Ser. 2 10, 61147, 162166.
Dubar, G., 1948. Etudes paleontologiques sur le Lias du Maroc.
La faune Domerienne du Djebel Bou-Dahar. Notes Mem.
Serv. Geol. Maroc 68, 1250.
Efimova, A.F., Kinasov, V.P., Paraketsov, K.V., Polubotko,
I.V., Repin, Yu.S., Dagis, A.S., 1968. Field atlas of the Jurassic fauna and flora of the northeastern part of the USSR.
Ministry of Geology RSFSR, Severo-vostochnoye ordena
trydovogo krasnogo Znameni geologicheskoe upravlenie,
Magadanskoe Knizhnoe-Izdatelstvo, 378 pp. (in Russian).
Fucini, A., 1920. Fossili domeriani dei dintorni di Taormina.
Parte I. Palaeontogr. Ital. 26, 75116.
Fursich, F.T., 1993. Palaeoecology and evolution of Mesozoic
salinity-controlled benthic macroinvertebrate associations.
Lethaia 26, 327346.
Gardet, G., Gerard, C., 1946. Contribution a letude paleontologique du Moyen-Atlas septentrional. Notes Mem. Serv.
Geol. Maroc 64, 188.
Gemmellaro, G.G., 18721882. Sopra alcune faune giuresi e
liasiche della Sicilia. Studi paleontologici. Lao. Palermo,
434 pp.
Geyer, O.F., 1973. Das prakretazische Mesozoikum von
Kolumbien. Geol. Jb. B 5 155 pp.
Goldfuss, G.A., 1833. In: Petrefacta Germaniae. II (1). Arnz,
Dusseldorf, pp. 168.
Goldfuss, G.A., 1835. In: Petrefacta Germaniae. II (2). Arnz,
Dusseldorf, pp. 69140.
Gradstein, F.M., Agterberg, F.P., Ogg, J.G., Hardenbol, J., van
Veen, P., Thierry, J., Huang, Z., 1994. A Mesozoic time
scale. J. Geophys. Res. 99, 24,05124,074.
Hallam, A., 1977. Jurassic bivalve biogeography. Paleobiology
3, 5873.
Hallam, A., 1983. Early and mid-Jurassic molluscan biogeography and the establishment of the central Atlantic seaway.
Palaeogeogr., Palaeoclimatol., Palaeoecol. 43, 181193.
Hallam, A., 1987. Radiations and extinctions in relation to environmental change in the marine Lower Jurassic of northwest
Europe. Paleobiology 13, 152168.
Hallam, A., 1988. A reevaluation of Jurassic eustasy in the light
of new data and the revised Exxon curve. In: Wilgus, C.K.,
Hastings, B.S.Posamentier, H., Wagoner, J.V., Ross, C.A.,
Kendall, C.G.S.C. (Eds.), Sea-level Changes An Integrated Approach, SEPM Spec. Publ. 42, 261273.
Hallam, A., 1992. Phanerozoic Sea-Level Changes. Columbia
University Press, Columbia.

Hallam, A., 1994. An outline of phanerozoic biogeography.

Oxford Biogeogr. Ser. 10, 1246.
Hallam, A., 1998. The determination of Jurassic environments
using palaeoecological methods. Bull. Soc. Geol. Fr. 169,
681687.
Hay, W.W., Behensky, J.F., Barron, E.J., Sloan II, J.L., 1982.
Late TriassicLiassic paleoclimatology of the proto-central
North Atlantic rift system. Palaeogeogr., Palaeoclimatol.,
Palaeoecol. 40, 1330.
Hayami, I., 1975. A systematic survey of the Mesozoic Bivalvia
from Japan. Univ. Mus., Univ. Tokyo, Bull. 10, 1249.
Hillebrandt, A.v., 1973. Die Ammonitengattungen Bouleiceras
und Frechiella im Jura von Chile und Argentinien. Eclog.
Geol. Helv. 66, 351363.
Hillebrandt, A.v., 1981. Kontinentalverschiebung und die
palaozoogeographischen Beziehungen des sudamerikanischen Lias. Geol. Rdsch. 70, 570582.
Hillebrandt, A.v., Schmidt-Effing, R., 1981. Ammoniten aus
dem Toarcium (Jura) von Chile (Sudamerika). Zitteliana
6, 174.
ber die Pectiniden-Gattung Parvamussium
Holder, H., 1978. U
im Jura. Stutt. Beitr. Naturk., Ser. B 38 37 pp.
ber die Muschelgattung Placunopsis (PectiHolder, H., 1990. U
nacea, Placunopsidae) in Trias und Jura. Stutt. Beitr.
Naturk., Ser. B 165 63 pp.
Jakobs, G.K., 1995. New occurrences of Leukadiella and Paroniceras (Ammonoidea) from the Toarcian (Lower Jurassic)
of the Canadian Cordillera. J. Paleont. 69, 8998.
Jeletzky, J.A., 1980. Dicoelitid belemnites from the Toarcian
Middle Bajocian of western and Arctic Canada. Geol. Surv.
Can. Bull. 338, 171.
Johnson, A.L.A., 1984. The palaeobiology of the bivalve families Pectinidae and Propeamussiidae in the Jurassic of
Europe. Zitteliana 11, 1235.
Joly, H., 1936. Les fossiles du Jurassique de la Belgique avec
description stratigraphique de chaque etage. Deuxie`me
partie. Lias inferieur. Mem. Mus. R. Hist. Nat. Belg. 79,
1244.
Kristan-Tollmann, E., Tollmann, A., 1982. Die Entwicklung
der Tethystrias und Herkunft ihrer Fauna. Geol. Rdsch.
71, 9871019.
Kuhn, O., 1935. Weitere Beitrage zur Fauna des untersten Lias
in Schwaben und Franken. Jb. Ver. Vaterland. Naturk.
Wurtt. 91, 218.
Kuhn, O., 1936. Die Fauna des Amaltheentons (Lias ) in Franken. N. Jb. Min. Geol. Palaont. B 75, 231311.
Kuhn, O., 1938. Die Fauna des Dogger der Frankenalb (Mit
Nachtragen zum ubrigen Jura). Nova Acta Leopold. N.F.
6, 125170.
Kutzbach, J.E., Gallimore, R.G., 1989. Pangaean climates:
megamonsoons of the megacontinent. J. Geophys. Res. 94,
33413357.
Lanquine, A., 1929. Le Lias et le Jurassique des chaines
provencales. I. Le Lias et le Jurassique Inferieur. Bull. Serv.
Carte Geol. Fr. 32, 41425.
Lentini, F., 1973. I molluschi del Lias inferiore di Lorigi (Sicilia
nordorientale). Boll. Soc. Paleont. Ital. 12, 2375.

M. Aberhan / Palaeogeography, Palaeoclimatology, Palaeoecology 165 (2001) 375394

Leymerie, A., 1881. Description geologique et paleontologique
des Pyrenees de la Haute-Garonne. Edouard Privat.
Toulouse 1010 pp.
Lissajous, M., 19071912. Jurassique Maconnais. Description
des fossiles caracteristiques et des espe`ces les plus communes. Bull. Soc. Hist. Nat. Macon 3 496 pp.
Liu, C., Heinze, M., Fursich, F.T., 1998. Bivalve provinces
in the Proto-Atlantic and along the southern margin of
the Tethys in the Jurassic. Palaeogeogr., Palaeoclimatol.,
Palaeoecol. 137, 127151.
Loriga, C., Neri, C., 1976. Aspetti paleobiologici e paleogeografici della facies a Lithiotis (Giurese Inf.). Riv. Ital.
Paleont. Stratigr. 82, 651706.
Mancenido, M.O., Dagys, A.S., 1992. Brachiopods of the
circum-Pacific region. In: Westermann, G.E.G. (Ed.), The
Jurassic of the Circum-Pacific. Cambridge University Press,
Cambridge, pp. 328333.
Mancenido, M.O., Damborenea, S.E., 1990. Corallophilous
micromorphic brachiopods from the Lower Jurassic of west
central Argentina. In: MacKinnon, D.I., Lee, D.E., Campbell, J.D. (Eds.), Brachiopods through Time. Balkema, Rotterdam, pp. 8996.
McKenna, M.C., 1973. Sweepstakes, filters, corridors, Noahs
arks, and beached Viking funeral ships in palaeogeography.
In: Tarling, D.H., Runcorn, S.K. ( Eds.), Implications of
Continental Drift to the Earth Sciences. Academic Press,
New York, pp. 295308.
Milova, L.V., 1976. Stratigraphy and bivalve molluscs of the
TriassicJurassic deposits of northern Priokhotya. Akademiya Nauk SSSR, Dalnevostochnyy Nauchnyy Tsentr,
Trudy Severo-Vostochnyy Kompleksnyy Nauchno-Issledovatel-skiy Instituta 65, pp. 3110 (in Russian).
Milova, L.V., 1985. New Pliensbachian bivalve molluscs of
North Priokhotya. In: Bivalve and Cephalopod Mollusks
of the North-East of the USSR. Akademiya Nauk SSSR,
pp. 4256. in Russian.
Milova, L.V., 1988. Early Jurassic bivalve molluscs of northeastern USSR. Akademiya Nauk SSSR, Dalnevostochnyy
Otdeleniye, Trudy Severo-Vostochnyy Kompleksnyy
Nauchno-Issledovatel-skiy Instituta, 152 pp. (in Russian).
Nauss, A.L., Smith, P.L., 1988. Lithiotis (Bivalvia) bioherms in
the Lower Jurassic of east-central Oregon, USA. Palaeogeogr., Palaeoclimatol., Palaeoecol. 65, 253268.
Newton, C.R., 1988. Significance of Tethyan fossils in the
American Cordillera. Science 242, 385391.
Olsen, P.E., 1997. Stratigraphic record of the Early Mesozoic
breakup of Pangea in the LaurasiaGondwana rift system.
Annu. Rev. Earth Planet. Sci. 25, 337401.
Oppel, A., 1853. Der mittlere Lias Schwabens. Jh. Ver. Vaterland. Naturk. Wurtt. 10, 39136.
Oppel, A., 1856. Die Juraformation Englands, Frankreichs und
des sudwestlichen Deutschlands. Wurtt. Naturwiss. Jh. 12,
1438.
Palmer, C.P., 1989. Larval shells of four Jurassic bivalve molluscs. Bull. Br. Mus. Nat. Hist. (Geol.) 45, 5769.
Parrish, J.T., 1992. Jurassic climate and oceanography of the
Pacific region. In: Westermann, G.E.G. ( Ed.), The Jurassic

393

of the Circum-Pacific. Cambridge University Press, Cambridge, pp. 365379.

Parrish, J.T., 1993. Climate of the supercontinent Pangea.
J. Geol. 101, 215233.
Parrish, J.T., Ziegler, A.M., Scotese, C.R., 1982. Rainfall patterns and the distribution of coals and evaporites in the
Mesozoic and Cenozoic. Palaeogeogr., Palaeoclimatol.,
Palaeoecol. 40, 67101.
Pedersen, G.K., 1986. Changes in the bivalve assemblage of an
Early Jurassic mudstone sequence (the Fjerritslev Formation in the Gassum 1 Well, Denmark). Palaeogeogr., Palaeoclimat., Palaeoecol. 53, 139160.
Phillips, J., 1829. Illustrations of the Geology of Yorkshire.
Wilson and Sons, York. 199 pp.
Phillips, J., 1871. Geology of Oxford and the Valley of the
Thames. Clarendon Press, Oxford. 523 pp.
Polubotko, I.V., Repin, Yu.S., 1988. Lower and Middle Jurassic
of the North-East. In: Westermann, G.E.G., Riccardi, A.C.
( Eds.), Jurassic Taxa Ranges and Correlation Charts for
the Circum Pacific. 1. Soviet Union, Newslett. Stratigr. 19,
117.
Quenstedt, F.A., 18511852. Handbuch der Petrefaktenkunde.
first ed., Laupp and Siebeck. Tubingen 792 pp.
Quenstedt, F.A., 18561857. Der Jura. Tubingen Laupp.
842 pp.
Quenstedt, F.A., 18651866. Handbuch der Petrefaktenkunde.
second ed., Laupp and Siebeck. Tubingen 982 pp.
Quenstedt, F.A., 18821885. Handbuch der Petrefaktenkunde.
third ed., Laupp. Tubingen 1239 pp.
Riccardi, A.C., 1991. Jurassic and Cretaceous marine connections between the Southeast Pacific and Tethys. Palaeogeogr., Palaeoclimatol., Palaeoecol. 87, 155189.
Riccardi, A.C., Damborenea, S.E., Mancenido, M.O., 1990.
Lower Jurassic of South America and Antarctic Peninsula.
Newslett. Stratigr. 21, 75103.
Rollier, L., 1915. Fossiles nouveaux ou peu connus des terrains
secondaires (mesozoiques) du Jura et des contrees environnantes. Mem. Soc. Paleont. Suisse 41, 345500.
Sandy, M.R., Stanley Jr., G.D., 1993. Late Triassic brachiopods
from the Luning Formation, Nevada, and their palaeobiogeographical significance. Palaeontology 36, 439480.
ber Lias- und Doggeraustern. Geol. Palaont.
Schafle, L., 1929. U
Abh. N.F. 17, 63150.
Schlonbach, U., 1863. Ueber den Eisenstein des mittleren Lias
im nord-westlichen Deutschland, mit Berucksichtigung der
alteren und jungeren Lias-Schichten. Zeitschr. dt. Geol. Ges.
15, 465566.
Schmidt-Effing, R., 1980. The Huayacocotla aulacogen in
Mexico (Lower Jurassic) and the origin of the Gulf
of Mexico. In: Pilger, R.H. ( Ed.), The Origin of the Gulf of
Mexico and the Early Opening of the Central North Atlantic
Ocean. Louisiana State University, Baton Rouge, pp. 7986.
Sey, I.I., 1984. The Late Pliensbachian bivalves of Bureya
Trough (Far East of Russia). In: Poyarkova, Z.N., Zonovalov, V.P. ( Eds.), New Data on the Detailed Biostratigraphy
of the Phanerozoic of the Far East. Akademiya Nauk SSSR,
pp. 8697. in Russian.

394

M. Aberhan / Palaeogeography, Palaeoclimatology, Palaeoecology 165 (2001) 375394

Simpson, G.G., 1940. Mammals and land bridges. J. Washington Acad. Sci. 30, 137163.
Smith, P.L., 1983. The Pliensbachian ammonite Dayiceras dayiceroides and Early Jurassic paleogeography. Can. J. Earth
Sci. 20, 8691.
Smith, P.L., 1988. Paleobiogeography and plate tectonics.
Geosci. Can. 15, 261279.
Smith, P.L., Tipper, H.W., 1986. Plate tectonics and paleobiogeography: Early Jurassic (Pliensbachian) endemism and
diversity. Palaios 1, 399412.
Smith, P.L., Westermann, G.E.G., Stanley, G.D., Yancey, T.E.,
1990. Paleobiogeography of the ancient Pacific. Science
249, 680681.
Staesche, K.v., 1926. Die Pectiniden des Schwabischen Jura.
Geol. Palaont. Abh. N.S. 15 136 pp.
Stanley Jr., G.D., 1994. Late Paleozoic and early Mesozoic reefbuilding organisms and paleogeography: the Tethyan
North American connection. Cour. Forsch.-Inst. Senckenberg 172, 6975.
Stanley Jr., G.D., Beauvais, L., 1994. Corals from an Early
Jurassic coral reef in British Columbia refuge on an oceanic
island reef. Lethaia 27, 3547.
Terquem, O., 1855. Paleontologie de letage inferieur de la formation liasique de la province de Luxembourg, GrandDuche (Hollande) et de Hettange, du Departement de la
Moselle. Mem. Soc. Geol. Fr., Ser. 2 5, 219343.
Terquem, O., Piette, E., 1865. Le Lias inferieur de lest de la
France comprenant la Meurthe, la Moselle, le Grand-Duche
de Luxembourg, la Belgique et la Meuse. Mem. Soc. Geol.
Fr., Ser. 2 8, 1175.
Trechmann, C.T., 1923. The Jurassic rocks of New Zealand. Q.
J. Geol. Soc. London 79, 246312.
Troedsson, G., 1951. On the Hoganas Series of Sweden

(Rhaeto-Lias). Lunds Univ. Arsskrift, N.F. Avd. 2 (47),

1268.
ber die Fauna der Oolithe von Cap. S. VigiVacek, M., 1886. U
lio verbunden mit einer Studie uber die obere Liasgrenze.
Abh. k. k. Geol. Reichsanst. 12, 1212.
van Andel, T.H., 1994. New Views on an Old Planet. Cambridge Univerity Press, Cambridge. 439 pp.
Waller, T.R., 1991. Evolutionary relationships among commercial scallops (Mollusca: Bivalvia: Pectinidae). In: Shumway,
S.E. ( Ed.), Scallops: Biology, Ecology and Aquaculture.
Elsevier, Amsterdam, pp. 173.
Waller, T.R., 1993. The evolution of Chlamys (Mollusca:
Bivalvia: Pectinidae) in the tropical western Atlantic and
eastern Pacific. Am. Malacol. Bull. 10, 195249.
Westermann, G.E.G., 1977. Comments to Hallams conclusion
regarding the first marine connection between the eastern
Pacific and western Tethys. In: West, R.M. (Ed.), Paleontology and Plate Tectonics, Milwaukee Publ. Mus. Spec. Publ.
Biol. Geol. 2, 3538.
Westermann, G.E.G., 1993. Global bio-events in mid-Jurassic
ammonites controlled by seaways. In: House, M.R. ( Ed.),
The Ammonoidea: Environment, Ecology and Evolutionary
Change, System. Assoc. Spec. Vol. 47, 187226.
Westermann, G.E.G., Riccardi, A., 1985. Middle Jurassic
ammonite evolution in the Andean Province and emigration
to Tethys. In: Bayer, U., Seilacher, A. ( Eds.), Sedimentary
and Evolutionary Cycles. Springer, Berlin, pp. 434.
Ziegler, A.M., Parrish, J.M., Yao, J., Gyllenhaal, E.D., Rowley,
D.B., Parrish, J.T., Nie, S., Bekker, A., Hulver, M.L., 1993.
Early Mesozoic phytogeography and climate. Philos. Trans.
R. Soc. London, Ser. B: 341, 297305.
Zieten, C.H.v., 1833. In: Die Versteinerungen Wurttembergs.
Schweizerbart Stuttgant, pp. 65102.