Cretaceous Research 54 (2015) 243e254

Contents lists available at ScienceDirect

journal homep age: ww w.e ls ev ie r. co m / l oc at e /C ret Re s

AptianeAlbian rudist bivalves (Hippuritida) from the Chilean Central

Andes: Their palaeoceanographic signicance
Jean-Pierre Masse
a
b

a, *

, Francisco-Amaro Mourgues , Mukerrem Fenerci-Masse

CEREGE, Aix-Marseille University, Centre Saint-Charles, 13331 Marseille Cedex 03, France
Terra Ignota, Heritage and Geosciences Consulting, Dr. Ca diz 726, N~ un~oa, Santiago, Chile

a r t i c l e i n f o
Article history:
Received 30 September 2014
Accepted in revised form 7 December 2014
Available online

Keywords:
Rudist bivalves
Early Cretaceous
Taxonomy
Palaeobiogeography
South America

a b s t r a c t
The study of AptianeAlbian rudist faunas from the Chilean Central Andes documents the presence of
two forms, ascribed to the Monopleuridae: Douvillelia skeltoni an early Aptian species known
hitherto from Mexico, and Jerjesia chilensis, an endemic Albian species from Chile and Peru, with a
complex taxonomic history. The regional stratigraphy of the Central Andes, combining ammonites and
rudists, is consistent with the Caribbean stratigraphic distribution of Jerjesia and Douvillelia.
Andean occurrences of the two genera broaden their distributional area on the Pacic side of Americas,
and testify their biostratigraphic value. Oceanographic conditions of the Chilean Pacic margin
during the AptianeAlbian, including moderate but effective cold oceanic current, upwellings, high
productivity and thermal instability, may acknowledge the low taxonomic diversity of rudist
assemblages, which look impoverished when compared to their low latitude homologues from the
Caribbean regions which were functioning as a dispersal centre.
2015 Published by Elsevier Ltd.

1. Introduction
Cretaceous, rudist-bearing carbonate platforms, tend to be
globally restricted to 30oe35o south and north latitudes, so they
extend farther beyond the tropics than their modern
counterparts,
i.e. coral reefs (Coates, 1973; Masse, 1992a,b; Simo et al., 1993;
Philip, 1998; Skelton et al., 2003). However Eastern Asia, i.e. the
Japanese archipelago, and the Pacic side of North America,
where rudist occurrences are found at higher palaeolatitudes, do
not t this overall latitudinal distribution (Masse, 1992a,b; Sano,
2012; Sano et al., 2013; Skelton et al., 2013). Rudist occurrences
of Japan at a palaeolatitude in the range of 35oe45o N are
somewhat anomalic. Nevertheless some debate exists on the
palaeoposition of the different parts of the archipelago deduced
from palinspastic re- constructions, for instance Hokkaido is close
to its present day location (44oN) for Hay and De Conto (1999), at
50oN for Eldridge et al. (2000), and 35e36oN for Takashima et
al. (2007). Whatever its amplitude, this latitudinal anomaly
hardly corresponds to a late AptianeAlbian p.p. warm peak, as
suggested by Takashima et al. (2007),
because
this
event
cannot explain the presence of

* Corresponding author.
E-mail address: masse@cerege.fr (J.-P. Masse).

TithonianeBerriasian forms recorded from the same region (Sano

and Skelton, 2010). We favour the warming effect of a Cretaceous
precursor of the modern Kuroshio current (Japan current) is supported by simulations of Aptian oceanic circulation (e.g. Fluteau
et al., 2007). This present day warm (24oc annual average SST)
north owing current, sustaining coral reefs in southern Japan,
corresponds to the western branch of the North Pacic, clockwise
ocean gyre. Similarly, rudist occurrences of North California
(Sano et al., 2013; Skelton et al., 2013) with palaeolatitudes in the
range of 40oN, do not t the above mentioned latitudinal range and
their palaeoceanographic and/or palaeogeographic context is still
unclear.
The southernmost rudist (27oe30o S) from America:
Hippurites chilensis, was known from Chile since the rst half
of XIX century after the pioneer works of d'Orbigny (1842) about
the fossils and geology of South America. Subsequently this
species was either ignored or misinterpreted.
The objectives of the present paper are to revise the systematic
position and to analyse the taxonomic avatars of Hippurites
chi- lensis, the rst rudist to have been described in America.
It com- plements and supports the results briey exposed earlier
(Mourgues et al., 2010) placing this form in the genus Jerjesia.
In addition we provide data on some other forms, including the
genus Dou- villelia, and discuss the biostratigraphic and
palaeobiogeographic

http://dx.doi.org/10.1016/j.cretres.2014.12.00
4 0195-6671/ 2015 Published by Elsevier
Ltd.

].-P. Masse et al. / Cretaceous Research 54 (2015) 243-254

signicance of the rudist assemblages, focussing on comparisons

with Caribbean faunas. This study is based on the analysis of
historic collections and specimens collected by one of us
(F.A.M.), their dating is founded on
an up-to-date
biostratigraphy of the Chan~ arcillo Basin (Mourgues, 2007). The specimens are housed in
the
Paleontological Collection of the Geological and Mining Survey of
Chile, Santiago (SNGM).
A key problem we will address is that South America rudist
occurrences which match the overall latitudinal distribution of the
group, are somewhat anomalic if we consider the sea surface
oceanic currents. By reference to the modern regional oceanography characterized by the existence of the Peru (or Humbolt)
current, a cold current owing northward off western South
America, and forming the eastern branch of the South Pacic gyre,
is assumed to have existed during the Early Cretaceous (Hay, 1995).
2. Regional geological setting
Rudist bearing beds are present in the Pabello n and
Arqueros Formations of the Chan~ arcillo basin, one of the retroarc
Andean ba- sins lining South America. The inll of the
Chan~ arcillo Basin
(Atacama basin) (Fig. 1), comprises more than 2000 m of marine
carbonate rocks and minor volcaniclastics at the base and top of the
succession, known as the Chan~ arcillo Group in the sense
of
Segerstrom and Ruiz (1962), which ranges in age from late
Berriasian to early Albian (Mourgues, 2007). This Lower Cretaceous
succession crops out in the southern Atacama region as a NNEtrending belt

from La Serena to Copiapo , parallel to the current plate

margin (Fig. 2). Southward of this belt, there are only isolated
outcrops. During late Aptianeearly Albian times the Pacic
Gondwana active margin was characterized by a geodynamic
setting with a closing back arc basin which suffered a strong
tectonic and volcanic activity (Are valo et al., 2005). This activity
was reected in the record of synsedimentary tectonic features,
volcaniclastics ows and lava events into the regressive platform
carbonate sediments of the upper Pabello
n Formation
(Mourgues, 2007), that is the uppermost lithostratigraphic unit of the Chan~ arcillo Group. This group is
covered
disconformably eastwards by the Cerrillos Formation (Segerstrom
and Parker, 1959), a thick subaerial volcaniclastic sequence ascribed
to the early Albianeearly Maastrichtian (Maksaev et al., 2009).
3. Biostratigraphy of rudist bearing formations
The Pabellon Formation was originally (Tavera, 1956;
Corvalan, 1974) assumed to be upper Barremian, based on the
presence of Agria blumenbachi (formerly Hippurites
chilensis Jerjesia chi- lensis). The publication of an
ammonite
fragment
attributed
to
Parahoplites
gr.
nuteldiensis (J. Sowerby), subsequently raised the age of
formation to the early late Aptian (Pe rez et al., 1990). The
recent revision of ammonite specimens collected by Pe rez et
al. (1990) show the presence of Neodeshayesites, this result
combined with the discovery of Hypacanthoplites-bearing
beds at the Que- brada El Molle, lead to assign the top of the
Pabello n Formation to the lower Albian (Mourgues, 2007).

Fig. 1. Location map and stratigraphic succession of the Chan~arcillo Group (modied from Mourgues, 2007; numerical ages from Cohen et al., 2013).

Chaarci
lloBasi
n

/&

26
Kin

_/
,/

,/

,/

Quebrada
Carrizalillo
Que"brada El Molle

(.)

\,

..

,e,

}'

/.

('
,

l
,

-.J

:l:

I
1

(.,)

/_J
\

..,

.l'
(

'

1
\..
'

LA S ER
ENA

50 km

Marine carbonated
sedimentary formations
Volcan ic with
intercalated

)'

Miocene Pliocene alluvial deposits

(MPga)

Cerrillos Formation (Kic)

l/1J1J Pabelln Formation

(Kip)

/.
(!)

Quebrada Marquesa Formation (Kic)

Arqueros Formation

BSTI Member 5 (Kia5): volcanic

Member 4 (Kia4): carbonate platform

marine sedimentary

E'l

Porphyry andesite (KipA )

formations
platform Fossil locality

D
D

Totoralillo Formation (Kit)

Nantoco Formation (Kin)

Q'

Member 3 (Kia3): volcanic

Member 2 (Kia2): carbonate

Member 1 (Kia1): volcanic

Fig. 2. Location map and geological context of the study stratigraphic sections.

2
4

J.-P. Masse et al. / Cretaceous Research 54

(2015) 243e254

At Quebrada Carrizalillo (Fig. 3A) Jerjesia biostromes are part

of a transgressive succession that unconformably overlies
porphyry andesites. At Quebrada El Molle (Fig. 3B) Jerjesia e
bearing beds are observed as biostrome block components in a
calcareous breccia olistostrome. This conspicuous unit is capped
by clastic e carbonate marine sequences containing a mideearly
Albian Neodeshayesites horizon and is supported by
Hypacanthoplites bearing marine de- posits of latest Aptian age
(Aguirre-Urreta et al., 2007). The age for Jerjesia chilensis beds
is therefore early Albian.
The type locality of Jerjesia chilensis correspond to the
Arqueros ancient silver mine (29o490 S). In this locality the Lower
Cretaceous is represented by a volcanic succession with two well
dened marine carbonate intercalations (Fig. 3C). Aguirre and
Egert (1962) dened de Arqueros Formation as a volcanosedimentary succes- sion made of ve members, and
recognized precisely the strati- graphic situation of rudists in the
upper member of the formation. Recently detailed stratigraphic
sections were performed at the type locality and two rudists
biostromes were identied in the middle and top parts of the
Member 4 (Kia4) of the Arqueros Formation. The rst member of
the Arqueros Formation (Kia1) is composed by a thick porphyry
andesite lava ow with locally pillow structures. The overlying
unit is a massive bed of shoreface limestone and marlyesherty
(silty) hemipelagic limestone with gastropods, spa- tangoid
echinids and pectinids, with interbedded thin grainstones
containing trigoniids and other bivalves (Kia2). A mid homogeneous volcanic member (Kia3) is formed by porphyry andesite and
is covered unconformably by a new marine succession (Kia4) made
of a thinly rhythmic bedded mudstone and coarse sandstone, with
two massive rudist biostromes having a muddy matrix. At the
upper part of the section a calcareous breccia is present. This carbonate interval is covered by a thin manganese-rich volcaniclastic
red sandstone which passes upward to a thick andesitic lava succession (Kia5).
Douvillelia beds are present in the uppermost marine strata
of the Pabellon Formation at Cerro Pajonales (Fig. 3D).
Stratigraphic indices, i.e. ammonites, were identied below and
mark the upper Barremian Antarcticoceras domeykanus zone
and the lower Barre- mian Emericiceras and Shasticrioceras
zones (Mourgues, 2007). Douvillelia beds are observed just
below the regional unconformity underlying the Cerrillos
Formation. In this locality the upper part of the Pabello n
Formation appears to be missing by erosion, this interpretation is
supported by the absence of porphyry andesite (see above) or
any volcanic episode recognized regionally, e.g. Quebrada
Carrizalillo, Quebrada El Molle and Llano de Arqueros, dated as
late Aptian, i.e.-115 Ma (Morata et al., 2001; Morata and Aguirre,
2003). The foregoing suggests that the Douvillelia- bearing
beds are early Aptian in age. The identication of Douvillelia
skeltoni Alencaster and Pantoja-Alor corroborates this dating
(see discussion below).
4. Systematic palaeontology
The classication used herein refers to Skelton (2013a,b).
Abbreviations used for systematic descriptions. LV, left
valve; RV, right valve; Ant, anterior side; Post, posterior side;
Vent, ventral; Dors, dorsal; t, tooth; at, anterior tooth; pt,
posterior tooth; ct, central tooth; am, anterior myophore; pm,
posterior myophore; lg, ligament groove; bc, body cavity; ol,
outer shell layer; il, inner shell layer; AB, anterior band; PB,
posterior band; IB, interband; Dap, antero-posterior diameter;
Ddv, dorso-ventral diameter; L, length.
Order Hippuritida Newell, 1965
Suborder Hippuritidina Newell, 1965, in Skelton, 2013b
Superfamily Radiolitoidea d'Orbigny, 1847, in Skelton, 2013a
Family Monopleuridae Munier-Chalmas 1873
Genus Jerjesia Alencaster 1986

Type species. Jerjesia encina Alencaster 1986

Emended diagnosis. Thick-shelled cylindro-conical RV, LV
low conical. Radial bands inconspicuous. Myocardinal apparatus
strong, myophores on both valves located on transverse
thickenings of the shell margin, parallel or slightly oblique to the
commissural plan, am larger than pm with an antero-ventral
elongation. Small body cavity with a rounded transverse outline
in RV, small conical body cavity with a limited cardinal platform
in the dorsal side of LV. Commissural plan strongly oblique to the
shell axis. Ligament ridge marked externally by a ligament groove.
Discussion. In her original description Alencaster (1986)
mentioned: 1 e an accessory cavity anked by a transverse
plate in the LV (see g. 3, plate 1 and g. 6, plate 3), 2 e
central tooth with a curved peaked termination engaged in a small
canal at the base of the opposite anterior tooth (see g. 2, p.
55). Figures provided by the author show that:
1 e the so-called transverse plate and accessory cavity represent the
section of the cardinal platform and adjacent body cavity of the
LV; this controversial interpretation rules out the placement of
Jerjesia in the Caprotinidae Gray emend. Skelton (2013a); it
also precludes, as suggested by Skelton (2013a), the assignment
of the genus to the Polyconitidae;
2 e the peculiar relationships of the central and anterior teeth and
sockets, illustrated on g. 2 (p. 55), hardly correspond to the
photographs of plates 1e3; this point is important because it was
considered as one of the key attributes of the genus (Alencaster, p.
49).
The emended diagnosis and the foregoing remarks suggest that
Jerjesia must belong to the Monopleuridae. The myophoral
orga- nization of Jerjesia departs from those of the
petlalodontid mono- pleurids including: Petalodontia Pocta,
Debrunia Masse and Fenerci- Masse, Pseudopetalodontia
Masse et al., Araeopleura Cox and Mathesia Mainelli, having
all one or two myophoral plates on the LV, protruding into the
opposite valve. It also differs from Mono- pleura Matheron by
the wide transverse, myophoral, anterior and posterior
thickenings of the RV, and from Bicornucopina Hofmann by its
low domal LV and teeth-sockets morphology. The genus
Glossomyophorus Masse et al. has a myophoral plate on the
LV, protruding in a cavity of the opposite valve; a myophoral
organi- zation much alike that of the Caprotinidae than the
Monopleuridae (Masse et al., 1994). Arnaudia Fischer, placed in
the Monopleuridae by Skelton (2013a), but assigned to the
Radiolitidae by Khn (1944), has a depressed LV and lacks a
ligament ridge. Gyropleura Douville has a posterior myophoral
plate in the RV. The foregoing shows that Jerjesia is a valid
taxonomic entity.
Jerjesia chilensis
(d'Orbigny) Fig. 4
1842 e Hippurites chilensis d'Orbigny, p. 107, pl. 22, g. 16
1929 e Agria blumenbachi Studer, Steinmann, p. 116e117,
g. 128 1967 e Agriopleura sp., Herm, p. 662, pl.1, g. 1; pl.
2, gs. 2 and 3 1977b e Agriopleura, Jurgan, p. 422e423, g.
24
1999 e Agriopleura sp. aff. A. blumenbachi (Studer),
Mourgues, p.
156. Pl. 5, g. 7, Figs. text 33 and 34
Type locality. Llanos de Arqueros, northeast of La Serena,
Chile (Fig. 2).
Type level. Member 4 of the Arqueros Formation (Kia4),
lower Albian.
Study material and localities. 4 isolated bivalve specimens,
(SNGM-1991 to 1994), from Llano de Arqueros, 1 isolated specimen
(SNGM 1185-1), two blocks cut in slabs (SNGM 1185-2 and 4), and
two clusters of silicied specimens (SNGM 1185-3 and 4), from
Quebrada El Molle.
Taxonomic avatars of Hippurites chilensis d'Orbigny.
D'Orbigny (1842) described Hippurites chilensis from fossils
samples collected by Ignacio Domeyko near the Arqueros ancient

Fig. 3. Stratigraphic sections of the upper part of the Pabello n Formation at (A) Quebrada Carrizalillo, and (B) Quebrada el Molle, and the upper part of the Arqueros Formation
at (C) Llanos de Arqueros, with position of Jerjesia chilensis bearing beds. The Cerro Pajonales (D) section shows the lower part of the Pabellon Formation and the position of
the Douvillelia skeltoni bearing beds.

Fig. 4. Jerjesia chilensis (d'Orbigny) from Quebrada El Molle (SNGM 1185-1). A, lateral view of a bivalve specimen in a bouquet (SNGM 1185-1a). B,
isolated specimen, showing the ligament groove (lg) of the RV and the attened LV (SNGM-1b). CeD, Longitudinal section of B specimen, showing the
and their interpretation (E, F). G, Longitudinal section of a well preserved bivalve specimen showing the myocardinal elements and their interpretation.
cardinal organization gured in (G) (Llanos de Arqueros, SNGM-1991). I, Transverse section of the RV of a well preserved specimen of Llanos de Arqueros
Antero- posterior section of an isolated LV, partly silicied, showing the myophores. L, M, Ibid., dorsal section showing the teeth (slabs from Quebrada el
2 and 4). Scale bar 1 cm.

mine, northeast of La Serena city. Darwin (1846) assigned to the

d'Orbigny species some specimens coming from the same locality.
During the XX century the d'Orbigny's taxon was either ignored or
re-interpreted. So Hippurites chilensis was ignored by
Fritzsche (1923) who ascribed the forms of Arqueros to Agria
blu- menbachi (Studer), he identied the same taxon at Potrero
Seco in the Copiapo valley; and in the Lower Cretaceous of Per,
namely at Huallanca and Huaraz. Steinmann's work (1929) tends
to bear out the former identications for both Chile and Peru
specimens. Subsequently Herm (1967) studied the shell
microstructure of the bivalves collected from the Arqueros and Lo
Prado formations and similarly, identied Agriopleura sp.. The
ensuing works made in Chile mentioned the same taxon with
different nomenclatural

Dorsal view of an
internal characters,
H, close up of the
(SNGM-1992). J, K,
Molle SNGM 1185-

combinations all revolving around Agriopleura (Tavera,

1956; Corvala n, 1974; Jurgan, 1977a,b; Mourgues, 1999, 2000).
The following deals with the assignment of Hippurites
chilensis and the so-called Agriopleura blumenbachi from
Chile and Peru, to a single taxon Jerjesia chilensis (d'Orbigny).
Description (Fig. 4). RV cylindrical or cylindro-conical, length
up to 8e9 cm, with a rounded elliptical transverse outline (Dap >
Ddv). Average dimensions Dap 2.6 cm, Ddv 2.3 cm.
Outer shell smooth, ligament groove well dened, marked by
downward deected growth lines; radial bands inconspicuous.
Flattened domal LV. Myophores of LV corresponding with
slightly oblique transverse thickenings, small conical body cavity
restricted to the dorsal side, teeth relatively short and subequal.
Myophores of RV

oblique (inward inclination) transverse thickenings. Body cavity of

RV relatively small and rounded. Compared to the outer shell layer,
originally calcitic and frequently silicied, the inner shell layer is
relatively thick except on the ventral side where the two layers are
nearly of same thickness.
Remarks. Well preserved (i.e. non-silicied) specimens from
Lla- nos de Arqueros document the teeth and sockets architecture
whereas myophores exhibit a peculiar colour. Serial longitudinal
sections of a specimen (Fig. 4G, H) show: teeth with a convex laminated microstructure, whereas sockets are lled by a concave
upward (towards the commissure) laminated microstructure, and
myo- phores on both valves are in black. Growth banding is also
recorded in the shell of the Mexican species Jerjesia encina and
the black coloured myophores (bandas oscuras) (Alencaster,
1986) as well. We assume that these features merely reect a
diagenetic context favouring their preservation, i.e. possibly the
preservation of aragonite. Usually they are not preserved, and the
inner shell layer has a homogeneous sparry structure (Fig. 4C,
D), see also Herm (1967).
Discussion. The above descriptions show that Hippurites
chilensis d'Orbigny lack the generic attributes of this genus:
mainly the shell infoldings of the RV and the canaliculated and
porous structure of the LV (Skelton, 2013a). Similarly it does not
match the characters of Agriopleura: mainly the protruding
myophores of the LV, usually depressed, and the salient bands,
depressed interband and incon- spicuous ligament groove of the
RV (Masse and Fenerci-Masse, 2014).
Jerjesia was hitherto represented by a single species J.
encina found in lower Albian beds from the Pihuamo region
(Jalisco) of the southwestern Pacic side of Mexico (Alencaster,
1986). The overall morphology of J. chilensis is similar to J.
encina the dimensions of

which are, however, signicantly higher (D > 5 cm, L up to 10 cm),

moreover in the Mexican species the inner shell layer is relatively
much thicker and the LV more oblique to the shell axis. We ascribe
to Jerjesia chilensis dense tubular congregations (Fig. 5C, D)
found by M. Floquet (personal communication, 2013) in the
Huayhuash cordillera (Chiquian region, Peru). The Jerjesia beds
are overlain by ammonite bearing marls with Prolyelliceras
ulrichi, an index of the middle Albian (L. Bulot, personal
communication, 2013). We as- sume that the monopleurid
rudists from Peru, mentioned by Fritzsche (1923) as Agria, and
especially those gured by Steinmann (1929) as Agria
blumenbachi, actually represent Jerje- sia chilensis. This
species is morphologically comparable to Mono- pleura
marcida White (1884) from the middle Albian of Texas and
Mexico (Garcia-Barrera, 2006). Internal characters of the LV are also
somewhat similar but the myophores of the RV of M. marcida
are represented by ledges (see Skelton, 1978). Monopleura sp.
(p. 518, g. 5) gured by Skelton et al. (2013) from the dredged
material (assumed to be Albian) of the Darwin Guyot, in the
Mid-Pacic Mountains, has a low capuloid LV and is characterized
by the commarginal elongation of the posterior tooth and nearly
sym- metrical myophores, both features depart from Monopleura
s.s., this form has therefore some afnity with Jerjesia.
The Chilean record of the Jerjesia chilensis matches the age of
the Mexican J. encina, and broadens the geographic distribution
of the genus, represented by two vicariant species, which extends
on the Pacic side of both central and south America, over about
48o in latitude (18oN to about 30oS).
Jerjesia chilensis, J. encina, Pseudopetalodontia felixi
(Douville ), Tepeyacia gregaria (Palmer) (see Masse et al.,
2007) and Monopleura marcida (Perkins, 1974; Scott, 1981)
are sub-cylindrical elevators

Fig. 5. Jerjesia chilensis assemblages from Chile (Llanos de Arqueros) (A, B), and Peru (Chiquian region) (by courtesy of M. Floquet, Aix-Marseille University) (C, D). A,
Packed assemblage with upright oriented individuals. B, Silicied bouquet (SNGM 1185-4). C, D, Plan view of irregularly packed assemblages in growth position (eld views).

250

J.-P. Masse et al. / Cretaceous Research 54

(2015) 243e254

(sensu Skelton and Gili, 2002) which tend to build dense aggreLV poorly preserved. Large body cavity in RV. Myocardinal
gations, a mode of assemblage remarkably developed during the
apparatus badly preserved but aligned on the dorsal side of
Albian in the New World (Fig. 5). As elevators they may represent
the RV.
fast growing, somewhat opportunistic species, able to rapidly
The placement of the Chilean specimens in Douvillelia is
colonize shallow water settings, a key adaptative strategy in unmainly based on the characters of the ventral side of the RV and
stable environments with, possibly, high nutrient contents due to
its commissural festoons. The average commissural dimensions
the oceanographic (i.e. coastal upwellings) and volcanic context, i.e.
given below support the identity of the specimens from Mexico
conditions assumed to increase trophic resources (Allmon and
and Chile (Ddv 3.9 cm and Dap 6.0 cm for the Mexican
Martin, 2014).
specimens; Ddv 4.0 cm and Dap 5.0 cm for the Chilean
Genus Douvillelia Alencaster and Pantoja-Alor 1998
specimens) which are therefore assigned to Douvillelia skeltoni.
Type species. Douvillelia skeltoni Alencaster and Pantoja-Alor
Discussion. Douvillelia skeltoni was hitherto known from a
1998 (Fig. 6A)
single area: the Pacic side of the Huetamo region of the
Emended diagnosis. Thick-shelled form with an exogyriform
Mexico (Michoacan), in beds of early Aptian age (Alencaster and
RV (apex twisted) having two longitudinal grooves on the ventral
Pantoja- Alor, 1998). Geological data provided by Abad (1976,
side bounding three rounded salient ridges with convex upward
1980) on rudists collected from the lower part of the Pabellon
growth lamellae, marked by two swellings of the commissure;
Formation at Vallenar, and by Jurgan (1977a) from Cerro
interband a furrow; moderately convex LV. Well developed body
Pajonales, and all ascribed to Agria, suggest that the rudists
cavities, dorsally oblique and conical in both valves, larger in
in question actually belong to Douvillelia skeltoni. The Chilean
RV. Myo- cardinal apparatus aligned parallel to the dorsal margin.
record of the species matches the age of the Mexican one and
Myo- phores on both valves located on transverse thickenings of
broadens the geographic distribution of the species which extends
the shell margin, parallel (posterior side) or slightly oblique
on the Pacic side of both central and south America, over about
(anterior side) to the commissural plan, am transversally
48o in latitude (18oN to about 30oS).
elongated and larger than pm. Central tooth with a crescentic
transverse section, with ante- rior socket located in the concave 5. Palaeoenvironmental, palaeogeographical and
anterior side; narrow apex of anterior tooth obliquely inserted in
palaeoclimatic signicance of Chilean rudist
its socket. Ligament groove and inner crest poorly dened.
assemblages
Discussion. In their original description of the genus
Alencaster and Pantoja-Alor (1998) noticed the overall similarity
AptianeAlbian rudist from Chile are characterized by their very
of Douville- lia with Horiopleura and Polyconites,
low taxonomic diversity, limited to two genera Jerjesia and
notwithstanding the absence of pediculated myophores in the RV
Dou- villelia, belonging to a single family, the Monopleuridae;
of the former genus. They also recognized some afnities of D.
and the monospecic state of the corresponding assemblages,
skeltoni with Monopleura salazari Palmer but rejected the
characters allowing to dene a palaeobiogeographic entity
placement of their taxon in Monopleura. As suggested by Skelton
equivalent to the modern PerueChile Province sensu Fernandez
(2013a) the myophoral characters of Dou- villelia do not match
et al. (2000). These features contrast with those of the Caribbean
those of the Polyconitidae, and the above emended diagnosis
regions where rela- tively high diversity assemblages are
leads to assign this genus to the Monopleur- idae. In placing
documented for the early Aptian, including 4 or 5 families and 10
Douvillelia in the Caprotinidae, Alencaster and Pantoja-Alor
genera (Harris and Hodson, 1922; Alencaster and Pantojareferred to the denition of the family given by Dechaseaux et al.
Alor,1996, 1998; Chartrousse and Masse, 2004; Mitchell, 2013a),
(1969), which included several genera which have been formerly
and the Albian with 6 families and 22 genera (Palmer, 1928;
or subsequently assigned to distinctive families, mainly the
Alencaster, 1986; Chartrousse, 1998; Scott,
Polyconitidae MacGillavry (1937) and the Caprinidae d'Orbigny
2002; Garcia-Barrera, 2006; Masse et al., 2007; Mitchell, 2013a,b,
(Skelton and Masse, 1998; Skelton, 2013a). The family
Skelton et al., 2013). However the two Chilean rudists: Jerjesia
Caprotinidae Gray is presently restricted to two closely related
and Douvillelia are Caribbean genera, and one of them is a
genera Caprotina and Chaperia (Skelton, 2013a). For the
Caribbean species. This similarity suggests that the Caribbean
denition of the genus we focus on the monopleurid
domain was a dispersal centre and probably a centre of origin for
myocardinal organiza- tion and the peculiar morphology of the
both the northern and southern American Pacic margins, as it
RV with an exogyriform habit and a trilobate, lamellar, ventral
was for the main part of the Pacic domain (Skelton et al., 2013).
side associated with commissural swellings. The salient radial
The south- ward dispersion of Jerjesia and Douvillelia from the
bands and depressed
interband depart from that of
Caribbean to the southern Pacic American margin shows that
Monopleura. Noticed that the myo- cardinal traits of
sea surface circula- tion, conveying larvae, did not conform the
Douvillelia are much alike those of Jerjesia (see above).
trajectory imposed by oceanic gyres owing towards the equator;
Description of the Chilean specimens of Douvillelia
consequently coastal eddy currents counteracting the gyres,
Douvillelia skeltoni Alencaster and Pantoja-Alor
were probably the main agents for dispersal. This dispersal mode
1998 Fig. 6 (BeF)
contrasts with the island hopping mode invoked for the
Study material and localities. 10 RV, most of them partly
colonization by rudists of the Pa- cic domain (Skelton et al.,
embedded in a ferrigeneous rocky matrix and more or less silicied,
2013).
some partly preserved; all from Cerro Pajonales (numbers SNGM
The stratigraphic distribution of rudists on the Pacic side of
1983e1990).
America conforms their Caribbean and Pacic mode (Skelton et al.,
Thick-shelled form with an exogyriform RV (apex twisted)
2013): rudists are absent in the late Aptian and disappeared in the
having two longitudinal grooves on the ventral side, bounding
late Albian. In Chile, the regional geology shows that a late Aptian
three rounded or attened salient ridges with convex, festooned,
active volcanism and a late Albian widespread continental regime,
upward growth lamellae, marked by two swellings of the
were the main causal factors for rudist demise.
commissure; interband a furrow. Antero-posterior elongation,
The sketch diagram of Fig. 7, modied from Ramos and Aleman
dorsal side attened and smooth, ventral side convex and lamellar,
(2000), illustrates the palaeogeographic location of rudist-bearing
angular junction between the dorsal and posterior sides. Well
platform carbonates in the island and back-arc settings associated
marked ligament groove, located close to the junction in question.
with the Andean subduction, it also acknowledges the peculiar
oceanographic context which characterizes the Chilean margin
during
the
mid-Cretaceous.
As
stated
above,
the
palaeoceanographic

Fig. 6. Douvillelia skeltoni Alencaster and Pantoja-Alor. A, type gure of Douvillelia skeltoni from Huetamo (Mexico), reproduced from Alencaster and Pantoja-Alor (1998), g.
7 (1),
p. 24, ventral view of RV showing the longitudinal salient lamellar ridges, i.e. ventral bands, bounded by grooves. Be F, Specimens from Cerro Pajonales (Chile). B, Ventral view of
RV showing the ventral bands and interband. C, Transverse section of RV and position of ventral bands and interband (SNGM 1985). D, Close-up of the commissure of the
specimen in B, showing the ventral, festooned morphology corresponding to the bands and interband (SNGM 1983). E, Antero-ventral view of RV showing the lamellar habit
(SNGM 1986). F, Dorsal side (RV) with position of the ligament groove (SNGM 1984). Scale bar 1 cm.

situation of Chile during the Cretaceous suggests that the Humbolt

current may have been present off the Chilean shelf (Hay, 1995), a
model supported by climatic simulations (e.g. Fluteau et al., 2007).
This model postulates SST lower than 18oc, potentially detrimental

for coastal rudist communities, as they are presently for zooxantellate coral communities (Montaggioni, 2007). Nevertheless the
following observations may contribute to explain the presence of
rudist communities.

2
5

J.-P. Masse et al. / Cretaceous Research 54

(2015) 243e254

Fig. 7. Palaeogeographic sketch of the central Andean region showing the association of rudist bearing platform carbonates with volcanic arc settings and their structural and
palaeoceanographic context.

1 e Applying the zooxantellate coral model to rudist communities is misleading. The Chilean shelf is not only characterized by a
low SST (<20oc) due to oceanic currents but mainly by a high
productivity, both induced by vigorous upwellings, and both
detrimental to coral growth which needs relatively high temperatures and low nutrient waters (Montaggioni, 2007). The Humbolt
current system (20oe40oS) presently determines the Temperate
Pacic Realm or PerueChile province of shallow marine ecosystems, breaking at 42oS (Fernandez et al., 2000). As suspensions
feeders (Gili et al., 1995) rudists, may have been, to some extent,
tolerant to high productivity waters. Moreover, all rudists were not
tropical, i.e. strictly associated to warm waters (Masse and FenerciMasse, 2008) and some of them were adapted to subtropical conditions. Notice that the Barremian to early Albian ammonite faunas
from the Chanarcillo basin are dominated by widely distributed
taxa with a Tethyan character (Aguirre-Urreta et al., 2007).
2 e The reconstruction of Fig. 7 postulates the location of rudistbearing platform carbonates in the inner part of the volcanic arc,
this situation may acknowledge a distinctive climatic/oceanographic regime between the offshore and inshore portions of the
arc, the latter being in gross equilibrium with the atmospheric
thermal regime at 30oS. Data from Chumakov (2004) indicate
that in AptianeAlbian times the region was located in the warm
arid belt zone, and because the Gondwana land mass was not
yet fully dislocated, seasonal contrast may have been signicant
(Fluteau et al., 2007). Assuming that the AptianeAlbian palaeoCO2 values were in the range of 700e1400 ppm (Berner and
Kothavala, 2001; Fluteau et al., 2007; Passalia, 2009; Wang et
al., 2014; Godde ris et al., 2014), that is, on average, 2e4 times
those of today, imply a global cool Greenhouse regime in
the sense of Kidder and Worsley (2012). This regime postulates
that wind velocities, wind shear and therefore wind-driven
oceanic circulation must have been more reduced and coupled
with a lesser high to low latitude temperature gradient than
today (Hay and Floegel, 2012). Climatic instability appears to
have been, and still is, an attribute of the Chilean-Peru shelf.
This is mainly due to the El Nino-Southern Oscillation (ENSO)
marked by the temporary increase or decrease

of the trade winds leading to the interruption (El Nino) or intensication (La Nina) of upwellings (Montaggioni, 2007). Instability is
also suggested by oceanographic simulations for the Early Cretaceous (Hay and Floegel, 2012) which document the role of poorlyorganized meso-scale eddies rather than a well-organized ocean
circulation, due to weakened and unstable winds.
Pooling the foregoing observations dealing with moderate
Humbolt current and associated upwellings, leading to some
oceanographic instability, plus continental seasonal contrast, show
that the overall palaeoenvironment, i.e. climatic conditions, were
not fully detrimental for rudist growth in Chile during the
AptianeAlbian, but the above stressing conditions did not allow the
development of high diversity assemblages. Low diversity assemblages are not only observed for rudists but also for corals, exemplied by Hauterivian coral bodies from the adjacent Neuquen
basin in Argentina (Garberoglio et al., 2013). The role of volcanism
as a potential, additional agent for instability, is not obvious
because polytaxic rudist assemblages are known from similar volcanic arc settings in the Caribbean regions (e.g. Mexico).
6. Conclusions
The study of AptianeAlbian rudist faunas from the Chilean
Central Andes documents the presence of two forms:
Douvillelia skeltoni an early Aptian species known hitherto
from Mexico, and Jerjesia chilensis, an endemic Albian species
from Chile and Peru, with a complex taxonomic history. First
described by d'Orbigny as an Hippurites, Jerjesia chilensis, the
rst rudist described from the New World, was then erroneously
ascribed to Agriopleura and this placement led to the wrong
biostratigraphic attributions of the associated beds. The denition
of the genera Jerjesia and Douvillelia is emended, moreover they
are placed in the Monopleuridae rather than in the Caprotinidae
or the Polyconitidae as suggested by former authors. The regional
stratigraphy of the Central Andes, combining ammonites and
rudists, is consistent with the Caribbean stratigraphic distribution
of Jerjesia and Douvillelia. Andean occur- rences of the two
genera broaden their distributional area on the

Pacic side of Americas, and testify their biostratigraphic value.

Oceanographic conditions of the Chilean Pacic margin during the
AptianeAlbian, are assumed to have been somewhat comparable to
modern ones, (cold oceanic current, upwellings, high productivity
and thermal instability), but in the context of a cool greenhouse
regime which implies a moderate intensity of the above phenomenon. The oceanographic regime and volcanic arc settings as well,
were not fully detrimental for the development of rudists which are
nevertheless characterized by a low taxonomic diversity, indicative
of stressing conditions. The assemblages are impoverished when
compared to their low latitude homologues from the Caribbean
regions which were functioning as a dispersal centre.
Acknowledgements
We warmly thank M. Floquet and L. Bulot (Aix-Marseille University) for the data regarding the specimens and age of
Jerjesia bearing beds from Peru. We appreciated the help of
A. Rubilar (SERNAGEOMIN), curator of the paleontological
collection, for making the fossils at his care available for our study.
We are grateful to S.I. Sano for his careful and helpful review of an
earlier version of the manuscript.
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