Palaeogeography, Palaeoclimatology, Palaeoecology 250 (2007) 216 – 236

www.elsevier.com/locate/palaeo

The La Voulte Lagerstätte (Callovian): Evidence for a deep water

setting from sponge and crinoid communities
Sylvain Charbonnier ⁎, Jean Vannier, Christian Gaillard,
Jean-Paul Bourseau, Pierre Hantzpergue
UMR 5125 PEPS, CNRS, France; Université Lyon 1, Campus de la DOUA, Bâtiment Géode, 69622 Villeurbanne Cedex, France
Received 27 November 2006; received in revised form 13 March 2007; accepted 19 March 2007

Abstract

The siliceous sponges and crinoids from the Chénier Ravine (France, Lower Callovian) are used here as biological markers to
characterize the palaeoenvironment of the adjacent and contemporaneous La Voulte Lagerstätte that is remarkable for its unique
soft-bodied fauna (e.g. worms, cirrate octopods, vampire squids). Sponges are abundant with dominant hexactinellids (80%) and
lithistids (20%). Four lines of fossil evidence, supported by Recent analogues, indicate that this sponge community inhabited deep-
water setting under dysphotic or aphotic conditions: (1) the dominance of hexactinellids; (2) the prevalence of cone-shaped and
erect morphologies that usually characterize Recent bathyal sponges; (3) the close similarities with Recent hexactinellids from the
continental slope and; (4) the lack of encrustation by photophilic organisms. Attachments of sponges on hard substrate (e.g.
crystalline basement) and their preservation in soft sediments (e.g. muds) indicate heterogeneous bottom conditions. The Chénier
fauna also contains small stalked, asymmetric cyrtocrinid crinoids that are known to live on hard substrates in bathyal environments
such as the SW Pacific steep seamounts. Convergent palaeoenvironmental clues obtained from both crinoids and siliceous sponges
support the notion that the La Voulte area, including the La Voulte Lagerstätte, was situated in the upper part of the bathyal zone
near the slope-basin transition with a water depth most probably exceeding 200 m. Supporting evidence from heterogeneous
substrates and complex fault systems indicate a depositional environment along the external part of the slope where steep
topographies and blocks usually favour the settlement of cyrtocrinid crinoids and hexactinellid sponges. La Voulte may therefore be
one of the rare and extremely precious Mesozoic Lagerstätten to have fossilized a deep marine environment.
© 2007 Elsevier B.V. All rights reserved.

Keywords: Siliceous sponges; Stalked crinoids; Bathyal; Lagerstätte; Jurassic; Callovian; France

1. Introduction first mentioned the presence of articulated ophiuroids in

this locality. Indeed, this Lagerstätte contains a diverse
Interest in the Middle Jurassic (Callovian) La Voulte and remarkably preserved fauna dominated by ophiur-
Lagerstätte dates back to the nineteenth century and the oids and crustaceans, with also numerous soft-bodied
pioneer works of Fournet (1843) and Lory (1854) who organisms. By far, the most frequent animals of the La
Voulte biota are the delicate endemic ophiuroid Ophio-
⁎ Corresponding author. Fax: +33 4 724 483 82. pinna elegans (Hess, 1960) and small thin-shelled
E-mail address: [email protected] bivalves (Bositra buchi), both found in large numbers
(S. Charbonnier). on bedding planes. Crustaceans are represented by
0031-0182/$ - see front matter © 2007 Elsevier B.V. All rights reserved.
doi:10.1016/j.palaeo.2007.03.013
 S. Charbonnier et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 250 (2007) 216–236 217

abundant decapods, mysids and, cumaceans, that are small size of exposures, a homogenous facies with very
most frequently three-dimensionally preserved in side- few sedimentological features, fossils concentrated
ritic nodules (van Straelen, 1922, 1923, 1925; Secrétan, within less than 5–6 m of marls and, above all, a
1983; Secrétan and Riou, 1983, 1986; Carriol and Riou, tectonic configuration with steeply dipping (dip ca. 60°).
1991; Schweigert et al., 2006). More enigmatic and also Considering that previous studies were unsuccessful and
very abundant at La Voulte are the so-called thylacoce- that the La Voulte Lagerstätte itself had low potential to
phalans (see Secrétan, 1985; Vannier et al., 2006 for provide a precise palaeobathymetry, we focussed our
complete references) that were unusually large bivalved research on the adjacent and contemporaneous outcrop
arthropods armed with prehensile appendages and of the Chénier Ravine (ca. 150 m from the La Voulte
bearing huge visual organs. Both in terms of diversity Lagerstätte) that offered better conditions of exposure
and abundance, the arthropod fauna from La Voulte (relatively thick deposits and larger outcrops). This
rivals that of the celebrated Solnhofen Lagerstätte (van locality yielded abundant and well-preserved siliceous
Straelen, 1922, 1925; Etter, 2002a). Soft-bodied organ- sponges and small stalked crinoids that, via comparisons
isms are represented by exceptional specimens of cirrate with their Recent analogues, turned out to be important
octopods (Fischer and Riou, 1982a) and other coleoid biological indicators of environmental conditions such
cephalopods (Fischer and Riou, 1982b), vampire squids as the water depth. The ecological significance of these
(Fischer and Riou, 2002), worms (possibly sipunculans; two groups is analysed in the present paper and leads to
Alessandrello et al., 2004) and sea spiders (Pycnogo- the first accurate interpretation of the palaeoenviron-
nida; S. Charbonnier and collaborators, in progress). ment of the La Voulte Lagerstätte.
Although several spectacular cases of soft-bodied
preservation have been succinctly described over the 2. Geological setting and fossil localities
years (e.g. cephalopods, worms; Fischer and Riou,
1982b; Alessandrello et al., 2004), no comprehensive The La Voulte Lagerstätte and the nearby Chénier
analysis of the organisms and communities present in Ravine are both located in south-eastern France
this exceptional biota has been made so far by using (Ardèche) along the right bank of the Rhône River
modern techniques and detailed comparisons with (ca. 150 km south of Lyon; Fig. 1) and belong to the
modern ecosystems. Equally surprising is the lack of eastern sediment cover of the Massif Central (Elmi,
knowledge concerning the palaeoenvironmental setting 1967; Fig. 1). The Massif Central is an Hercynian
of the La Voulte Lagerstätte and the taphonomic pro- crystalline complex whose eastern margin was faulted
cesses involved in the exceptional preservation of the and tilted during the Middle Jurassic. A major sub-
fauna. Elmi (1990), and Alméras and Elmi (1996) vertical normal fault, the so-called La Voulte fault (N
proposed a complex submarine topography consisting 54°, inherited Hercynian direction) was very active
of tilted blocks possibly lying at the transition between during the Callovian and cuts through the whole area
slope and basin in the bathyal zone. According to Etter (Fig. 1). The two studied localities (La Voulte
(2002b) the La Voulte area, during the Early Callovian, Lagerstätte and Chénier Ravine) are also separated by
was situated slightly below the storm wave base. The a transverse normal fault (the Col de Viau fault, N 120°)
interpretation of Fischer (2003), partly derived from the with very limited horizontal displacement (ca. 25–
Sargasso Sea model, is that of a bathyal environment 50 m). The fact that the La Voulte Lagerstätte and the
with most organisms supposedly attached to algae Chénier Ravine belong to two separate blocks explains
floating in the uppermost levels of the water column. part of the differences observed in the sedimentological
Alessandrello et al. (2004) interpreted the palaeoenvir- features and faunal composition of these two localities
onment of La Voulte as a restricted coastal lagoon with (Fig. 1).
stagnant waters. The most recent palaeogeographic reconstructions
At the beginning of the 1980s, successive field for the Callovian place the La Voulte area along the
campaigns conducted by the Natural History Museum western margin of the Tethys Ocean and adjacent to the
Paris (Fischer, 2003), yielded a substantial amount of Massif Central which was probably submerged at that
new fossil material and confirmed the exceptional time (Enay et al., 1993a) (Fig. 2). The Tethyan margin
richness of the La Voulte biota but, unfortunately, failed running east of La Voulte was characterized by a
to provide precise information on the palaeoenviron- complex submarine palaeotopography of tilted blocks
mental parameters and ecological organization of the generated by a series of inherited Hercynian and trans-
biota. Several factors have obviously hindered accurate verse faults (Elmi, 1967, 1990). Platforms lying on the
palaeoenvironmental studies at La Voulte. These are the submerged Massif Central were the principal source of
218 S. Charbonnier et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 250 (2007) 216–236

Fig. 1. Locality map (modified from Elmi, 1967).

carbonate sediments whereas terrigenous inputs originat- lopods, and more occasional occurrences of sea stars,
ed partially from the sub-marine erosion of the Hercynian worms, pycnogonids, marine crocodiles, and sharks.
basement and its Early Mezosoic (e.g. Triassic) sediment The deposits of the Chénier Ravine crop out in
cover (Elmi, 1967, 1990). badlands 150 m SW of the La Voulte Lagerstätte and are
The biostratigraphy of the La Voulte area (both separated from it by the Col de Viau transverse fault
Lagerstätte and Chénier Ravine) was established by (Fig. 1). They consist of ca. 20 m of marls and marly
Elmi (1967) and is based on ammonite biozonation. The limestones (Fig. 3). The first 15 m of the section are
fossiliferous layers of the two localities belong to the characterized by alternations of marls and limestones
Lower Callovian Gracilis Biozone (Elmi, 1967). yielding siliceous sponges and stalked crinoids. Abundant
The fossiliferous layers of the La Voulte Lagerstätte sponges occur in the uppermost 5 m of the section forming
crop out in the Mines Ravine (Fig. 1) SW of the town of the so-called Lentille à Hexactinellides (Sayn and Roman,
La Voulte-sur-Rhône (Fig. 1) and occur within a 1928; Fig. 3). This conspicuous fossiliferous horizon is
relatively thin interval (ca. 5–6 m; Fig. 3). Their overlain by very limited black layers rich in phosphatized
lithology consists of marls topped with ca. 15 m thick ammonites, followed by thin iron carbonate layers and
iron deposits that were exploited in the 19th century. The slumped calcareous beds possibly generated by synsedi-
basal marls contain numerous sideritic nodules which mentary slides (Elmi, 1990) (Fig. 3).
frequently preserve uncompacted crustaceans and more The small number and poor preservation of ammo-
or less undeformed soft-bodied cephalopods. Some nites (Gracilis Zone) do not allow high-resolution
marly horizons are also locally rich in soft-bodied fossils biostratigraphical correlations between the two sections.
(e.g. cephalopods, worms) that occur more flattened. Lithological correlations are also limited because of the
Thin iron carbonate layers with abundant ophiuroids lack of well-defined marker-beds both in the monoto-
occur throughout the succession (Fig. 3). Although an nous facies of the Chénier Ravine (marls and marly
updated faunal inventory of the La Voulte Lagerstätte is limestones) and the Lagerstätte (marls with nodules and
beyond the scope of the present paper, preliminary field sideritic horizons).
observations (SC) confirm that the most typical elements The siliceous sponges from the Chénier Ravine were
of the biota are: (1) decapod and mysid crustaceans, (2) first described by Dumortier (1871; ca. 12 species)
thylacocephalan arthropods, (3) ophiuroids, (4) cepha- and revised by Moret (1926, 1928; 16 species) that
 S. Charbonnier et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 250 (2007) 216–236 219

Fig. 2. Palaeogeography of the Western Tethys during the Callovian (162 to 158 Ma) showing the La Voulte area (redrawn from Enay et al., 1993b).

both established systematic inventories. A new taxon Dumortier (1871) and 48 by Moret (1928) were also
(Trochobolus perfectus) was also described by Lagneau- examined. The whole fossil material consists of approx-
Hérenger (1951). The stalked crinoids were reported to imately 240 specimens of siliceous sponges (both hexa-
occur in this locality by Sayn and Roman (1928, faunal ctinellids and lithistids) and very abundant remains of
list) but, so far, have neither been described nor illustrated stalked crinoids: several hundreds of isolated columnals
in detail. (stem elements) and brachials (arm elements) of cyrtocri-
nids and isocrinids, and cyrtocrinid aboral cups (=calices,
3. Materials and methods 125 specimens).
We revised and updated the previous inventories of
Our study is based on fossils collected during suc- the sponge fauna made by Dumortier (1871), Moret
cessive field excursions in 2003 and 2004 (SC) from (1926, 1928) and Lagneau-Hérenger (1951) in the light
several sections of the Chénier Ravine. This new of personal observations (CG, SC) and recent works
material adds to siliceous sponges and stalked crinoids dealing with the systematics of the group (Melh, 1992;
deposited in the collections of the Université Lyon 1 (72 Pisera, 1997; Tabachnick and Reiswig, 2002). We are
specimens) and the Institut Dolomieu, Université de using here the terminology and updated classification
Grenoble (44 specimens). Eight specimens published by proposed by Reid (2004a,b,c). The diversity and relative
220 S. Charbonnier et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 250 (2007) 216–236

Fig. 3. Lithological logs of the Chénier Ravine and the La Voulte Lagerstätte with fossil distribution and biostratigraphic correlations.

abundance of hexactinellid and lithistid species were observation of the external shape of the fossil sponges
calculated in order to establish comparisons with the (Fig. 4). The relative percentage of these morphotypes
composition of Recent sponge communities. We were was calculated from ca. 240 specimens from the Chénier
able to distinguish ten different morphotypes from the Ravine.
 S. Charbonnier et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 250 (2007) 216–236 221

1979; Žítt, 1983) were used for the identification of the

specimens from the Chénier Ravine. Counts of crinoid
cups were used to evaluate the relative abundance of
crinoid species.
The fact that most crinoids from the Chénier Ravine
belong to extant groups enabled precise morphological
and ecological comparisons with Recent crinoids both
from the Order Cyrtocrinida (Suborder Cyrtocrinina)
and the Family Isocrinidae sensu Rasmussen, 1978
(= Pentacrinidae sensu David et al., 2006).

4. Results

4.1. Siliceous sponges

4.1.1. Diversity and relative abundance

The sponge fauna from the Chénier Ravine consists
of 17 different species (Table 1). Eleven of them are
hexactinellids, the rest being assigned to the lithistids.
The hexactinellids are represented by 4 hexactinosoan
and 2 lychniscosan families, the lithistids only by the

Table 1
List of siliceous sponges from the Chénier Ravine (Lower Callovian)
Hexactinellids Lithistids
HEXACTINOSA RHIZOMORINA
Craticulariidae Azoricidae
Craticularia dichotomans Azorica calloviensis
(Dumortier, 1871) Moret (1928)
Craticularia parallela Cytoracia cylindrica
(Goldfuss, 1826) (Dumortier, 1871)
Craticularia rhizoconus Cnemidiastridae
(Quenstedt, 1878) Cnemidiastrum stellatum
Cribrospongiidae (Goldfuss, 1826)
Gevreya synthetica Scleritodermatidae
Moret (1926) Verruculina gevreyi
Staurodermatidae Moret (1926)
Rhodanospongia robusta Verruculina multiforis
Moret (1926) (Dumortier, 1871)
Saynospongia palmicea Jereicidae
(Dumortier, 1871) Moretispongia praegnans
Stauroderma inversa (Dumortier, 1871)
(Dumortier, 1871)
Family uncertain
Porospongia fungiformis
(Goldfuss, 1826)
LYCHNISCOSA
Sporadopylidae
Fig. 4. Sponge morphotypes present at the Chénier Ravine. Sporadopyle micropora
Moret (1926)
Sporadopyle obliqua
(Goldfuss, 1826)
Previous monographs on stalked crinoids (de Loriol, Pachyteichismatidae
1882–1884; Jaekel, 1907; Rasmussen, 1961; Arendt, Trochobolus perfectus
1974; Hess, 1975; Rasmussen, 1978; Pisera and Dzik, Lagneau-Hérenger (1951)
222 S. Charbonnier et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 250 (2007) 216–236

Fig. 5. Siliceous sponges from the Chénier Ravine, Lower Callovian. (a) Gevreya synthetica (ID), general view and lateral view, note the encrusting
base; (b) Saynospongia palmicea (FSL 710001), general view and lateral view, palmate shape; (c) Azorica calloviensis (FSL 710002), lateral view
and upper face view, cup shape; (d) Verruculina gevreyi (FSL 710003), fragmentary specimen, lower and upper face view, flabellate shape, note the
numerous pustular pores on both surfaces; (e) Verruculina multiforis (FSL 710004), upper face view, earlike shape, note the pustular pores on the
upper surface; (f ) Verruculina multiforis (FSL 710005), holotype, lower and upper face view, earlike shape; (g, h, i) Moretispongia praegnans (ID),
lateral view, club-like shape with pustular pores; (j) Stauroderma inversa (FSL 710006), internal face view showing the siliceous skeleton, sub-
cylindrical shape; (k) Stauroderma inversa (FSL 710007), general view, vase shape, note the encrusting base; (l) Stauroderma inversa (FSL 710008),
general view, conical shape; (m) undetermined species (FSL 710034), lateral view and upper or lower face view, biconvex shape. Collections:
ID = Institut Dolomieu, Université de Grenoble (Moret Collection); FSL = Université Lyon 1.
 S. Charbonnier et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 250 (2007) 216–236 223

Fig. 6. Siliceous sponges from the Chénier Ravine, Lower Callovian. (a) Stauroderma inversa (FSL 710009), fragmentary specimen, general view,
narrow tubular shape; (b) Craticularia parallela (FSL 710010), general view, narrow tubular shape; (c) Craticularia parallela (FSL 710011), general
views, sub-cylindrical shape; (d) Craticularia parallela (FSL 710012), general view, tubular shape with regular constrictions; (e) Craticularia
parallela (FSL 710013), lateral view, branching shape; (f, g) Craticularia dichotomans (FSL 710014, 710015), fragmentary specimens, surface
views, cup shape; (h) Attachment by encrusting base (FSL 710017), lateral view and top side view; (i) Attachment by encrusting base (FSL 710016),
lateral view, note the bivalve shell as substrate; (j) Attachment by encrusting base (FSL 710018), lateral view and under side view, note the mica-
schist fragment as substrate (white arrows); (k) Craticularia sp. (FSL 710019), general view of the base; (l) Serpulids on outer surface of Craticularia
parallela (ID), general view, note the two worm tubes; (m) Small oyster valve (FSL 710005) fixed on the upper edge of V. multiforis holotype
(see Fig. 5f ); (n) Cyrtocrinid holdfast (n1) and small encrusting sponge (n2) (white arrows) (FSL 710020) attached to sponge fragment, upper surface
view. Collections: ID = Institut Dolomieu, Université de Grenoble (Moret Collection); FSL = Université Lyon 1.
224 S. Charbonnier et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 250 (2007) 216–236

rhizomorine group (4 families). Most species such as (4.6%) dominate the lithistid fauna, other species being
Gevreya synthetica (Fig. 5a), Rhodanospongia robusta, below 3% (Table 2).
Saynospongia palmicea (Fig. 5b), Azorica calloviensis
(Fig. 5c), Verruculina gevreyi (Fig. 5d), Verruculina 4.1.2. Preservation, morphotypes, mode of attachment
multiforis (Fig. 5e, f ), Moretispongia praegnans and epizoans
(Fig. 5g–i), and most probably also Stauroderma The siliceous sponges from the Chénier Ravine are
inversa (Figs. 5j–l and 6a) are known only from the not preserved in life position and are found scattered in
La Voulte area. In this locality, the three sponges with sediments. However, many delicate specimens such as
the most remarkable morphologies are: (1) the upright branching and narrow tubular like ones are complete
hexactinellid G. synthetica (Fig. 5a) with its broad basal (Fig 6b, e) and a number of them have their basal
attachment, (2) the small club-like lithistid M. praeg- attachment preserved (Fig. 5a). Rare basal attachments
nans (Fig. 5g) and, (3) the palmate hexactinellid on fragments of crystalline rocks (Fig. 6j) provide
S. palmicea (Fig. 5b). Several sponges exhibit very evidence that these sponges grew on the submarine
uncommon external morphologies, for example small exposed areas of the crystalline basement. Their original
biconvex forms (Fig. 5m) that may represent a new habitat was obviously situated at some distance from
taxon. However, the small number of available speci- their depositional area which is overwhelmingly dom-
mens does not allow formal descriptions. inated by marls. However, their excellent preservation is
Hexactinellids account for 80.1% of the sponge not consistent with a post-mortem transportation driven
biodiversity, lithistids representing only 19.9% of the by high-energy currents. The siliceous sponge fauna
total number of sponge species (Table 2). The most of the Chénier Ravine is interpreted here as being
abundant hexactinellid species are Craticularia paral- parautochthonous.
lela (34.0%; Fig. 6b–e), S. inversa (19.9%) and, Cra- Recent sponges display a great plasticity of body
ticularia dichotomans (10.8%; Fig. 6f, g), followed by form which may be linked to environmental parameters,
G. synthetica (5.8%). All the other hexactinellids are such as water flow dynamics, sedimentation rate and,
minor elements of the sponge fauna with relative substrate (Tabachnick, 1991; Finks, 2003). Changes in
abundances lower than 3.0%. Compared with hexacti- body form can arise during the life span of individual
nellids, the diversity and numerical abundance of organisms in response to environmental pressures.
lithistids is low. M. praegnans (6.6%) and V. gevreyi Sponge morphotypes therefore appear as promising

Table 2
Siliceous sponges from the Chénier Ravine (Lower Callovian): results of quantitative analysis (number of samples and percentages)
Taxa This work Collections FSL Collections ID Samples Samples Total samples Percentages (%)
Moret (1928) Dumortier (1871)
Craticularia parallela 44 29 4 5 82 34.0
Stauroderma inversa 8 20 14 5 1 48 19.9
Craticularia dichotomans 9 12 1 3 1 26 10.8
Gevreya synthetica 5 4 1 4 14 5.8
Sporadopyle micropora 4 3 7 2.9
Sporadopyle obliqua 1 5 6 2.5
Saynospongia palmicea 1 1 2 4 1.7
Rhodanospongia robusta 3 3 1.2
Craticularia rhizoconus 1 1 0.4
Porospongia fungiformis 1 1 0.4
Trochobolus perfectus 1 1 0.4
Total hexactinellids 80.1
Moretispongia praegnans 10 5 1 16 6.6
Verruculina gevreyi 1 6 4 11 4.6
Cnemidiastrum stellatum 2 2 2 2 8 3.3
Cytoracia cylindrica 1 5 6 2.5
Verruculina multiforis 3 1 1 5 2.1
Azorica calloviensis 1 1 2 0.8
Total lithistids 19.9
Total 69 72 44 48 8 241 100.0
Collections: ID = Institut Dolomieu, Université de Grenoble (Moret Collection); FSL = Université Lyon 1.
 S. Charbonnier et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 250 (2007) 216–236 225

and reliable palaeoenvironmental indicators. We were Table 3

able to distinguish 10 different morphotypes among the List of crinoid species from the Chénier Ravine (Lower Callovian)
sponge fauna from the Chénier Ravine (Figs. 4 and 7). Stalked crinoids
The narrow tubular (28.8% of specimens) and conical CYRTOCRINIDA
(16.3%) morphologies are dominant, followed by sub- Eugeniacrinitidae
cylindrical (10.8%), club-like (10.8%) and, branching Eugeniacrinites dumortieri de Loriol (1882)
Lonchocrinus sp. Jaeckel (1907)
(10.0%) morphotypes. Much less frequent are cup-like
Sclerocrinidae
(7.9%), earlike (7.1%), globular (4.2%), biconvex Cyrtocrinus nutans (Goldfuss, 1829)
(2.5%) and palmate (1.7%) sponges (Fig. 7). Cyrtocrinus nutans var. voultensis (Sayn and Roman, 1928)
Today, the type of substrate (e.g. mud, hard rock) Gammarocrinites compressus (Goldfuss, 1829)
largely determines the mode of attachment of sponges Phyllocrinidae
Phyllocrinus colloti de Loriol (1882)
and the success of their settlement. Some attach to a hard
Uncertain family
substrate by encrusting, others anchor in soft sediments Dolichocrinus aberrans (de Loriol, 1882)
by a basal spicule tuft. Numerous sponges from the ISOCRINIDA
Chénier Ravine have no basal part preserved making Isocrinidae
their mode of attachment uncertain. All are embedded in Balanocrinus dumortieri de Loriol (1887) (in de Loriol 1884–1889)
marly sediments but with no evidence of anchoring into
the soft sediment. Nevertheless, several specimens have
clearly an encrusting base which confirm their attach- even the dead parts of living sponges, are also frequently
ment on a hard substrate (Figs. 5a, k and 6h–k). Their encrusted by various small sessile invertebrates and
encrusting bases are substantially enlarged and may microbial organisms. This phenomenon is very common
incorporate either fragments of crystalline rock (mica- in the Recent and was also recognized in the fossil
schist; Fig. 6j) or shell fragment (bivalve; Fig. 6i). record, especially among the Upper Jurassic siliceous
Commensalism is extremely common in sponges sponge communities from the Tethys Ocean, that are
(Brusca and Brusca, 2002) and the body of numerous typically associated with sponge/microbialite circalit-
species is utilized as a temporary refuge or a more toral bioherms (Gwinner, 1976; Gaillard, 1983; Pisera,
permanent settlement by a number of small invertebrates 1997; Olivier et al., 2004). The circalittoral Upper
(crustaceans, ophiuroids, worms). Dead sponges, or Jurassic sponges show a selective distribution of their

Fig. 7. Pie-diagram showing the sponge morphotype frequency from the Chénier Ravine (ca. 240 specimens; results expressed in percentages of
specimens).
226 S. Charbonnier et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 250 (2007) 216–236

encrusters. Stromatolitic microbial crusts develop in the different types of encrusters were recognized: (1) serpulids
upper parts of the body exposed to the light, whereas (ten isolated tubes on different sponge fragments; Fig. 6l),
encrusters such as serpulids, bryozoans and thecideid (2) small oysters (Fig. 6m), (3) small stalked crinoids (one
brachiopods concentrate in the sheltered parts facing cyrtocrinid holdfast; Fig. 6n1) and, (4) encrusting sponges
downwards. This polarity is a valuable indicator for (Fig. 6n2).
light penetration (euphotic zone) and sedimentation rate
in ancient marine environments. By contrast with these 4.2. Stalked crinoids
Upper Jurassic circalittoral sponges, those from the
Chénier Ravine are devoid of photophilic microbial 4.2.1. Diversity and relative abundance
crusts and animal encrusters are extremely rare. When The crinoid fauna from the Chénier Ravine consists
present, these encrusters are invariably found in the of cyrtocrinids with a lesser proportion of isocrinids
upper parts of the external surfaces of the sponges. Four (=pentacrinids) (Table 3). Both groups have Recent

Fig. 8. General morphology of stalked crinoids. (a) Cyrtocrinid, reconstruction of Lonchocrinus (composite drawing from Jaekel (1907) and Pisera and
Dzik (1979)); (b) Cyrtocrinid, reconstruction of Eugeniacrinites (redrawn from Jaekel (1907) and Hess (1975)); (c) Cyrtocrinid, reconstruction of
Cyrtocrinus (redrawn from Jaekel (1907) and Hess (1975)); (d) Cyrtocrinid, reconstruction of Phyllocrinus (redrawn from Arendt (1974));
(e) Isocrinid, living specimen from Bahamas (depth: 692 m) (redrawn from Messing (1985)); (f ) Cyrtocrinid, recent specimen of Gymnocrinus richeri
(FSL 710035) from New Caledonia (MUSORSTOM 6 cruise, Loyalty Rise, Station DW 471, depth: 470 m, courtesy N. Améziane and M. Roux).
 S. Charbonnier et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 250 (2007) 216–236 227

representatives (e.g. Améziane and Roux, 1997). Numer- rinids (Fig. 8e) display a long heteromorphic (nodals with
ous recent oceanographic expeditions and in situ obser- prominent cirri) stem surmounted by a rather small and
vations conducted with submersibles or camera sledges low cup with a long multi-armed crown. Brachial crown
have provided new data on their morphology and ecology works in currents like a parachute (Macurda and Meyer,
(see David et al., 2006). Cyrtocrinids are small, stalked 1974, 1976). Their length generally exceeds 10 cm.
and sturdy crinoids (length ca. 4 to 5 cm) with short arms The cyrtocrinid crinoids from the Chénier Ravine are
that can be enrolled for protection (see Fig. 8 for typical represented by isolated remains (cups, brachials, colum-
morphologies). The aboral cup (or calyx) may be asym- nals and holdfasts) that fall within three different families:
metric. The stem is short with few columnals (b 10) and its (1) the Eugeniacrinitidae with Eugeniacrinus dumortieri
distal end cemented to the substrate. By contrast, Isoc- (Figs. 8b and 9a) and Lonchocrinus sp. (Figs. 8a and

Fig. 9. Cyrtocrinids from the Chénier Ravine, Lower Callovian. (a) Eugeniacrinus dumortieri (FSL 710036), cup, general view; (b) Lonchocrinus sp.
(FSL 710037), primibrachial, ventral view and lateral view; (c) Lonchocrinus sp. (FSL 710038), primibrachial, dorsal view; (d) Cyrtocrinus nutans
(FSL 710039), primibrachial, general view and dorsal view, note the convex verrucose surface; (e) Cyrtocrinus nutans (FSL 710040), spoon-like cup
with fused proximal columnals, lateral view; (f, g) Cyrtocrinus nutans (FSL 710041, 710042), cups, oral views; (h) Cyrtocrinus nutans (FSL
710043), asymmetric cup with pronounced pits, dorsal view; (i) Cyrtocrinus nutans var. voultensis (FSL 710044), asymmetric cup with verrucose
surface, dorsal view; (j) Gammarocrinites compressus (FSL 710045), cup, oral view; (k) Eugeniacrinites? barrel-shaped columnal, (FSL 710046),
lateral view; (l, m) Cyrtocrinus? (FSL 710047, 710048) biconcave-shaped columnals, lateral views. All scanning electron micrographs. Collections:
FSL = Université Lyon 1.
228 S. Charbonnier et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 250 (2007) 216–236

9b, c), (2) the Sclerocrinidae with Cyrtocrinus nutans electron microscopy revealed extremely well-preserved
(Figs. 8c and 9d–h), Cyrtocrinus nutans var. voultensis microstructures (Fig. 10g) with no abrasion marks of
(Fig. 9i) and Gammarocrinites compressus (Fig. 9j) and, the ossicles. This type of preservation is in no way con-
(3) the Phyllocrinidae with Phyllocrinus colloti (Figs. 8d sistent with long and/or high-energy transportation of
and 10b, c). Dolichocrinus aberrans (Fig. 10a) is a rare crinoid ossicles by currents or mudflows. Similar with
small crinoid of uncertain affinities although superficially their sponge associates (see previous chapter), cyrtocrinids
resembling cyrtocrinids. Circular columnals of the iso- are most likely autochthonous or parautochthonous, their
crinid (=pentacrinid) Balanocrinus dumortieri (Fig. 10e, habitat lying probably close to their area of deposition.
f ) were also found among the assemblages of crinoid The cyrtocrinid remains are diverse. Cups are the most
ossicles. Contrasting with cyrtocrinids, Balanocrinus was abundant elements and were used here for quantitative
a large isocrinid stalked crinoid (length ca. 30 to 50 cm) analyses. The very frequent asymmetry of their shape most
that anchored to the sea bottom by means of distal cirri probably results from the action of marine currents
(Fig. 8e). (Ausich et al., 1999a). Spoon-like pattern (often obtained
C. nutans (71% of cyrtocrinid remains, Table 4) is by the fusion of cup with the proximal columnals; Fig. 9e)
numerically the most abundant species, the rest of the would suggest the prevalence of unidirectional currents in
crinoid fauna representing minor components (less than the area where the cyrtocrinids once lived (see also Roux
10.0%). et al., 1991a, for Recent data from CALSUB cruise). Other
cups bear more or less prominent swellings that have
4.2.2. Preservation, morphological diversity of ossicles circular pits corresponding to the ichnogenus Tremichnus
The cyrtocrinid elements present a low degree of attri- (Brett, 1985; Feldman and Brett, 1998) made by borers of
tion. Columnal surfaces, brachial facets and cup surfaces unknown identity (Fig. 9h). Columnals (stem elements)
are neither abraded nor rounded or bio-eroded. Scanning are frequent but less abundant than cups. They are either

Fig. 10. Crinoids from the Chénier Ravine, Lower Callovian. (a) Dolichocrinus aberrans (FSL 710049) cup with fused proximal stem, general view;
(b, c) Phyllocrinus colloti (FSL 710050, 710051) cups, general views; (d) Cyrtocrinid holdfast (FSL 710052), top side view with distal columnal;
(e, f) Balanocrinus dumortieri (FSL 710053, 710054), columnals, general views; (g) Stereom in the Lonchocrinus sp. ossicle (FSL 710037), detail of
Fig. 9b. All scanning electron micrographs. Collections: FSL = Université Lyon 1.
 S. Charbonnier et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 250 (2007) 216–236 229

Table 4 Janussen et al., 2004). Recent lithistid sponges live at

Crinoids from the Chénier Ravine (Lower Callovian): results of much shallower depths (200 to ca. 700 m) than hexa-
quantitative analysis on the cyrtocrinids cups (number of samples and
percentages)
ctinellids, the deepest habitats being occupied by
rhizomorines (van Soest and Stentoft, 1988; Pisera,
Cyrtocrinid remains This Collections Total Percentages
(cups) work FSL samples (%)
1997). Exceptionally high concentrations of siliceous
sponges occur along the slope areas where water flow
Eugeniacrinites 3 6 9 7.1
dynamics offer optimal conditions (current velocity
dumortieri
Lonchocrinus sp. 2 2 1.6 gradient, sufficient organic particles in suspension, low
Cyrtocrinus nutans 13 77 90 70.9 sedimentation rate) to their settlement and development
C. nutans voultensis 4 4 3.1 (Gage and Tyler, 1991). In the Mediterranean Sea, the
Gammarocrinites 11 11 8.7 abrupt cliffs that occur along the fringes of the con-
compressus
tinental shelf host a remarkably rich fauna of siliceous
Phyllocrinus colloti 9 9 7.1
Dolichocrinus 2 2 1.6 sponges (Vacelet, 1969, 1979). In general, the uneven
aberrans topography of the bathyal benthic environments is
Total 16 111 127 100.0 propitious to the development of high-density popula-
Collections: FSL = Université Lyon 1. tions of hexactinellid sponges (Pérès, 1972). Submarine
observations indeed reveal the presence of high con-
centrations of these organisms near rocky steps where
barrel-shaped (i.e. low simple cylindrical shape; Fig. 9k) topographic breaks generate turbulence and currents.
or biconcave-shaped (i.e. cylindrical shape with medial Regular, symmetrical and erect shapes usually charac-
groove; Fig. 9l, m). Goldfuss (1826–1833) and de Loriol terize siliceous sponges living at greater depths
(1882–1884) suggested that these two types of columnals (Tabachnick, 1991; Finks, 2003). These morphotypes
may belong to Eugeniacrinites and Cyrtocrinus, respec- probably represent the most adequate adaptive response
tively. The only cyrtocrinid holdfasts (Fig. 10d) from the of sponges to living in habitats characterized by high
Chénier Ravine are those attached to siliceous sponges sedimentation rate and soft bottom sediments. Recent
(Fig. 6n1). They clearly indicate that cyrtocrinids had siliceous sponges obviously adopt erect shapes to raise
ecological preferences for hard substrates. Although very their body high above the substrate and to maintain their
rare, typical primibrachials (ossicles of proximal series of aquiferous system away from turbid or flocculent layers.
brachials) of cyrtocrinids displaying verrucose external Moreover, tubular morphologies allow to separate in-
convex surfaces (Fig. 9d) were recognized among the coming and outcoming water systems thus protecting
ossicle assemblages. Lonchocrinus sp. that bears peculiar sponges from ingesting their own wastes in calm water
axillary primibrachials (brachial supporting two arm settings (Trammer, 1983). Hexactinellids often anchor
branches), with a long spiny process overhanging the to soft sediment by means of a single long siliceous
secundibrachial rows, is also present (Fig. 9b, c). For spicule (e.g. Hyalonema, see Roux et al., 1991b, pl. IV,
Jaekel (1907), this feature typifies the genus Lonchocrinus Fig. 6) or a tuft of spicules (Roux et al., 1991b, Fig. 2).
whereas de Loriol (1882–1884) considered these brachial However, if certain hexactinellids do live on soft,
ossicles as merely a characteristic of Eugeniacrinites. muddy substrate, many species prefer firmer grounds
such as fragmentary shells or hard substrates such as
5. Discussion rocks (Tabachnick, 1991).

5.1. Recent deep-sea siliceous sponges and crinoids 5.1.2. Cyrtocrinid crinoids and other deep-sea crinoids
The stalked crinoid fauna of New Caledonia (France,
5.1.1. Siliceous sponges SW Pacific) has long been considered as the most
Numerous aspects of the biology and ecology of archaic of all Recent oceans because of close similarities
siliceous sponges remain obscure to us partly due to with the Mesozoic crinoid fauna of the Tethyan Ocean
their deep-sea habitat that allows only brief observations (Améziane-Cominardi et al., 1987; Bourseau et al.,
via remote controlled cameras or submarine dives. 1988; Améziane and Roux, 1997). This is confirmed by
Hexactinellids range from 100 to ca. 6000 m but remain other faunal elements such as Nautilus, pleurotomarian
rare at depths between 100 and 200 m. They reach a gastropods, sponges, brachiopods, and echinoderms
peak of numerical abundance and diversity between 200 (Améziane-Cominardi et al., 1987; Roux et al., 1991b).
and 2000 m, in the bathyal zone beyond the continental Cyrtocrinids form an extant group of crinoids with
shelf (Sara and Vacelet, 1973; Leys and Lauzon, 1998; well-documented Jurassic ancestors (Manni and Nicosia,
230 S. Charbonnier et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 250 (2007) 216–236

1996; Ausich et al., 1999b). Although scarce in the off the Philippines Islands (South China Sea) at depths
Lower Jurassic, the group becomes more diverse and exceeding 900 m, (Bourseau et al., 1991), displays
abundant in the Middle Jurassic and is represented by striking similarities with some Mesozoic ancestors of
seven Recent species distributed in four genera: Gym- the isocrinid representatives such as Balanocrinus
nocrinus, Holopus, Cyathidium and, Proeudesicrinus which has been used as an indicator of deep-sea palaeo-
(Bourseau et al., 1991). environment (e.g. Roux et al., 1988; Bourseau et al.,
Key-information concerning the biology and ecology 1988, 1998). Based on oceanographic expedition
of modern cyrtocrinids was obtained in the 1980s owing results, Roux et al. (2006) noted that the minimal
to successive oceanographic expeditions by French sci- depth for Recent balanocrinids (e.g. Neocrinus decorus
entists along the bathyal slope of New Caledonia (SW (Meyer et al., 1978)) is around 250 m, with species
Pacific, Bourseau et al., 1991). These expeditions led to living deeper down to 400 m.
the discovery of a rich cyrtocrinid fauna living at depths
between 300 and 500 m. Submarine dives (Roux et al., 5.2. Palaeobathymetry and palaeosubstrate of the La
1991a,b) made it possible to observe these crinoids in Voulte area: evidence from sponges and crinoids
their natural habitat for the first time. For example,
Gymnocrinus richeri (Bourseau et al., 1987; Fig. 8f) is a 5.2.1. Sponges
small, robust cyrtocrinid that lives attached to hard- A series of fossil evidence supported by detailed
grounds on the top of very abrupt seamounts (Norfolk comparisons with modern and fossil morphological and
Rise, Loyalty Rise, Richer de Forges et al., 1986; Richer ecological analogues underpins the notion that the sponge
de Forges and Laboute, 1989; Bourseau et al., 1991) communities of the Chénier Ravine once flourished
where sediment deposition is moderate and currents in dysphotic or aphotic, relatively deep, possibly bathyal
relatively strong. These studies also revealed extremely settings. These are:
close morphological similarities between Recent and
Mesozoic cyrtocrinids. G. richeri is morphologically (1) the dominance of hexactinellids (80.1%) that, in
close to Cyrtocrinus (Fig. 8c, f) and Hemicrinus from Recent marine environments, characterizes deeper
the Jurassic and would indeed deserve the status of water conditions;
“living fossil”. The cyrtocrinid fauna from New (2) the presence of lithistids and especially rhizomor-
Caledonia also confirms the ecological preference of ines whose modern representatives occur mostly
the group for deeper water settings, strong slopes (ca. in the bathyal zone;
20°), hard substrates (moderate sedimentation rate) and (3) the prevalence of conical shapes erect morphol-
high energy (Roux et al., 1991b), all ecological ogies (narrow tubular: 28.8%; conical: 16.3%)
characteristics that had long been suspected by genera- that typifies most of modern deep-sea sponges;
tions of palaeontologists before Recent species could be (4) the absence of low and flat morphotypes that
observed in situ. Monospecific populations of echino- usually thrived in Upper Jurassic circalittoral
derms also frequently colonize the bathyal abrupt slopes environments (Gaillard, 1983; Beresi, 2003);
of New Caledonia (Roux et al., 1991b). Cidarid sea (5) the lack of photophilic organisms (microbialites)
urchins, probably with similar tolerance to physical encrusting the upper part of the external wall of
parameters, tend to occupy the same ecological niches sponges and the lack of sciophilous (=capable of
as those of cyrtocrinids and, in some cases, may replace thriving in shade) epizoans (e.g. bryozoans,
them (Roux et al., 1991b). It is worth noting that a thecideid brachiopods) in parts facing down-
comparable situation exists in the Callovian marls of the wards. This feature does not indicate a precise
Chénier ravine (see Thiéry et al., 1928) where both bathymetrical range but is consistent with dys-
cyrtocrinid and cidarid remains co-occur in skeletal photic or aphotic conditions. The benthic envi-
assemblages. ronment of the Chénier Ravine was obviously
Other crinoids occur in the deep-sea. Among Recent much deeper than the circalittoral zone and may
crinoids, the Bathycrinidae live at greatest depths. For correspond to the slope-basin transition probably
example, Naumachocrinus was collected at depths in the upper bathyal zone (ca. 200–400 m).
between 800 and 1000 m (Bourseau et al., 1991;
Améziane and Roux, 1997). Rasmussen (1978) and The mode of attachment of certain sponges (those
Bourseau et al. (1991) suggested that Dolichocrinus with an encrusting base on crystalline rock) provides
might be the oldest representative of the family Bath- some clues concerning the bottom conditions that
ycrinidae. Similarly, Hypalocrinus naresianus, collected once prevailed in the Chénier Ravine. It suggests the
 S. Charbonnier et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 250 (2007) 216–236 231

existence of both submarine exposures of crystalline suggested, from the brachiopod fauna and sedimento-
rocks and muddy areas where the sponges were pre- logical analyses, that the palaeoenvironment at La Pouza
served. The crystalline exposures may find their origin was relatively deep (possibly a slope setting at ca. 150 m
in the major fault of La Voulte and the fault complex of depth). The Middle Oxfordian sponge facies from the
associated with it. The fault system is likely to have Swiss Jura and Swabian Jura are also rich in cyrtocrinid
played a key-role in creating a sub-marine palaeotopo- crinoids (Quenstedt, 1874–1876; de Loriol, 1877–
graphy with substantial breaks and steep reliefs that 1879; Hess, 1975). Evidence from facies associated
favoured the colonization by siliceous sponges. Such with sponge bioherms (Ausich et al., 1999b) points to a
substrate and topographic heterogeneity is typical of the bathymetrical range possibly exceeding 100 m. The
Recent bathyal slope environments that are character- Middle Tithonian of Rogoznik (Pieniny Klippen Belt,
ized by a relatively steep relief, and are scoured by Poland) also yielded a remarkable cyrtocrinid fauna in a
currents and mudflows (Roux et al., 1991a). depositional environment that Pisera and Dzik (1979)
and Ausich et al. (1999b) view as corresponding to the
5.2.2. Crinoids top of a seamount below the wave base. The most
Recent crinoid morphology and distribution (see famous cyrtocrinids are probably those of the Stramberg
previous chapter) provide compelling evidence that the fauna (Valanginian, Czech Republic) first described by
crinoids from the Chénier Ravine lived in relatively Jaekel (1891). This rich fauna occurs in reddish cal-
deep-sea habitats (at least exceeding 200 m). These are: careous mudstones that form in-fillings along the eroded
surface of Tithonian limestones. Žítt (1983) interpreted
(1) the dominance of cyrtocrinids almost identical to this facies as the irregular surface of palaeo-seamounts.
the Recent bathyal representatives of the group; Ausich et al. (1999b) hypothesized that the assumed
(2) the presence of Dolichocrinus and Balanocrinus creviced bottoms of the Stramberg Sea offered optimal
that closely resemble typical Recent deep-sea habitats for the settlement of cyrtocrinids and compared
crinoids (the Bathycrinidae and the Isocrinidae this palaeoenvironment with Recent Pacific seamounts
Balanocrininae, respectively). colonized by G. richeri. Other fossil cyrtocrinid faunas
from Italy, Hungary and Turkey are also known in
Fossil evidence from cyrtocrinid occurrences also similar palaeoenvironments located on structural heights
gives support to our interpretation. This group is well with hardgrounds swept by currents (see Manni and
documented in the Mesozoic of Europe (see examples in Nicosia, 1996 for references).
Table 5). One of the richest cyrtocrinid faunas comes D. aberrans is present at La Pouza (Lower Bathonian,
from La Pouza (Lower Bathonian, SE France, Dumor- France, de Loriol, 1882–1884) associated with numer-
tier, 1871; Ausich and Simms, 1999; Ausich et al., ous cyrtocrinids and also occurs in the French “Terres
1999b) close to La Voulte (Fig. 1). This locality yielded Noires” Formation in a facies consistent with deeper
numerous brachiopods and undetermined siliceous water conditions (Rolin et al., 1990). Numerous speci-
sponges (Elmi, 1967). Alméras and Elmi (1996) mens were indeed collected (CG) near the carbonate

Table 5
Examples of Mesozoic cyrtocrinid occurrences
Outcrops (1) La Pouza, France (2) Swiss Jura, Swabian Jura (3) Rogoznik, Poland (4) Stramberg, Czech Republic
Age Lower Bathonian Middle Oxfordian Middle Tithonian Valanginian
Cyrtocrinid faunas Cyrtocrinus Cyrtocrinus Apsidocrinus Cyrtocrinus
Dolichocrinus Dolichocrinus Eudesicrinus Eugeniacrinites
Eugeniacrinites Eugeniacrinites Hemicrinus Hemibrachiocrinus
Gammarocrinites Pilocrinus Lonchocrinus Hemicrinus
Lonchocrinus Tetracrinus Phyllocrinus Phyllocrinus
Phyllocrinus Psalidocrinus Sclerocrinus
Sclerocrinus Torynocrinus
Tetanocrinus
Palaeoenvironment steep slope sponge bioherms seamont top seamount top
Palaeobathymetry about 150 m 100 m or more important important
References Elmi (1967) Ausich et al. (1999) Pisera and Dzik (1979) Žítt (1983)
Alméras and Elmi (1996) Ausich and Simms (1999)
(references in the text).
232 S. Charbonnier et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 250 (2007) 216–236

structures (with associated benthic fauna) previously raphy of the Callovian palaeo-slope. Siliceous sponges
interpreted as sponge bioherms and now regarded as and cyrtocrinids were most probably inhabiting this type
induced by hydrothermal springs located along the main of environment because it provided them appropriate
faults of a passive margin (Gaillard et al., 1985; Rolin hydrodynamic conditions and adequate substrates. In
et al., 1990). addition, this fault-controlled escarpment may have
C. nutans that dominates the crinoid assemblages of generated local sedimentary slides leading to parau-
the Chénier Ravine, may have occupied, along with tochthonous burial of sponges and crinoids. This setting
others robust cyrtocrinids, habitats and ecological was certainly favourable to numerous others organisms
niches comparable with those of the Recent G. richeri present in the Chénier Ravine such as sea stars,
(relatively small endemic populations colonizing bathy- brachiopods, bivalves, ammonites or, belemnites. The
al seamounts and hardgrounds where energy is high). block to which the present-day La Voulte Lagerstätte
belongs, possibly benefited from local specific condi-
6. Implications for the palaeobathymetry of the La tions that allowed the exceptional preservation of a vast
Voulte Lagerstätte array of organisms (e.g. deep-sea cephalopods, varied
crustaceans, ophiuroids, sea stars and sharks). These
The double set of convergent evidence shown by favourable conditions may have been induced by the
siliceous sponges and crinoids allows an accurate vicinity of hydrothermal vents. Although direct evi-
assessment of the bathymetry of the Chénier Ravine dence for such vents has not been found yet, the pre-
and by extension of the La Voulte area during the sence of substantial iron deposits clearly related to the
Callovian. The La Voulte area most likely laid under a activity of the La Voulte fault brings some support to the
substantial water depth, exceeding 200 m (most hypothesis of hydrothermal activity. Moreover, many of
probably in the upper bathyal zone) around the slope- the minerals (e.g. pyrite, siderite) preserving the La
basin transition. The distinctive submarine topography Voulte fossils are often present at marine hydrothermal
that characterizes this particular domain in Recent settings (see Haymon et al., 1984; Oudin and Con-
oceans (heterogeneous substrates, active faulting, steep stantinou, 1984; Banks, 1985). The mineralization of the
relief with blocks, currents) was likely to have been soft-bodied organisms was obviously rapid and suggests
present in the La Voulte area. This has important that unusually high concentrations of dissolved metals,
bearings on the palaeoenvironmental reconstruction of phosphates, carbonates, sulfates, and sulfides were
the associated La Voulte Lagerstätte that, taken by itself, immediately available (Wilby et al., 1996). Rapid
lacks precise sedimentological and faunal criteria. This interaction between the sediment and the carcasses
first attempt to characterize its palaeoenvironmental promoted early diagenetic mineralization, probably
setting and to tentatively place it within the bathyal zone mediated by bacteria (Wilby et al., 1996). The fossil
is consistent with the faunal composition of the La assemblages from the Lagerstätte and the Chénier
Voulte Lagerstätte. Indeed, its fauna contains vampire Ravine are virtually contemporaneous and have a
squids (Fischer and Riou, 2002), cirrate octopods similar bathymetry. Their major differences may be
(Fischer and Riou, 1982a), sea spiders, great span sea explained by the occurrence of hydrothermal activity in
stars (Fischer, 2003) that are all indicators of relatively the vicinity of the Lagerstätte area that created con-
deep-sea conditions (Gage and Tyler, 1991). The ditions favourable to: (1) the settlement of a specific
enigmatic thylacocephalans (Secrétan, 1985; Vannier community of organisms and, (2) the exceptional
et al., 2006) bearing hypertrophied visual organs preservation of the fauna. These conditions were most
resembling those of deep-sea crustaceans, if not direct probably absent in the Chénier Ravine and allowed
indicators of palaeobathymetry, clearly point to the neither the colonization of the area by the same organ-
dysphotic or aphotic conditions. isms as those from the Lagerstätte, nor soft-bodied
A tentative reconstruction of the La Voulte area is preservation. Another possible effect of the hydrother-
proposed (Fig. 11) that shows the assumed relations mal activity on the marine fauna is the high concentra-
between the La Voulte Lagerstätte and the Chénier tion of flocculent metal minerals that possibly inhibited
Ravine. The overall submarine topography is seen as the development of sponges and crinoids in the
being dominated by steep blocks generated and con- Lagerstätte area.
trolled by the La Voulte fault and its cortege of Concerning the palaeobathymetry, Etter (2002b)'s
transverse faults (e.g. Col de Viau fault separating the view of a palaeoenvironment situated slightly below the
Chénier Ravine from the Lagerstätte). The La Voulte storm wave base is probably underestimated. Fischer
fault appears as a major break in the submarine topog- (2003)'s interpretation of a bathyal palaeoenvironment
 S. Charbonnier et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 250 (2007) 216–236 233

Fig. 11. Palaeoenvironmental reconstruction of the La Voulte area during the Lower Callovian. (1) Chénier Ravine with siliceous sponges and
crinoids (black triangles, circles, stars = assumed natural habitat of sponges and crinoids); (2) La Voulte Lagerstätte with diverse crustaceans,
echinoderms and, soft-bodied cephalopods. Note steep topography (tilted blocks) generated by local faults.

at La Voulte may be correct but his “Sargasso Sea consistent with those of Elmi (1990) and Alméras and
model” is not supported by existing evidence. The Elmi (1996). If our interpretations are correct, the La
hypothesis of a coastal lagoon (Alessandrello et al., Voulte Lagerstätte may be one of the rare and extremely
2004), which obviously does not take geological precious Lagerstätten to have fossilized a deep marine
evidence into account, is rejected. Our results are fauna.
234 S. Charbonnier et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 250 (2007) 216–236

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nie. In: Crosnier, A. (Ed.), Résultats des Campagnes MUSORS-
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Kimméridgien inférieur de La Rochelle (Charente-Maritime,
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