GEOLOGICA CARPATHICA, 51, 2, BRATISLAVA, APRIL 2000
ROVEACRINUS BERTHOUI NOV. SP., EARLY HAUTERIVIAN
REPRESENTATIVE OF ROVEACRINIDAE (ROVEACRINIDA,
CRINOIDEA) OF BUSOT (ALICANTE, SPAIN)
and BRUNO GRANIER
2 rue Guy de Maupassant, F-76800 Saint Étienne du Rouvray, France; firstname.lastname@example.org.
TOTAL Scientific and Technical Center, TEP/DE/CST/G-CARB, Domaine de Beauplan, Route de Versailles,
F-78470 Saint Rémy lès Chevreuse, France
(Manuscript received December 3, 1998; accepted in revised form December 12, 1999)
Abstract: A new roveacrinid microcrinoid, Roveacrinus berthoui nov. sp., is described from the Lower Hauterivian
marly limestones at Busot (Alicante, SE Spain). It represents the first record of this family from pre-Albian strata, and as
such, constitutes a major stratigraphic and taxonomic advance in our knowledge of this unusual fossil crinoid group.
Key words: Cretaceous, Hauterivian, Spain, Alicante, Echinodermata, Roveacrinida, Roveacrinidae.
In the province of Alicante (SE Spain), Mesozoic series crop
out due to structural anomalies referred to extrusions à
noyau dur (Polvêche 1963). In the Prebetic and Subbetic
domains, the Hauterivian series are usually incomplete and
condensed, in contrast to the relatively thick series devel-
oped at Busot (Fig. 1; Granier 1993). This is explained by the
peculiar position of this locality at the edge of the Berriasian
platform. During the Early Valanginian, an important trans-
gressive wedge related to the platform flexure and subse-
quent drowning developed at the edge, on the slope basin-
wards and far beyond (Busot, Fig. 2). During the Late
ValanginanHauterivian interval, the initiation of block-tilt-
ing induced condensation and/or erosion phenomena on the
one hand, particularly on local highs, and sediment trapping
in grabens and/or half-grabens on the other (Fig. 2). Such an
environment favoured the preservation of Hauterivian de-
posits at Busot. Twenty kilometres NNE of Alicante, the
village of Busot is located at the SW end of a Mesozoic belt
called bande à anomalies structurales de Busot-Altea
(Granier 1987). There, Lower Cretaceous (Valanginian to
Aptian) deposits crop out within a hardcore extrusion com-
plex. South of the village, two Cretaceous hills consist of
Hauterivian marly limestones. These bioclastic carbonates
with wackestone texture yield abundant small benthic fora-
minifers, echinoderm fragments, ammonites and calpionel-
lids as autochthonous assemblage components. They also
contain reworked, worn and micritized microfossils, mostly
large benthic foraminifers and calcareous algae. However,
reworking is limited and neither slumps nor conglomerates
have yet been documented.
While the wackestone marly limestone displays a micritic
matrix, the high-magnesium calcite (HMC) network of the
echinoderm ossicles does not appear to have been affected by
overgrowth or syntaxial cement. If micritization or corrosion
of the echinoderm ossicles should occur, it is only moderate
and does not alter noticeably the morphology of the HMC
section considered below, nor its subsequent reconstruction.
No additional ossicles were found in the numerous thin
sections processed to further document this new species. De-
structive petrophysical measurements on the original sub-
sample made any further research of conspecific plates by se-
rial sections impossible. Our attempt to extract additional
material from the original field sample was unsuccessful: the
washing residues were devoid of any other thecal plate
*Present address: PDD/GEOL 7W4, ADMA-OPCO, P.O. Box 303, Abu Dhabi, United Arab Emirates; email@example.com
Fig. 1. Map showing the location of the study area (SE Spain).
Fig. 2. Tectonic cross-section through the Busot-Altea hardcore extrusion complex reconstructed from field-section observations (from
Granier et al. 1995).
Fig. 3. Lithology, chronostratigraphy and ammonite occurrences in section Busot III, southeast of Busot, Alicante-Spain (adapted from
Granier et al. 1995).
ROVEACRINUS BERTHOUI NOV. SP., HAUTERIVIAN ROVEACRINIDAE OF BUSOT 103
while, amongst the echinoderm ossicles, some rare rove-
acrinid plates could not be identified to species.
The present study describes one of these echinoderm
components as the oldest stratigraphic record of the family
Roveacrinidae to date, sections of which have commonly
been referred erroneously to as the Lombardia-microfacies
(Brönnimann 1955; Verniory 1956) or as Cretaceous sacco-
comids (for a review see Ferré 1997).
Morphological terminology follow Rasmussen (1961) and
Scott et al. (1977). For the orientation of section planes and
description of thin sections (Fig. 4), the microfaciological ter-
minology of Ferré & Berthou (1994) has recently been re-
fined and completed by Ferré & Granier (1997a, in press).
Order Roveacrinida Sieverts-Doreck, in Moore et al. 1952
Family Roveacrinidae Peck 1943 (emend. Rasmussen 1961)
?Subfamily Roveacrininae Peck 1943 (emend. Peck, in
Ubaghs et al. 1978)
Genus Roveacrinus Douglas 1908
(=Drepanocrinus Jaekel 1918)
Type species Roveacrinus communis Douglas 1908
(=Drepanocrinus sessilis Jaekel 1918 =Roveacrinus
westphalicus Sieverts 1933)
Roveacrinus berthoui nov. sp.
Pl. I: Figs. 14
1995 Rovéacrinidé: Granier et al. Fig. 4.3.
1997b Roveacrinus berthoui nov. sp. Ferré & Granier: p. 338339
Holotype: By monotypy is Thin section No. AL 945, in
the Office National de Gestion des Collections Paléon-
tologiques (O.N.G.C.P., Villeurbanne, France) under registra-
tion number FSL 411 599, the original negative film being la-
belled nb 07730A.
Material: Type specimen only (a unique oblique medial
Derivatio nominis: Dedicated to the late Dr. Pierre-Yves
Berthou (Université Pierre and Marie CurieParis VI) for his
outstanding contribution to our knowledge of the roveacrinid
Type locality: Section Busot III (Granier et al. 1995), south-
east hill of Busot (Alicante, Spain).
Type level: Bed 945, bioclastic wackestone carbonate
bed, Los Villares Formation (Vera et al. 1982), Lower Hau-
terivian (Fig. 3).
Age: Early Hauterivian, Angulicostata Zone, Tintinnopsel-
la Zone (sensu Pop 1980).
Dimension: Standard measurements cannot be taken. Ap-
proximate thecal width and height: ca. 550
m and ca. 650
m, respectively; H/W ratio: ca. 1.181.23.
Diagnosis: Theca medium to small, width equalling height,
subconical to rounded, almost pentagonal to rounded from
basal view. No subdivision between radials and basals, nor
Fig. 4. Orientation of sections shown on a reconstructed rove-
acrinid (from Ferré & Berthou 1994; additional data from Ferré &
Granier 1997 in press).
prominent constriction between dorsal and ventral cavities,
nor interradial recess visible. Theca deeply indented between
radial facets, which are large and slope outwards. Dorsal liga-
ment pits thick and crescentic, nerve channels medium-sized
to small. Interradial pegs without any relief. Radials strongly
ornamented by a plume or acanthus-leaf pattern proceeding
upwards from the polar edge and continuing to nearly up to the
radial facets. Secondary plume flanges branching off along
each side of the primary radial edge, and merging aborally to
the polar edge. These secondary flanges grow laterally and
slightly outwards, their bases maintaining distal connections
with the main flange. Lateral flanges from adjoining rays nev-
er coalesce or efface. No interradial processes appear above
the general edge of the calyx. Ventral cavity completely en-
cased in the radial frame. Thecal microstructure of dense
HMC. Arm structure, plating and ornament unknown, but
most likely to be of the same type as thecal ornament.
Description: This HMC section unquestionably represents
a roveacrinid crinoid (Ferré 1997) since a ventro-dorsal
stretching appears from the geometrical reconstruction. In-
deed, this oblique section in part shows an elongated ventro-
dorsal theca which corroborates attribution to the family
Roveacrinidae. For the time being, a possible assignment to
the subfamily Roveacrininae is speculative, since there is no
evidence of an additional inner set of plates (Schneider 1987,
1989). However, there is no evidence of their absolute absence
eitherthey may have fallen victim to post-mortem decay or
during transport as they are neither fused nor articulated to the
Despite the section being oblique, we may estimate the
cavity ratio: the dorsal cavity is slightly larger. Although two
tiny indentations formed by the upper side of the basals (cav-
ity wall) appear on the inner part of the section, indicating
the presence of an inner partition, we cannot determine
whether there is an additional set of plates defining cryptodi-
cyclicity as defined by Schneider (1987). The dorsal cavity is
rather large compared to the radial expansions preserved in
this oblique plane, whilst the ventral cavity is rather de-
pressed. The dorsal cup is rather smooth, if we consider it to
be encased in the radials and the weak radial ornament effac-
ing at the distal point of the basals.
The pentamerous symmetry induces irregular ridges and
small flanges all around the theca to represent the respective
set of five radials. Only two of them are available for full
examination. Radial 2 (Pl. I: Fig. 4) is preserved articulated
to and thus covered by the first primibrach: its articular fac-
et is displayed in full in the oblique section at hand; thus the
radial articular facet must have been oblique. Consequently,
the preserved radial expansions and the associated first
primibrach document the outward sloping of the articular
facet, showing the specimen at hand to be a representative of
the genus Roveacrinus.
The section area featuring Radial 1 (Pl. I: Fig. 4) shows a
twisted ornament pattern. Due to the section obliquity and the
radial position, this undulated frame must be interpreted as in-
dependent parallel flanges. These flanges, oblique to the radial
vertical ridge, branch off along this very same vertical radial
ridge. Geometrically, these oblique ridges occur just below the
radial facet and continue down to the thecal base. They form a
plume pattern which shows up as wavy ornament in oblique
section; their proximal sides join weakly to the polar edge to
result in a relatively smooth lower theca. There is no interradi-
al ornament nor obvious secondary ornament on the radials,
apart from the plume pattern appearing in between the radial
Unfortunately, our material does not contain any brachial
ossicle section associated with the thecal remains, with the
exception of the partial oblique section of the first primi-
brach connected to Radial 2. Based on that, as well as on a
specimen from the Sergipe (Ferré & Bengtson 1997) and ma-
terial from Texas (Scott et al. 1977), we may infer a similar,
though less marked, ornament on brachial plates, i.e. either a
coarse, rugose primibrach pattern and a smooth pinnule pat-
tern, or a granulated brachial frame.
Discussion: The short twisted aboral knob, the plume radi-
al ornament, as well as (to a lesser extent) the stratigraphical
position, characterize this new find, and justify the erection
of a new species of the genus Roveacrinus Douglas. With
reference to the radial expansions preserved in this oblique
section, we may infer a rather heavy ornamentation and
strong arms consisting of large brachial plates. The ventral
(or brachial or adoral) cavity is rather depressed compared to
the dorsal cavity, explaining the relatively high dorsal/ven-
tral cavity ratio.
The inferred marked ornament and the general low form of
this theca rule out a benthic inhabitant of agitated shallow
waters. This ecological assignment is based on observations
of a complete Roveacrinus specimen (Ferré & Bengtson
1997) which is consistent with microfacies analyses. It has
now become clear that the worldwide occurrence of rove-
acrinid microfacies and the light structure of their skeletons
no longer support their previously widely accepted plank-
tonic mode of life (Milsom 1989, 1999). These arguments
were based exclusively on the early planktonic stages that
characterize any echinoderm group. Therefore under favour-
able environmental conditions such as hypoxic high-produc-
tivity events, roveacrinids produced opportunistic broods
which spread over the whole Tethys; then during a later
stage, these broods sunk to the seafloor to become benthic
suspension-feeding adult colonies. Thecal structure (mostly
proportions and ornament), allows us to assume a rather shal-
low, open-marine environment since the accretion of the ra-
dials covering the dorsal cavity might have been related to
the sinking of planktonic roveacrinid brood on the seafloor.
This environment is consistent with the reconstruction elabo-
rated by Granier (1987).
Although it displays the typical roveacrinid thecal stretch-
ing, its low profile recalls Tithonian Saccocomidae. Howev-
er, in the present state of knowledge, much has to be still
done to make Roveacrinus berthoui, nov. sp., the link from
classic Jurassic Saccocomidae to Cretaceous Roveacrin-
idae. Finally, with regard to the geographical location,
Busot lay at the crossroads of Tethyan seaways. Hence, by
promoting the development of roveacrinid populations
(comparable to Triassic Osteocrinus-facies; see Kristan-
Tollmann 1970), the central Tethys might have actually
been at the root of Cretaceous Saccocomidae (Peck 1973),
as well as fostering a new strain towards Roveacrinidae
(Ferré et al. 1997).
Age: The senior author has identified in thin sections Tin-
tinnopsella carpathica Murgeanu & Filipescu and T. longa
(Colom) indicative of the Tintinnopsella Zone (Pop 1980).
The ammonite assemblage at the Busot locality comprises
Olcostephanus sp., Pseudothurmannia grandis Busnardo,
Pseudothurmannia cf. pseudomalbosi Sarasin & Schöndel-
mayer and Pseudothurmannia sp. (identifications by R.
Busnardo & L.G. Bulot), dating this level as Early Hauteriv-
ian (Angulicostata Zone).
Assemblage: As stated above, this part of the section yield-
ed an unusual faunal assemblage of both ammonites and calpi-
onellids (Granier et al. 1995). The sedimentary texture of this
marly carbonate is a bioclastic wackestone containing numer-
ous benthic foraminifers (Neotrocholina sp., Trocholina gr. al-
pina (Leupold), Montsalevia salevensis (Charollais, Brönni-
Plate I: Roveacrinus berthoui nov. sp., Early Hauterivian, Anguli-
costata Zone, Tintinnopsella Zone, Busot III, Alicante, Spain. Fig.
1. Holotype (ca.
120). Fig. 2. Interpretative line drawing of the ho-
lotype section. Scale bar: ca. 250 µm. Fig. 3. Tentative reconstruc-
tion of the theca. The dihedrals indicate the plane section at hand.
Fig. 4. Simplified version of 2. CW cavity wall separating dorsal
and ventral cavities; Rad Exp radial expansion showing the radi-
al edge cut slightly by the oblique plane section; RAF radial ar-
ticular facet of the first primibrach as preserved articulated to radial
PLATE I 105
mann & Zaninetti, Lituolidae, and others), various echinoderm
ossicles (starfish, ophiuroid and crinoid) and reworked calcar-
eous algae (among others Macroporella? praturloni Dragastan
and Pseudocymopolia pluricellata Bakalova), with additional
silt-size detrital quartz grains.
Stratigraphical distribution: Roveacrinus berthoui nov.
sp. represents the oldest member of the family Roveacrinidae
known to date, previous records involving Early Albian forms
(Dias-Brito & Ferré 1997, in press) and thus fills the gap be-
tween the Late Tithonian Saccocomidae and the latter Early-
Middle Albian Roveacrinidae. As the stratigraphic gap is pro-
gressively being closed, the new species once again calls into
question the previously assumed relationships between these
two families (for a review see Ferré 1997).
To date, the specimen studied represents the oldest known
representative of the family Roveacrinidae, previously re-
corded from the Early Albian (Dias-Brito & Ferré 1997, in
press) and the MiddleLate Albian (Peck 1943, 1955; Ras-
mussen 1961; Schmid 1971; Jäger 1981; Destombes 1984;
Griffiths 1989; Ferré & Granier 1997, in press) to the Late
Maastrichtian (Jagt 1992, 1999). It fills the gap between the
Late Tithonian Saccocomidae, the dubious Microcalam-
oides diversus Bonet 1956 (see Ferré 1997) and the later Ear-
lyMiddle Albian Roveacrinidae. However, additional mate-
rial (particularly isolated calyces, or at least orthogonal/axial
thecal sections) will be needed to refine our present diagno-
sis and a more definitive phyletic statement.
As the stratigraphical gap between these sister groups is
closing, the new material again casts doubts on the phyletic
relationships between Saccocomidae and Roveacrinidae,
and puts a much earlier date on the branching if these fami-
lies are really related. Furthermore, from a paleogeographic
point of view, as Kristan-Tollmann (1970) promoted the Os-
teocrinus-facies in the Late Triassic Tethys Ocean, this
new material stresses the role played by the Tethyan seaway
in the appraisal of Cretaceous roveacrinid microfacies
(Dias-Brito & Ferré 1997, in press; Ferré et al. 1997).
The presence of identifiable diagnostic roveacrinid re-
mains in thin sections greatly enhances the chronostrati-
graphical value of roveacrinid microfacies.
The previously established framework of roveacrinid mi-
crofacies, still being refined, offers a means of rapid and re-
liable identification of local stages in the Tethyan Realm and
beyond (Gulf of Mexico, Brazilian and West African mar-
ginal basins). The relative proportions and the occurrence of
roveacrinid taxa appear to be of supraregional chronostrati-
Acknowledgements: Both support and permission to publish
of TOTAL are gratefully acknowledged. Drs J.M. Villain, D.
Michoux (TOTAL Scientific and Technical Center, France),
É. Fourcade and P. Cros (CNRS-ESA7073, Université P. and
M. CurieParis VI, France) provided useful advice and con-
structive comments on a preliminary draft. We extend our
warm thanks to Prof. M. Miík (University of Bratislava, Slo-
vakia) and two anonymous reviewers for their comments and
suggestions. We are greatly indebted to Drs. R. Busnardo
(Université Claude Bernard-Lyon I, France) and L.G. Bulot
(Université de Provence, Marseille, France) for ammonite
identifications. The bibliographical assistance of Mrs. F.
Ozanne (Librarian, Société Géologique de France, Paris,
France) and of Ms M.A. Lançon (Data Manager, Université P.
and M. CurieParis VI, France) is gratefully acknowledged.
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