Jurassic biostratigraphy and paleoenvironmental evolution of the ...

Abstract
Jurassic biostratigraphy and paleoenvironmental evolution
of the Malaguide complex from Sierra
Espuña (Internal Betic Zone, SE Spain)
Biostratigraphie du Jurassique et évolution paléoenvironnementale
du complexe Malaguide de la Sierra Espuña
(Zone bétique interne, SE de l’Espagne)
Jesús E. Caracuel a, *, José Sandoval b , Manuel Martín-Martín a ,
Antonio Estévez-Rubio a , Iván Martín-Rojas a
a Departamento Ciencias de la Tierra y del Medio Ambiente, Universidad Alicante, Apdo 99, 03080 San Vicente del Raspeig, Alicante, Spain
b Departamento Estratigrafía y Paleontología, Universidad Granada, Avenida Fuentenueva s/n, 18002 Granada, Spain
Received 22 December 2003; accepted 28 September 2004
Available online 09 November 2005
Jurassic studies in the Internal Zones of the Betic Cordillera are scarce since this zone is composed mainly of pre-Jurassic metamorphic
rocks. Only the “Dorsal” and the Malaguide domains include fossiliferous Jurassic successions, as in Sierra Espuña (SE Spain), which is one
of the bigger and well-exposed Jurassic outcrops of the Internal Zones. Collected Ammonite assemblages update and improve the precision of
previous biostratigraphic data by the recognition of: the Domerian (= Upper Pliensbachian, in the Mediterranean Domain) Lavinianum (Cornacaldense
Subzone), Algovianum (Ragazzoni, Bertrandi, Accuratum and Levidorsatum Subzones) and Emaciatum (Solare and Elisa Subzones)
Zones; the Lower Toarcian Polymorphum and Serpentinum Zones; the Middle Toarcian, Bifrons and Gradata Zone; the Upper Toarcian
Reynesi Zone; the Lower/Upper Bajocian, the Lower Callovian Bullatus and Gracilis Zones; the Middle/Upper Oxfordian Transversarium,
Bifurcatus, Bimammatum and Planula Zones; and the Lower and Upper Kimmeridgian Platynota, Strombecki, Divisum and Beckeri Zones.
The paleoenvironmental evolution of the Malaguide Jurassic at Sierra Espuña shows similarities with other Mediterranean Tethyan paleomargins.
The biostratigraphic precision along with the litho- and biofacies analyses has enabled the interpretation that the Malaguide paleomargin
evolved as a passive margin, developing shallow carbonate platforms, until the Domerian (Lavinianum Zone). Then, the platform
broke up (Domerian, Lavinianum Zone–Upper Toarcian, Reynesi Zone) with the beginning of the rifting stage, beginning the development of
horst–graben systems and the coeval drowning of the area. This stage ended in the upper Lower Callovian (Gracilis Zone) to the Middle
Oxfordian (Transversarium Zone) interval, starting the drifting stage, which accentuated the horst–graben systems, leading to the deposition
of condensed nodular limestones in the raised sea bottom.
© 2005 Elsevier SAS. All rights reserved.
Résumé
Les études sur le Jurassique des Zones Internes de la Cordillère bétique sont très peu abondantes du fait que les Zones Internes sont
essentiellement composées de roches métamorphiques plus anciennes. Seuls, la Dorsale bétique et le Domaine Malaguide, où les roches sont
en général épargnées par le métamorphisme, comprennent des successions fossilifères jurassiques d’intérêt biostratigraphique. C’est le cas de
la Sierra Espuña où on peut observer un des affleurements les plus complets et mieux exposés des Zones Internes. Les associations d’ammonites
permettent d’améliorer et de préciser les données biostratigraphiques préalables. Nous avons reconnu les Zones à Lavinianum (sous-zone à
* Corresponding author.
E-mail address: jesus.caracuel@ua.es (J.E. Caracuel).
0016-6995/$ - see front matter © 2005 Elsevier SAS. All rights reserved.
doi:10.1016/j.geobios.2004.09.002
Geobios 39 (2006) 25–42
http://france.elsevier.com/direct/GEOBIO/

26 J.E. Caracuel et al. / Geobios 39 (2006) 25–42
Cornacaldense), Algovianum (sous-zones à Ragazzoni, Bertrandi, Accuratum et Levidorsatum) et Emaciatum (sous-zones à Solare et Elisa)
dans le Domérien; les zones à Polymorphum et Serpentinum dans le Toarcien inférieur ; les Zones à Bifrons et Gradata dans le Toarcien moyen
et la Zone à Reynesi dans le Toarcien supérieur ; la partie supérieur du Bajocien inférieur et le Bajocien supérieur ; le Callovien inférieur,
Zones à Bullatus et Gracilis ; les Zones à Transversarium, Bifurcatus, Bimammatum et Planula dans l’Oxfordien moyen/supérieur et les Zones
à Platynota, Strombecki, Divisum et Beckeri dans le Kimméridgien.
L’évolution de l’environnement du Jurassique Malaguide dans la Sierra Espuña montre beaucoup d’affinités avec les autres paléomarges
de la Téthys méditerranéenne. La précision biostratigraphique et l’analyse des lithofaciès et des biofaciès nous ont permis d’interpréter la
paléomarge Malaguide comme une paléomarge passive, avec le développement d’une plate-forme carbonatée sommaire jusqu’au Domérien
(Zone à Lavinianum). Ensuite eut lieu la fracturation de cette plate-forme (Domérien inférieur, Zone à Levinianum-Toarcien supérieur, Zone
à Reynesi) liée au commencement de la phase de « rifting », caractérisée par le développement d’un système de horsts et grabens et l’enfoncement
de cette région. L’extension se poursuit entre le Callovien inférieur (Zone à Gracilis) et l’Oxfordien Moyen (Zone à Transversarium), en
indiquant le début d’une phase de « drifting » qui aurait déclenché la réactivation du système de horsts et grabens, conduisant au dépôt de
calcaires nodulaires condensés dans les parties les plus élevées du fond marin.
© 2005 Elsevier SAS. All rights reserved.
Keywords: Jurassic; Ammonite biostratigraphy; Paleoenvironmental evolution; Internal Betic Zone; Malaguide Complex; Southeastern Spain
Mots clés : Jurassique ; Biostratigraphie des ammonites ; Évolution de l’environnement ; Zone Interne Bétique ; Complexe Malaguide ; Sud-Est de l’Espagne
1. Introduction
The Jurassic evolution of the Betic Cordillera took place
under distensive tectonic conditions related to Tethyan rifting.
Although, the rifting age has been considered quite similar
throughout the cordillera, controversy continues concerning
the synchronism versus diachronism for the pre-, synand
postrifting periods in the different domains into which
the Betic Cordillera has traditionally been divided: the Internal
and the External Zones.
The External Zones belong to the South Iberian Paleomargin
and consist of post-Triassic unmetamorphosed rocks.
Thus, the Jurassic paleogeographic evolution and the age of
the onset of the rifting (Carixian = Lower Pliensbachian), and
the post-rifting phase (boundary Middle/Upper Jurassic) has
been well-characterized (Vera, 1988). On the contrary, the
Internal Betic Zones belong to a microplate (Mesomediterranean
Terrain, Guerrera et al., 1993) derived from the North
African continental margin, which collided against the External
Zones during the Early Miocene. These zones are built up
by the stacking of four complexes (in order upwards): Nevadofilabride,
Alpujarride, Malaguide and “Dorsal”. The Nevadofilabride
and Alpujarride are composed mainly of Paleozoic
and Triassic (and older), metamorphic rocks, while the
Malaguide and “Dorsal” Complexes include unmetamorphosed
Paleozoic (Malaguide) and Meso-Cenozoic (Malaguide
and “Dorsal”) sedimentary rocks. In general, the scarcity
of Jurassic unmetamorphosed successions, together with the
intense tectonic activity, have hampered the dating of the main
Jurassic events in the Internal Zones.
The Malaguide is the uppermost Complex (and, consequently,
the most internal) of the Internal Zones, which outcrops
mainly from Malaga Province (westward) to Murcia
Province (eastward). In contrast to the other complexes, the
Malaguide Complex includes Jurassic sedimentary covers,
favoring the analysis of the Jurassic evolution of the Internal
Betic Zones. The Sierra Espuña area, in Murcia Province, is
probably the most extensive, and the best exposed Jurassic
outcrop belonging to the Malaguide in the Betic Cordillera.
Except for the pioneer work by Fallot (1945), the main
papers on the Jurassic of Sierra Espuña are from the 1960s
and 1970s (Peyre and Peyre, 1960; Mac Gillavry et al., 1963;
Navarro and Trigueros, 1963; Paquet, 1962, 1969; Geyer and
Hinkelbein, 1971, 1974; Kampchuur et al., 1974; Seyfried,
1978, among others). Almost no recent studies have been
made on the biostratigraphy and the paleogeographic evolution
of the Jurassic of Sierra Espuña. In the present work, we
analyze some classical Jurassic sections, together with new
sections, in order to update the biostratigraphic framework
and to approach the paleoenvironmental evolution of the
Jurassic Malaguide from Sierra Espuña. Moreover, this will
improve the knowledge of the Jurassic of the Internal Betic
Zones, in addition to providing a fuller understanding of the
External Betic Zones, and their relationships.
2. Geographical and geological setting
The Sierra Espuña area is located in Murcia Province,
accessible by road from the villages of Alhama and Totana,
from the south, and Mula from the north (Fig. 1). The Malaguide
outcropping area of Sierra Espuña bounds tectonically
with the Alpujarride Complex to the SE, and with the External
Betic Zones (Subbetic) to the NW. Laterally, the nearest
outcrop of Jurassic Malaguide in the region is located 40 km
westward, near Vélez Rubio in the Province of Almeria
(Castillón Fm., Geel, 1973).
The outcropping Malaguide Complex in Sierra Espuña is
composed of two tectonic units (Martín-Martín, 1996); Morrón
de Totana and Perona (Fig. 1). The Morrón de Totana
Unit is the footwall while the Perona Unit is the hanging wall,
which, paleogeographically came from a more proximal position
(Martín-Martín and Martín-Algarra, 1997). Both units
include a marine Jurassic sedimentary cover, although in the

Perona Unit only outcrops Liassic sediments and have a
reduced areal extension. In this unit, only one section was
selected for studying (Perona section; Fig. 1). In the Morrón
de Totana Unit, the Jurassic is well developed, outcropping
with lateral continuity along more than 12 km (Fig. 1). Here,
four sections were analyzed, these being spaced 2–5 km from
each other in an E-W direction (Malvariche, Tres Carrascas,
Prat Mayor and Morrón Chico sections; Fig. 1).
J.E. Caracuel et al. / Geobios 39 (2006) 25–42
Fig. 1. Geographical (upper) and Geological (lower) sketches with the location of the 5 studied sections at Sierra Espuña. Legend: 1. Undifferentiated Quaternary.
2. Eocene platform carbonates. 3. Lower Cretaceous marls and marly limestones. 4. Upper Jurassic marl, limestones and nodular limestones. 5. Domerian–
Lower Callovian cherty rhythmic limestones. 6. Liassic p.p. shallow water limestones. 7. Triassic/Liassic? Dolostones. 8. Triassic conglomerates and sandstones.
Fig. 1. Croquis géographique (supérieur) et géologique (inférieur) avec l’emplacement des 5 sections étudiées dans la Sierra Espuña. Légende : 1. Quaternaire
indifférencié. 2. Plate-forme carbonatée éocène. 3. Marnes et calcaires marneux du Crétacé inférieur. 4. Marnes, calcaires et calcaires noduleux du Jurassique
supérieur. 5. Calcaires rythmiques à silex du Domérien inférieur - Callovien. 6. Calcaires d’eaux peu profondes du Lias p.p. 7. Dolomies du Trias/Lias?. 8.
Conglomérats et sables du Trias.
3. State-of-the-art Jurassic Malaguide at Sierra Espuña
The first relevant data on the Jurassic from Sierra Espuña
come from Fallot (1929, 1945), who studied the Jurassic successions
at Morrón Chico and Prat Mayor. This author recognized,
within the carbonate succession of the platform, a
ferruginous oolith-rich interval (0.5–3 m thick) with late Liassic
ammonites (Dumortieria sp. and Pleydellia sp.). This
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28 J.E. Caracuel et al. / Geobios 39 (2006) 25–42
guide-level was first noted by Villasante (1912), who assigned
it a Lusitanian to Kimmeridgian age.
Later on, Peyre and Peyre (1960) and Navarro and
Trigueros (1963) fixed the age of the ferruginous oolith-rich
interval studied by Fallot (1945), based on the faunas (ammonites
and brachiopods) collected in the Prat Mayor section.
Peyre and Peyre (1960) recognized abundant ammonites of
the genera Lytoceras, Coeloceras, Reynesoceras, Arieticeras,
Harpoceras and Protogrammoceras, and brachiopods
(Spiriferina, Terebratula and Rhynchonella), and used
them to date the Middle Domerian (Domerian = Upper Pliensbachian,
in the Mediterranean Domain). No more macrofauna
was collected in the remaining Jurassic succession,
although the record of microfacies enriched in crinoids, foraminifers
(Involutina, Nodosaria and Lenticulina), Stomiosphaera
and Cadosina toward the upper part, make feasible
to assume the existence of the Middle and Upper Jurassic, as
in the section at Morrón Chico studied by Fallot (1945).
The synthesis by Paquet (1969), and precursor works
(Paquet, 1962), are the most exhaustive studies of the Jurassic
from Sierra Espuña, including the analysis of sections in
the Morrón de Totana and Perona tectonic units. According
to Peyre and Peyre (1960) and Paquet (1969) proposed a
Middle Domerian (Upper Pliensbachian) age for the ferruginous
oolith-rich level of the Liassic in the Prat Mayor section,
based on the record of Fuciniceras cf. curionii (Meneghini),
Protogrammoceras bassanii (Fucini), Arieticeras
bertrandi (Kilian) and Ar. fuccinii (Del Campana), among
others. In the Perona section, the same level was dated by
Paquet, using brachiopods as Lower Pliensbachian, leading
to the suggestion that the ferruginous oolith-rich interval might
be diachronous.
According to Paquet (1969), the Lower/Middle Jurassic
boundary in the Morrón de Totana Unit (probably studied in
the Prat Mayor section), begins with a thick oolitic-limestone
level (25 m) with gastropods and bivalve fragments, evolving
to whitish-gray micritic limestones with filaments (10 m), and
later on, well-bedded gray, slightly marly, pelagic limestones
(25 m) with “Cancellophycus”, filaments, and Globochaete
alpina Lombard belonging to the Middle Jurassic. In the
Upper Jurassic analyzed in the same localities, including the
Fuente Blanca section (here called Prat Mayor), Paquet (1969)
recognized well-bedded gray limestones with filaments at the
base, slightly marly limestones with Globochaete above and
grayish-white, more or less nodular, limestones with Calpionella
alpina Lorenz, Stomiosphera minutissima (Colom),
G. alpina Lombard, Textulariidae and Lageniidae towards
the upper part.
Geyer and Hinkelbein (1971, 1974) focused on the detailed
biostratigraphy of the Liassic ferruginous oolith-rich interval,
outcropping in the Morrón de Totana (Morrón de Alhama
section, for these authors) and Prat Mayor (Fuente Blanca
section, for these authors). Geyer and Hinkelbein (1971, 1974)
made a detailed correlation of this 5 m thick section, dating it
as the Upper Pliensbachian (on the basis of a few and badly
preserved forms of Arieticeras/Canavaria/Fontanelliceras,
Lioceratoides and Catacoeloceras) in the lower part of Prat
Mayor section. In the upper part of Morrón de Totana section,
it was dated as the Middle Toarcian [Peronoceras cf.
millarense Monestier, Catacoeloceras tethysi Géczy,
Hildoceras cf. graecum Renz, Hi. cf. bifrons (Bruguière),
among others], and the Upper Toarcian [Hugia cf. variabilis
(d’Orbigny), Grammoceras thouarsense (d’Orbigny), Dumortieria
sp. and Pleydellia sp., as more relevant]. Thus, these
authors assumed an Upper Pliensbachian–Aalenian? age for
the complete interval in both sections, avoiding the interpretation
of the diachroneity proposed by Paquet (1962, 1969).
Kampchuur et al. (1974) assigned 90 m of white oolitic
limestones with algae (Thaumatoporella), ostracods, and foraminifers
to the pre-Domerian Liassic. Over these shallow platform
carbonates appear a few meters of ferruginous oolithrich
limestones. These authors accepted the dating by Paquet
(1969) for this guide horizon as Middle Domerian in the Morrón
de Totana Unit, and then the subsequent diachrony of
such level. The Dogger encompasses 100–140 m thick of
oolitic limestones with crinoids, miliolids and ostracods,
evolving to biopelmicrites, rich in echinoderms, “filaments”,
Ammodiscus, and lagenids. The Upper Jurassic is composed
of 90 m of massive limestones, occasionally nodular, with
radiolaria, Globuligerina, G. alpina Lombard, and ostracods,
with Saccocoma and calpionellids in its upper part.
Seyfried (1978) studied some Jurassic sections at Casas y
Pozos de Murcia (equivalent, in part to the Prat Mayor and
Tres Carrascas sections here) recognizing the ferruginous
oolith-rich level with abundant Middle Domerian faunas. The
Middle Jurassic materials are represented by 65–70 m of
oolitic/crinoidal limestones, and gray laminated limestones
containing trace fossils, interbedded with turbiditic levels with
resedimented oolites and scarce Upper Bajocian ammonites
(Spiroceras sp., Nannolytoceras sp.). The overlying Upper
Jurassic (approximately 45 m thick) includes calcareous breccias,
pelites and fluxoturbidites with foraminifers, bivalves
(Inoceramus) and Oxfordian ammonites (Phylloceras sp.,
Holcophylloceras sp., Sowerbyceras sp., Arisphinctes sp. and
Dichotomosphinctes? sp.) at its base, with massive limestones
above and well bedded limestones (pelmicrosparite and
biopelmicrite) with foraminifers, crinoids, and echinoids in
its upper part.
Recently, Caracuel et al. (2001) and Martín-Rojas et al.
(2002) described the general stratigraphy of a new Jurassic
section near Malvariche, where the lithofacies are similar to
those of the classical outcrops at Prat Mayor and Morrón
Chico. In the Malvariche section, these authors found ammonite
assemblages indicating the Middle Domerian (Algovianum
Zone; ferruginous oolith-rich limestones), the
Lower Callovian [Bullatus and Gracilis (= Patina zone, sensu
Sequeiros, 1974)], and the uppermost Kimmeridgian (Beckeri
Zone).
4. The sections studied: stratigraphic data
Five Jurassic sections were studied in Sierra Espuña; the
three classical sections already mentioned from Fallot (1945)

to Seyfried (1978); Prat Mayor, Morrón Chico and Perona,
(Fig. 1), and two new ones (Malvariche and Tres Carrascas
sections; Fig. 1). Sections were selected to cover the Morrón
de Totana tectonic Unit (Malvariche, Tres Carrascas, Prat
Mayor and Morrón Chico sections; Figs. 1–3) and the Perona
Unit (Perona section; Figs. 1 and 3), evenly spaced along the
Sierra Espuña.
The present study was focused in the Prat Mayor and Malvariche
sections (Fig. 2) since they are easily accessible, are
little tectonized and contain the most complete, and fossilrich
successions. In sections at Tres Carrascas, Morrón Chico
and Perona, only the Liassic (especially the ferruginous oolithrich
limestone interval) was studied.
Along with the stratigraphical and sedimentological analysis,
more than 140 thin sections were used for characterizing
the microfacies, textures and microfossil content; particularly
the Globuligerina and calpionellids, during the Upper
Jurassic, which have potential biostratigraphic interest. Macroinvertebrates
(more than 500 specimens), mainly ammonites
and brachiopods, were sampled in favorable facies for
biostratigraphy.
4.1. Lower Jurassic
The Lower Jurassic was analyzed in all sections studied. It
is composed of oo-oncolitic limestones, sometimes brecciated,
evolving to crinoidal limestones, with an interval of ferruginous
silty limestones at the top. In the Malvariche and
Prat Mayor sections, the lower boundary is better exposed
and makes contact tectonically with massive saccaroid dolostones,
sometimes attributed to the earliest Jurassic. The thickness
of the outcropping Lower Jurassic, which sometimes can
be slightly dolomitized at its base, ranges from 70 to 125 m.
No dating was established for the oo-oncolitic limestones,
but, in any case, an Early Jurassic age is ruled out by the
presence of Lithiotis-rich levels, typical for the Liassic Perimediterranean
Tethys. Moreover, Kampchuur et al. (1974)
suggests a Sinemurian to Pliensbachian age for its microfauna.
By contrast, the upper part with ferruginous silty limestones
were easily dated in all sections, ranging from the
Domerian (= Upper Pliensbachian; Lavinianum Zone, Cornacaldense
Subzone) to the Upper Toarcian (Reynesi Zone),
according to the ammonite assemblages collected.
In all sections studied, the lower part of the succession is
composed of oo-oncolitic, evolving to crinoidal limestones,
showing decametric thickening and upwardly coarsening
parasequences (10–25 m thick), quite stable both in thickness
and lithofacies variation, with no evident stacking trend.
The elementary parasequence appears to be built up by white
oo-pisolitic limestones evolving to pinkish oncolitic–rodolitic
limestones, sometimes breccioids, and topped by an algal
crust and/or an intensely bioturbated/bored surface. Crossbedded
grainstones are usual in the lower part of the parasequences.
Microfacies are mainly grainstones to packstones (occasionally
rudstones) with oolites, pisolites and/or oncolites and,
J.E. Caracuel et al. / Geobios 39 (2006) 25–42
secondarily, algae [Cayeuxia piae Frollo, Palaeodasycladus
mediterraneus (Pia)], ostracods, sponges, and benthic foraminifers.
Some levels are rich in Textulariidae, Siphovalvulina
sp., and transitional forms of Mayncina termieri Hottinger
and Lituosepta compressa Hottinger, which are
characteristics of the protected Liassic platform of the Mediterranean
(Sartorio and Venturini, 1988). Benthic macrofauna
such as gastropods, bivalves (pectinids, ostreids and
Lithiotis), brachiopods, solitary corals, echinoderms are often
abundant (Fig. 2). Generally, they are found in living positions,
as for example the characteristic Lithiotis horizon of
the upper part of the elementary parasequences.
The upper part of the Lower Jurassic succession is built up
by 2–12 m of alternating yellowish marly/silty limestones,
occasionally slightly nodular, with levels of ferruginous
oolites and/or decimetric Fe–Mn oncoids, which are formed
by concentric laminae around a nucleus (meter levels 110–
122 in Malvariche and 60–65 in Prat Mayor; Fig. 2). Only in
section at Morrón Chico, does the base of this interval show a
karst-like irregular surface sinking more than 2monthe
underlying crinoidal limestones. This characteristic interval,
which can be used as a guide-horizon for geological mapping,
is well-developed and widespread in Sierra Espuña.
Some sections contain a similar interval made up of yellowish
marly/silty limestones, although thinner, less continuous
and fossiliferous (meter levels 88–92 in Malvariche and
20–25 in Prat Mayor; Fig. 2).
In the five sections studied, the interval of ferruginous silty
limestones with Fe-oolites is rich in well-preserved macrofauna
with neomorphosed ammonite and brachiopod shells,
together with oriented belemnites, bivalves, gastropods, and
echinoderms (Fig. 3). Trace fossils (Thalassinoides, Planolites
and Chondrites) are widespread in the marly levels. As
shown in Fig. 3, the ammonites collected have enabled the
characterization of discrete and discontinuous horizons within
the Domerian (Lavinianum, Algovianum and Emaciatum
Zones) and the Toarcian (Polymorphum, Serpentinum,
Bifrons, Gradata and Reynesi Zones).
4.2. Middle Jurassic
The studied Middle Jurassic sections at Malvariche and
Prat Mayor are 138 and 145 m thick, respectively (Fig. 2).
They are composed mainly of well-stratified micritic/crinoidal
limestones with abundant chert in nodules and ribbons,
increasing toward the upper part. At the base, the micritic/
crinoidal limestones alternate with thick oolitic limestones
levels, which resemble that of the Lower Jurassic. Toward
the upper part, some levels composed by micritic limestone
show incipient nodularization, with abundant trace fossils
(mainly Thalassinoides), developing two multiple hard
grounds at the top, with an accumulation of glauconite or
ferruginous crusts and faunal concentrations, including
ammonites and occasionally oriented belemnites (“belemnite
battlefields” sensu Doyle and MacDonald, 1993).
As a whole, the lower and middle part of the Middle Jurassic
is composed of upwardly thickening and coarsening
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30 J.E. Caracuel et al. / Geobios 39 (2006) 25–42

parasequences (2–5 m thick), with stacking of upwardly
thicker parasequences. Microfacies range from wackestones
to packstones (occasionally crinoidal grainstones) with
crinoids, along with thin-shelled bivalves (“filaments”), radiolaria,
benthic and planktonic foraminifers, and other macroinvertebrate
fragments. Apart from disarticulated crinoid
ossicles, benthic macrofauna are almost absent in the Middle
Jurassic, in contrast to the Lower Jurassic. Planktonic macrofauna
is also scarce, but in the top of some levels, especially
toward the upper part, cephalopods proved widespread
(ammonites and belemnites).
In these levels, better developed in the Malvariche section,
ammonites assemblages were collected, enabling recognition
of the Lower Bajocian, based on the record of a specimen
of Skirroceras sp. Also recognized were the Lower
Callovian [Bullatus Zone; based on the record of Homoeoplanulites
sp., Macrocephalites sp., and Kheraiceras cf. bullatus
(d’Orbigny)], and the Gracilis Zone (Patina Zone sensu
Sequeiros, 1974) with abundant and significant ammonites,
especially Macrocephalinae and Reineckeidae. No other
ammonite faunas were found in the Middle Jurassic.
4.3. Upper Jurassic
The Upper Jurassic features at Malvariche and Prat Mayor
differ somewhat with regard to lithofacies and faunal assemblages,
although the thicknesses are quite similar, 80 and 90 m,
respectively (Fig. 2). As a whole, the succession is composed
by finely stratified marl and marly limestones which evolved
to stratified limestones stacked in thickening upward parasequences
(1–2 m thick). Then, massive nodular limestones
(sometimes brecciated) alternate with stratified limestones and
marls, arranged in upwardly thinning parasequences. Textures
and microfacies are highly variable, ranging from pelloidal
mudstones to intraclastic packstones (occasionally
crinoidal grainstones) with Globuligerina (at the base), filaments,
Saccocoma, radiolaria, Globochaete, Stomiosphaera,
Cadosina, macroinvertebrate fragments (mainly ammonites
and belemnites) and calpionellids (in the upper part).
In both sections (Fig. 2), above the multiple Fe–Mn crusts
dated as Lower Callovian (Gracilis Zone) outcrops a 5–10 m
thick interval of marly limestones and marls, finely stratified,
and texturally mudstones to wackestones rich in Globuligerina,
attributed to the Callovian?–Oxfordian. In the Malvariche
section,7mofslightly nodular marly limestones appear
in upwardly thickening parasequences. Towards the top, there
is an irregular massive breccioid bank,4mofslightly nodular
marly limestones organized in upwardly thinning parasequences,
and a characteristic crinoidal bank with an erosive
base. Just below this crinoidal bank, ammonite assemblages
were collected from the uppermost Kimmeridgian (Beckeri
Zone). These slightly nodular marly limestones stacked in
Fig. 2. Correlation of the Jurassic sections at Malvariche and Prat Mayor.
Fig. 2. Corrélation des sections jurassiques de Malvariche et Prat Mayor.
J.E. Caracuel et al. / Geobios 39 (2006) 25–42
upwardly thinning parasequences continues 15 m more over
the encrinitic bank, and then change to upwardly thickening
parasequences; the facies change gradually to alternating
marly/calcareous nodular limestones, often brecciated. The
upper 20 m are composed of crinoidal levels and marly intervals
with microfacies enriched in hyaline calpionellids which
characterized the Upper Tithonian.
In the Prat Mayor section above the marly limestones with
Globuligerina, outcrop 50 m of ammonitico rosso nodular
limestones alternating with thick breccioids banks, organized
in upwardly coarsening and thickening parasequences
(2–4 m thick). In this interval, ammonite assemblages characterizing
the Middle–Upper Oxfordian and the Lower–
Middle? Kimmeridgian were collected. In contrast to the Malvariche
section, no ammonites from the Upper Kimmeridgian
were found. The upper part of section at Prat Mayor appears
similar to Malvariche, although more reduced and with characteristic
breccioid banks. In both sections, the calpionellids
Zones of Chitinoidella, Crassicollaria and Calpionella were
recorded.
5. Ammonite biostratigraphy
Jurassic ammonite faunas belonging to the Internal Zone
of the Betic Cordillera has been rarely and discontinuously
reported, and in all cases from the non-metamorphosed
Malaguide and “Dorsal” domains (Fallot, 1929, 1931–1934,
1945; Peyre and Peyre, 1960; Azema, 1960, 1961; Paquet,
1969; Geyer and Hinkelbein, 1971, 1974; Seyfried, 1978 and
Caracuel et al., 2001). Biostratigraphic data from Sierra
Espuña come mainly from Geyer and Hinkelbein (1974); Seyfried
(1978) and Caracuel et al. (2001), who described ammonite
assemblages belonging to the Domerian, Middle–Upper
Toarcian, Upper Bajocian, Oxfordian and Upper Kimmeridgian.
For the present paper, more than 500 macroinvertebrates,
mainly ammonites, were collected in order to complete
and to revise the biostratigraphic framework from the
Domerian to the Kimmeridgian in the three previously studied
and two new Jurassic sections at Sierra Espuña. The
ammonite zonal/subzonal scheme in Cariou and Hantzpergue
(1997) for the Mediterranean Domain was used in the
Domerian, Toarcian and Callovian stages.
Domerian ammonites show good preservation with neomorphosed
shells, sometimes ferruginous, and mostly with
preserved living chambers; phragmocone septa are consistently
preserved (Figs. 4 and 5). There appears to be no taphonomic
size bias, although the smaller ammonoids (occasionally
fragments) are sometimes found in the interior of Fe–Mn
oncoids, with a variable mode of preservation.
31

32 J.E. Caracuel et al. / Geobios 39 (2006) 25–42
Fig. 3. Detailed successions in the five Domerian–Toarcian sections studied; interval of alternating yellowish marly/silty limestones, occasionally slightly
nodular, with levels of ferruginous oolites and/or decimetric oncoids. Recognized Zones and Subzones (according to Cariou and Hantzpergue, 1997) are shaded
in the chronostratigraphic chart. Legend as in Fig. 1.
Fig. 3. Successions détaillées des cinq sections du Domérien–Toarcien étudiées ; intervalle d’alternance des calcaires marno-silteux jaunâtres, parfois légèrement
noduleux, avec des niveaux décimétriques d’oncoïds et/ou d’oolites ferrugineuses. Les zones et sous-zones reconnues (d’après Cariou et Hantzpergue,
1997) sont ombragées sur le tableau chronostratigraphique. Légende, cf. Fig. 1.

In the lower level of yellowish silty/limestones or
silty/marly limestones of the Malvariche section (meter levels
88–92), which contains abundant bivalves, ammonite are
scarce and fragmentary; we have recognized Lytoceras villae
Meneghini, Fuciniceras gr. isseli (Fucini), Fu. cornacaldense
(Taush) and Fieldingiceras fieldingii (Reynes), indicating
that, at least in this section, the Lower Domerian
(Lavinianum Zone, Cornacaldense Subzone) is represented
in Sierra Espuña.
In Malvariche (meters 110–122) and Prat Mayor (meters
60–65) the alternating yellowish marly/silty limestones, occasionally
slightly nodular, with levels of ferruginous oolites
and oncoids (Figs. 2 and 4), have supplied abundant, diversified
and relatively well preserved ammonites: Ly. villae
Meneghini (Fig. 4A), Fi. fieldingii (Reynes) (Fig. 4B), Fu.
cornacaldense (Taush) (Fig. 4C), Protogrammoceras aequiondulatum
(Bettoni), Pr. aff. ilurcense Braga, Arieticeras
algovianum (Oppel) (Fig. 4E), Ar. disputabile (Fucini)
(Fig. 4F), Ar. amalthei (Oppel), Ar. bertrandi (Kilian), Leptaleoceras
accuratm (Fucini), Le. ugdulenai (Gemmellaro),
Becheiceras bechei (Sowerby), Reynesoceras acanthoides
(Reynes), Re. ragazzonii (Hauer) (Fig. 4D), and Parstchiceras
aff. proclive (Rosemberg). According to Braga (1983),
this ammonite assemblage dates the Middle Domerian
(Algovianum Zone; Ragazzoni, Bertrandi and Accuratum
Subzones) and possibly the Lower Domerian [Lavinianum
Zone, as is shown the by the record of Fu. cornacaldense
(Taush), Fi. Fieldingii (Reynes) and Be. Bechei (Sowerby)].
Although most of the ammonites are clearly reworked, it is
possible that, with a detailed sampling, the three Middle
Domerian Subzones could be separated, given that different
samples (some with several ammonites species) can have different
lithologies.
In Tres Carrascas section, at 40 cm of the base (Fig. 3), we
have recorded Emaciaticeras levidorsatum Fucini (Fig. 4H),
Em. speciosum Fucini, Em. sp., and Dactyliocetatidae ind.,
which are representative of the upper Middle Domerian
(Algovianum Zone; Levidorsatum Subzone). Also in this section,
60 cm upward (samples TC.100C) we have found Pleuroceras
solare (Phillips), Em. sp., Lioceratoides fucinianus
(Haas), Li. exapatus (Gemmellaro), and Neolioceratoides
hoffmanni (Gemmellaro), which characterize the lower Upper
Domerian (Emaciatum Zone, Solare Subzone). The samplings
TC.100A and TC.100B, both in Tres Carrascas section
(Fig. 3) contain Em. archimedis Fuccini, Em. lottii (Gemmellaro)
(Fig. 4G), Em. timaei (Gemmellaro) (Fig. 4I), Em.
sp., Li. serotinus (Bettoni), Li. micetoi (Fucini) and Ne. hoffmanni
(Gemmellaro). This assemblage dates the uppermost
Domerian (Emaciatum Zone, Elisa Subzone).
Domerian/Toarcian transition has been recognized in the
lower part of the Morrón Chico section (Fig. 3), with the following
ammonite assemblage: Li. lorioli (Bettoni), Li. sp.,
Ne. hoffmanni (Gemmellaro) Ne. schopeni (Gemmellaro),
Canavaria sp., Ca. gregalis Fucini and Fontanelliceras fontanellense
(Gemmellaro). Braga (1983) proposed that this
assemblage may correspond to the uppermost Domerian (Elisa
Subzone)–lowermost Toarcian (Polymorphum Zone).
J.E. Caracuel et al. / Geobios 39 (2006) 25–42
As in the External Zones (Subbetic) of the Betic Cordillera
(see Braga, 1983), the Hildoceratidae (Harpoceratinae
and Hildoceratinae) and Lytoceratidae clearly dominate the
Domerian ammonite assemblages, indicating its typical Mediterranean
character. Dactylioceratinae are also abundant in
the Ragazzoni Subzone, whereas other groups such as Phylloceratidae,
Amaltheidae and Liparoceratidae, though also
present, are scarce.
Toarcian materials are clearly represented in the Perona
Unit (Fig. 3) where some specimens of Dactylioceras (Eodactylites)
sp. and Hildaites striatus Guex have been recorded,
pointing to an Early Toarcian age (Polymorphum and Sepentinum
Zones).
Middle and Upper Toarcian sediments are well represented
in the Morrón Chico section, where a minimum of
four fossiliferous levels can be differentiated (Fig. 3). The
lower one (40 cm upward of the base), reported above, represents
the Domerian/Toarcian transition. The following fossiliferous
level, located approximately 140 cm upward
(samples MC.180), shows reduced net sedimentation with
very thinly laminated Fe–Mn layers which encrust either the
discontinuity surfaces or very well-preserved fossils, especially
ammonites. The following ammonite species occur:
Hildoceras bifrons (Brugière) (Fig. 4J), Hi. semipolitum
Buckman, Pseudomercaticeras venzoi Pinna, Osperleioceras
bicarinatum (Zieten), Phymatoceras (Furloceras) chelussi
(Parisch and Viale) (Fig. 5A), Ph. (Fu.) venustulum
(Merla), Mouterdeiceras sp., Catacoeloceras crassum (Young
and Bird), Ca. sp., Platystrophites latusi Levi-Seti and Oxyparoniceras?
sp. This ammonite assemblage represents the
Middle Toarcian, Bifrons Zone (Bifrons Subzone) and possibly
the base of the Gradata Zone. A single specimen of
Mouterdeiceras sp. from 6.5 m from the base in the Morrón
Chico section indicates the Gradata Zone.According to Geyer
and Hinkelbein (1974), who reported Upper Toarcian ammonites
from this section, some specimens of Geczyceras speciosum
(Janensch) (Fig. 5B), Pseudogrammoceras sp. and
Lytoceras sp. appear from approximately 3.5 m upward,
which are indicative of the lower part of the Upper Toarcian,
lower part of Reynesi Zone (Fig. 5). A single fragment of ex
situ Dumortieria sp. indicates the upper part of the Reynesi
Zone.
Middle Jurassic ammonite assemblages were better registered
in the Malvariche section. Just below the lower level of
a multiple limonite crust (230 m in Fig. 2) we sampled scarce
and badly preserved fauna (Skirroceras sp.), which can be
attributed to the upper Lower Bajocian. The upper levels of
these crusts have provided articulated crinoids, oriented
belemnites, and some deformed specimens of Homoeoplanulites
sp., Macrocephalites sp., Kheraiceras cf. bullatus
(Orbigny) and Kh. (Bomburites) sp. from the Lower Callovian
(Bullatus Zone).
Some meters upward, also in the Malvariche section,
ammonite assemblages were collected from the Lower Callovian
(Gracilis Zone = Patina Zone, sensu Sequeiros, 1974).
These show taphonomic reworking with neomorphized shells
33

34 J.E. Caracuel et al. / Geobios 39 (2006) 25–42
Fig. 4. Lower Jurassic ammonites from Sierra Espuña. All the specimens are figured at natural size except for B (× 2). A. L. villae Meneghini, MI.121S.1,
Middle Domerian, Algovianum Zone, Malvariche section 121 m. B. F. fieldingii (Reynes), MI121S.2, Middle Domerian, Malvariche section 121 m. C. Fuciniceras
cornacaldense (Taush) MI.121S.3, Malvariche section 121 m, D. Reynesoceras ragazzonii (Hauer), MI.121S.4, Middle Domerian, Algovianum Zone,
Malvariche 121.5 m. E. A. algovianum (Oppel), MI.121S.5, Algovianum Zone, Malvariche section 121.5 m. F. Arieticeras disputabile (Fucini), PM.60.10,

together with inner molds, sometimes truncated, corroded,
occasionally fragmented, and usually imbricate. Both inner
molds and shells can occasionally be covered by Fe–Mn
crusts. The assemblage is dominated by Phylloceratina [Phylloceras
trifoliatum Neumayr, Holcophylloceras zignodianum
(Orbigny), Calliphylloceras disputabile (Zittel) and Ptychophylloceras
flabellatum (Neumayr)], which reach the 60%.
Macrocephallitinae as Macrocephalites gracilis (Spath)
(Fig. 5E), and Macrocephalites? sp. along with Haploceratidae
[Lissoceratoides jullieni (Douvillé)] are also abundant.
Other groups as Hecticoceratinae (Hecticoceras (Chanasia)
bannense Elmi (Fig. 5C), He. (Ch.) sp. and
Jeanneticeras? sp.), Reineckeiidae [Rehmannia (Re.) freii
(Jeannet) sensu Cariou (1980) (Fig. 5D), Collotia oxyptycha
(Neumayr)], Perisphinctidae (Choffatia waageni (Teisseyre),
Grossouvria sp., Parapatoceratinae, (Parapatoceras
tuberculatum (Baugier and Sauzé), Pa. sp.) and Oppeliinae
(Oxycerites sp.) are also common. Middle Jurassic ammonites
are scarcer in Prat Mayor section. Seyfried (1978)
reported Spiroceras sp. and Nannolytoceras sp. from the
Upper Bajocian, and we collected only two specimens (Hecticoceras
(Ch.) sp. and Grossouvria sp., 214 m) which probably
represent the Lower Callovian (Gracilis Zone).
Oxfordian ammonite assemblages were recorded in 7 m
of succession in the Prat Mayor section (237.5–244.5 m in
Fig. 2) at the top of nodular-limestone levels, sometimes brecciated.
As usual in ammonitico rosso facies, ammonites are
preserved as inner moulds with complete phragmocone and
incomplete body chambers. Sutures are poorly or not at all
preserved, indicative of relatively deep deposition (Fernández-
López, 2000). No significant taphonomic size bias was noted,
although the smaller ammonites are frequently lying oblique
(even vertical) with respect to the stratification, especially at
the top of the strata. As a whole, Phylloceratina reach the
37.5% (mainly Calliphylloceras and secondarily Sowerbyceras)
of the assemblage.
The Middle Oxfordian, Transversarium Zone, was registered
in a single horizon with the record of Calliphylloceras
silesiacum (Oppel), Ca. manfredi (Oppel), Sowerbyceras tortisulcatum
(d’Orbigny), Euaspidoceras (Eu.) gr.oegir
(Oppel), Taramelliceras (Ta.) cf. callicerum (Oppel),
Perisphinctes (Arisphinctes) cf. helenae De Riaz and Pe.
(Dichotomosphinctes) sp., among others. Some 2 m upward,
there are three horizons with scarce but significant fauna of
J.E. Caracuel et al. / Geobios 39 (2006) 25–42
Pe. (Dichotomoceras) bifurcatoides Enay (Fig. 6A), Pe.(Di.)
cf. bifurcatus (Quenstedt), Pe.(Dichotomosphinctes) cf. elisabethae
(De Riaz) and a specimen of Pe. (Dichotomosphinctes)aff.ultimus,
with atypical abundance of trifurcate ribs in
the body chamber, together with Calliphylloceras and Sowerbyceras,
which characterize the Bifurcatus Zone. The Upper
Oxfordian, Bimammatum Zone (or still the uppermost Bifurcatus
Zone) was unsurely recorded in a horizon with Passendorferia
(Passendorferia) cf. teresiformis (Brochwich-
Lewinski), Pa. (Pa.) sp. and Trimarginites gr. trimarginatus
Oppel, just 50 cm below the first record of Subnebrodites in
the Planula Zone. Upwardly, four ammonite-bearing horizons,
spaced within a thickness of 2.5 m, enabled the recognition
of the Planula Zone by the record of Subnebrodites cf.
schröederi (Wegele), Su. cf. minutum (Hehl) sensu Zieten
(1830–1834), Passendorferia (Pa.)aff.uptonoides (Fig. 6B),
Pa. (Pa.) sp., Physodoceras altenense (d’Orbigny), Orthosphinctes
sp. and abundant Phylloceratina (mainly Sowerbyceras).
Kimmeridgian ammonite assemblages in the Prat Mayor
section were registered from 246.4 to 260 m (Figs. 2 and 6).
As in the underlying Oxfordian, the lithofacies is composed
by nodular limestones with thicker and more brecciated levels.
The Kimmeridgian succession has less, and more widely
spaced, ammonite-bearing horizons. Nevertheless, the ammonite
mode of preservation is similar to that of the Oxfordian,
although, due to the higher sedimentation rate, some specimens
have lost the inner whorls (or have become flattened
due to the absence of sediment infill during early diagenesis).
The amount of Phylloceratina is higher, reaching the 50%,
mainly by the contribution of the eurythopic genus Sowerbyceras.
Almost 2 m above the last record of the Planula Zone, there
is a horizon attributed to the lowermost Kimmeridgian
(Platynota Zone?) based on the record of Sutneria gr. galar
(Oppel), Streblites tenuilobatus (Oppel), Glochiceras (Lingulaticeras)gr.nudatum
(Oppel)-lingulatum (Quenstedt), Gl.
(Li.) sp., Aspidoceras sp., Orthosphinctes (Orthosphinctes)
polygyratus (Reinecke) sensu Schairer morph. colubrinus
Olóriz (Fig. 6F), Or.(Or.) sp, Or.(Lithacosphinctes) sp. and
abundant Phylloceratina [So. silenum (Fontannes) as well as,
secondarily, Ly. orsinii (Gemmelaro) and Ho. mediterraneum
(Neumayr)]. Above appear Nebrodites (Nebrodites)gr.
hospes (Neumayr) (Fig. 6C), Ne.(Ne.) sp., Presimoceras sp.,
Middle Domerian, Algovianum Zone, Prat Mayor section 60 m. G. Emaciaticeras lottii Fucini, TC.100A.3, Upper Domerian, Emaciatum Zone, Tres Carrascas
section 1 m. H. E. levidorsatum Fucini, TC.40.1 Upper Domerian, Emaciatum Zone, Tres Carrascas section 0.4 m. I. Emaciaticeras timaei (Gemmellaro),
TC.100A.2, Upper Domerian, Emaciatum Zone, Tres Carrascas section 1 m. J. H. bifrons (Brugière), MC.180.1, Middle Toarcian, Bifrons Zone, Morrón Chico
section 1.8 m.
Fig. 4. Ammonites du Jurassique inférieur de la Sierra Espuña. Tous les spécimens sont représentés à grandeur naturelle sauf B (× 2). A. L. villae Meneghini,
MI.121S.1, Domérien moyen, Zone à Algovianum, section de Malvariche 121 m. B. F. fieldingii (Reynes), MI121S.2, Domérien moyen, section de Malvariche
121 m. C. Fuciniceras cornacaldense (Taush) MI.121S.3, section de Malvariche 121 m. D. Reynesoceras ragazzonii (Hauer), MI.121S.4, Domérien moyen,
Zone à Algovianum, section de Malvariche 121,5 m. E. A. algovianum (Oppel), MI.121S.5, Zone à Algovianum, section de Malvariche 121,5 m. F. Arieticeras
disputabile (Fucini), PM.60.10, Domérien moyen, Zone à Algovianum, section de Prat Mayor 60 m. G. Emaciaticeras lottii Fucini, TC.100A.3, Domérien
supérieur, Zone à Emaciatum, section de Tres Carrascas 1 m. H. E. levidorsatum Fucini, TC.40.1, Domérien supérieur, Zone à Emaciatum, section de Tres
Carrascas 0,4 m. I. Emaciaticeras timaei (Gemmellaro), TC.100A.2, Domérien supérieur, Zone à Emaciatum, section de Tres Carrascas 1 m. J. H. bifrons
(Brugière), MC.180.1, Toarcien moyen, Zone à Bifrons, section de Morrón Chico 1,8 m.
35

36 J.E. Caracuel et al. / Geobios 39 (2006) 25–42
Fig. 5. Lower and Middle Jurassic ammonites from Sierra Espuña. All the specimens are figured at natural size except for A (× 0.7). A. Phymatoceras (Furloceras)
chelussi (Parisch and Viale), MC.180.8, Middle Toarcian, Gradata Zone, Morrón Chico section 1.8 m. B. G. speciosum (Janensch), MC.1000.1, Upper
Toarcian, Reynesi Zone, Morrón Chico section 10 m. C. Hecticoceras (Chanasia) bannense Elmi, MI.251.1, Lower Callovian, Gracilis Zone, Malvariche

Taramelliceras (Ta.) cf. trachinotum (Oppel) (Fig. 6E), Ta.
(Ta.) sp., Glochiceras (Lingulaticeras) gr.crenosum (Quenstedt),
Pseudowaagenia aff. micropla (Oppel), Ataxioceras
(Schneidia?) sp., and Ataxioceras sp., along with abundant
Sowerbyceras silenum (Fontannes), which dated the uppermost
Platynota or the Strombecki Zone.
The remainder Lower Kimmeridgian (Strombecki or
Divisum Zone) was recorded 2.6 m above, with the record of
Nebrodites (Nebrodites)gr.hospes (Neumayr), Ne.(Ne.) sp.,
Ta. (Ta.) subcallicerum (Gemmellaro) (Fig. 6D), Ta.(Ta.) cf.
pseudoflexuosum (Favre), Ta.(Ta.) sp., and Streblites sp.
(Fig. 6). In the last 6 m appeared scarce and badly preserved
Ta. (Ta.)gr.pugile (Neumayr), Discosphinctoides (Di.) aff.
capillaceous (Dumortier) with Ho. mediterraneum (Neumayr)
and So. silenum (Fontannes), which belong to the
Divisum Zone.
The Upper Kimmeridgian was recognized in the top of
four nodular-breccioids levels at 260 m in the Malvariche section
(Figs. 2 and 6). Ammonites were badly preserved as
reworked inner moulds with common fragmentation, imbrication
and generalized disarticulation of body chambers and
phragmocones along the septa. The recorded assemblage is
dominated (more than 80%) by Phylloceratina [mainly So.
loryi (Munier-Chalmas) morphs loryi and pseudosilenum, and
few Ho. polyholcum (Benecke)].
The four horizons studied were attributed to the Beckeri
Zone, with a similar ammonite assemblage composed, as more
relevant, of Hybonoticeras (Hy.) beckeri beckeri (Neumayr)
(Fig. 6H), Hy. (Hy.) beckeri harpephorum (Neumayr)
(Fig. 6G), Hy. (Hy.) gr. beckeri (Neumayr), Ta. (Ta.) pugile
pugile (Neumayr), Ta.(Ta.) gr. pugile (Neumayr), Glochiceras
(Lingulaticeras) sp., Shaireria cf. episa (Oppel), Aspidoceras
cf. sesquinodosum (Fontannes), Torquatisphinctes cf.
laxus Olóriz, Biplisphinctes cf. uracensis Berckhemer and
Discosphinctoides (Discosphinctoides) sp. (Fig. 6)
No Tithonian ammonites were recorded in the Malvariche
or in the Prat Mayor section. Thus, the Upper Tithonian was
evidenced by the FAD of hyaline calpionellids, and the record
of the Crassicollaria Zone, at 295 m in the Malvariche and
285 m in the Prat Mayor sections (Fig. 2). Finally, the
Tithonian/Berriasian boundary was approached by the bloom
of large and isometric C. alpina (Calpionella Zone) at 330 m
in the Malvariche and 305 m in Prat Mayor sections.
6. Paleoenvironmental interpretation
The evolution of the Jurassic from the Malaguide at Sierra
Espuña is comparable to other sectors of the Tethyan paleo-
J.E. Caracuel et al. / Geobios 39 (2006) 25–42
margins belonging to Internal and External Zones: the Subbetic
(S Spain, Vera, 1988), the Venetian Alps (N Italy, Zempolich,
1993), the Apennines (Central Italy, Colacicchi et al.,
1999), the Trapanese, (W Sicily, Catalano et al., 2002)orthe
Ghomarids (N Africa, Maate, 1996). During the Jurassic it
evolves as a passive margin, beginning with the pre-rifting
stage, followed by the platform break-up of the rifting stage
(starting from the Domerian, Lavinianum Zone), and finally
the drifting stage (from the Lower Callovian, Gracilis Zone).
As shown in Fig. 7, over the earliest Liassic dolostones,
the outcropping pre-Domerian deposits are built up by
oo-oncolitic limestones, sometimes breccioids, evolving
upwards to crinoidal limestones. These are interpreted as
restricted inner-shelf deposits of oolitic shoals, nearby algal
and/or crinoidal meadows. Accordingly, the recorded faunas
are solely abundant and well-diversified shallow-water
benthos such as algae, crinoids, sponges, gastropods, bivalves
(pectinids, ostreids and Lithiotis), brachiopods, solitary corals,
echinoderms, and benthic foraminifers, among others.
These benthic faunal assemblages are dominated by suspension
feeders, with little resedimentation processes, even lying
in living position (Lithiotis, corals). In such a context, the
recorded upwardly thickening and coarsening parasequences
(more developed in the Malvariche section; Fig. 3) may be
interpreted as upwardly shallowing cycles.
Above, the Domerian-Toarcian guide interval of alternating
yellowish marly/silty limestones (slightly nodular) with
ferruginous oolites and oncoids, records the beginning of the
rifting stage with the break-up of the platform (drowning
unconformity). This event is linked to a tectonic pulse (active
listric faulting and tilting blocks), evidenced by the variable
thicknesses and lithofacies among sections (Fig. 3) and the
presence of neptunian dykes (e.g. Morrón Chico section,
Fig. 3). The restricted inner platform, where the underlying
oolitic limestones developed, may drown at relatively shallow
depth, leading to drastic reduction of carbonate productivity
with changes in current circulations and water chemistry,
probably with contribution of upwelling of eutrophic
water, rich in Fe–Mn oxi-hydroxides (influx of trophic
resources and plankton coming from the open sea into the
platform). Thus, the faunal assemblages are alternatively
dominated by benthos of low diversity (mainly brachiopods,
echinoderms, bivalves), and, for the first time, pelagic assemblages;
stunted ammonites and belemnites of low diversity,
with intense reworking, sometimes wrapped by ferruginous
centimetric-decimetric oncoids.
The interpreted depositional environment may be an open
shelf, with variable depth, water chemistry and hydrodynamics,
due to the intricate bottom topography (block faulting
section 251 m. D. Rehmannia (Rehmannia) freii (Jeannet) sensu Cariou (1980), MI.251.3, Lower Callovian, Gracilis Zone, Malvariche section 251 m. E. M. gracilis
(Spath), MI.251.2, Lower Callovian, Gracilis Zone, Malvariche section 251 m.
Fig. 5. Ammonites du Jurassique inférieur et moyen de la Sierra Espuña. Tous les spécimens sont représentés à grandeur naturelle sauf A (× 0.7). A. Phymatoceras
(Furloceras) chelussi (Parisch et Viale), MC.180.8, Toarcien moyen, Zone à Gradata, section de Morrón Chico 1,8 m. B. G. speciosum (Janensch),
MC.1000.1, Toarcien supérieur, Zone à Reynesi, section de Morrón Chico 10 m. C. Hecticoceras (Chanasia) bannense Elmi, MI.251.1, Callovien inférieur,
Zone à Gracilis, section de Malvariche 251 m. D. Rehmannia (Rehmannia) freii (Jeannet) sensu Cariou (1980), MI.251.3, Callovien inférieur, Zone à Gracilis,
section de Malvariche 251 m. E. M. gracilis (Spath), MI.251.2, Callovien inférieur, Zone à Gracilis, section de Malvariche 251 m.
37

38 J.E. Caracuel et al. / Geobios 39 (2006) 25–42
Fig. 6. Upper Jurassic ammonites. All the specimens at natural size, except for B (× 0.20). A. Perisphinctes (Dichotomoceras) bifurcatoides Enay, PM.241.6-1,
Middle Oxfordian, Bifurcatus Zone, Prat Mayor section 241.6 m. B. Passendorferia (Passendorferia)aff.uptonoides, PM.243.7-1, Upper Oxfordian, Planula
Zone, Prat Mayor section 243.7 m. C. Nebrodites (Nebrodites) gr.hospes (Neumayr), PM.252-1, Lower Kimmeridgian, Strombecki-Divisum? Zone, Prat
Mayor section 252 m. D. Taramelliceras (Taramelliceras) subcallicerum (Gemmellaro), PM.252-2, Lower Kimmeridgian, Strombecki-Divisum? Zone, Prat

and tilting). Fig. 3 shows three tectonic blocks, limited by
extensional faults. The intermediate block, which contains
the Tres Carrascas, Prat Mayor and Morrón Chico sections,
tends to sink eastward, developing less thickness (“ammonitico
rosso” condensed limestones) in the Tres Carrascas section
and a thicker succession with yellowish marly/silty limestones
containing ferruginous oolites and oncoids in the
Morrón Chico section. The generalized sediment starvation
caused by the drowning of the platform led to a faunal accumulation
as well as Fe–Mn oxi-hydroxides, which, together
with the few available sediments, may have been distributed
by waves/currents to the active depocenter (eastern parts of
the tilted blocks; Fig. 3), tending toward smoothing relief.
As in other Western Tethyan paleomargins, the tectonic
subsidence during the progressing rifting stage in the Dogger
(probably already from the uppermost Liassic) caused an
incipient half-graben system in the area (Fig. 7). Sierra Espuña
constitutes a relatively subsident depocenter, in an outer shelf
to upper-talus paleogeographic context. Thus, the sedimentation
during this period is dominated by pelagic or hemipelagic
sediments (gravity flows and periplatform ooze),
enriched in chert, coming from the contribution of siliceous
planktonic organisms (radiolaria), especially in the upper part.
Consequently, the recorded upwardly thickening and coarsening
parasequences, with stacking of thicker parasequences
above reflected the general progradation of the platform.
Fauna is scarce in these well-stratified micritic/crinoidal
limestones with chert. Benthos is represented almost exclusively
by the disarticulated crinoid knuckles, which, together
with oolites, came from the shallower part of the platform
transported by gravity flows of sediment. Pelagic faunal
assemblages are recorded only toward the upper part
(Lower/Upper Bajocian and lowermost Callovian), in some
condensed horizons at the top of strata. They are dominated
by belemnites (often oriented, “belemnite battlefield” sensu
Dolyle and Mac Donald, 1993) and/or complete ammonites
with body-chamber, but no septa preservation (mostly pelagic
free-swimmer forms such as Phylloceratids), which is interpreted
as in situ burial, in a relatively deep environment
(Fernández-López, 2000). On the contrary, the recorded
Lower Callovian ammonites, which occur in relation to stratigraphic
discontinuities, are truncated, corroded, fragmented
inner moulds that, at times, can hardly be distinguished from
J.E. Caracuel et al. / Geobios 39 (2006) 25–42
mere intraclasts, together with epigenized shelly specimens;
both are often domed by Fe–Mn laminae which are sometimes
concentric around of the nucleus. Similar cephalopod
concentrations appear in the Betic External Zones (Subbetic).
These have been interpreted (Sandoval and Checa,
2002) as reworked assemblages formed on an irregular substrate
subjected to energy pulses which would have led to the
formation and repeated destruction of accumulation beds with
different microfacies.
As in many other sectors of the Tethyan paleomargin, the
drifting stage starts around the Dogger/Malm boundary,
changing the generalized tectonics by thermal subsidence.
This stresses the differentiation of the horst and graben system,
and then the diversification of depositional environments
and facies (Figs. 2 and 7). During the Malm, the depositional
environment for the Malaguide in Sierra Espuña is a
mid-outer shelf that rises in some areas, becoming a distal
pelagic swell with sedimentation of condensed ammonitico
rosso, and related facies. In the study area, the condensed
sedimentation linked to this raised sea-bottom started gradually
from the Middle Oxfordian (Fig. 2).
Among the two sections studied, a single tilted block or
more likely, as in the Liassic, two different blocks can be
interpreted, western side of which was relatively uplifted (situation
of the Malvariche section; Fig. 2) while the eastern side
was relatively sunk (Prat Mayor; Fig. 2). Because of this, over
the interval with finely stratified marls and marly limestones,
which are, consequently, thicker in the Prat Mayor section,
the Middle–Upper Oxfordian to Kimmeridgian nodularbreccioid
limestones were reduced and hiatal in the Malvariche
section, while expanded and relatively with abundant
resedimentation (pebbly mudstones) in the Prat Mayor section.
Moreover, the relatively shallow depocenter at Malvariche
presents only pelagic assemblages with ammonites in
the uppermost Kimmeridgian, when the eustatic sea level was
the highest of the Upper Jurassic, while the relatively deeper
depocenter at Prat Mayor collected ammonites and other
pelagic faunas from the Middle Oxfordian.
In general, the Upper Jurassic ammonite assemblages are
dominated by phylloceratids (especially by the eurythopic and
ubiquist genre Sowerbyceras), recorded only in condensed
nodular limestones, ammonitico rosso and related facies. As
usual in this facies, cephalopods are variably preserved as
Mayor section 252 m. E. Taramelliceras (Taramelliceras) cf. trachinotum (Oppel), PM.249.4-12, Lower Kimmeridgian, Strombecki? Zone, Prat Mayor section
249.4 m. F. Orthosphinctes (Orthosphinctes) polygyratus (Reynecke) sensu Schairer morph colubrinus Oloriz, PM.246.4-15, Lower Kimmeridgian, Platynota?
Zone, Prat Mayor section 246.4 m. G. Hybonoticeras (Hybonoticeras) beckeri harpephorum (Neumayr), MIII.1230-1250-15, Upper Kimmeridgian, Beckeri
Zone, Malvariche III section 1230–1250 m. H. Hybonoticeras (Hybonoticeras) beckeri beckeri (Neumayr), MIII.1270-1280-53, Upper Kimmeridgian, Beckeri
Zone, Malvariche III section 1270–1280 m.
Fig. 6. Ammonites du Jurassique supérieur. Tous les spécimens sont représentés à grandeur naturelle sauf B (× 0.20). A. Perisphinctes (Dichotomoceras)
bifurcatoides Enay, PM.241.6-1, Oxfordien moyen, Zone à Bifurcatus, section de Prat Mayor 241,6 m. B. Passendorferia (Passendorferia) aff.uptonoides,
PM.243.7-1, Oxfordien supérieur, Zone à Planula, section de Prat Mayor 243,7 m. C. Nebrodites (Nebrodites)gr.hospes (Neumayr), PM.252-1, Kimméridgien
inférieur, Zone à Strombecki-Divisum?, section de Prat Mayor 252 m. D. Taramelliceras (Taramelliceras) subcallicerum (Gemmellaro), PM.252-2, Kimméridgien
inférieur, Zone à Strombecki-Divisum?, section de Prat Mayor 252 m. E. Taramelliceras (Taramelliceras) cf. trachinotum (Oppel), PM.249.4-12,
Kimméridgien inférieur, Zone à Strombecki?, section de Prat Mayor 249,4 m. F. Orthosphinctes (Orthosphinctes) polygyratus (Reynecke) sensu Schairer
morph. colubrinus Oloriz, PM.246.4-15, Kimméridgien inférieur, Zone à Platynota?, section de Prat Mayor 246,4 m. G. Hybonoticeras (Hybonoticeras)
beckeri harpephorum (Neumayr), MIII.1230-1250-15, Kimméridgien supérieur, Zone à Beckeri, section de Malvariche III 1230–1250 m. H. Hybonoticeras
(Hybonoticeras) beckeri beckeri (Neumayr), MIII.1270-1280-53, Kimméridgien supérieur, Zone à Beckeri, section de Malvariche III 1270–1280 m.
39

40 J.E. Caracuel et al. / Geobios 39 (2006) 25–42
Fig. 7. Synthetic Jurassic successions at Sierra Espuña, with indication of biofacies, microfacies and paleoenvironmental interpretation for the main lithofacies.
M (mudstones), W (wackestones), P (packstones), G (grainstones). Legend for bioclasts in Fig. 2.
Fig. 7. Succession synthétique du Jurassique de la Sierra Espuña, avec indication des biofaciès, microfaciès et interprétation paléoenvironnementale du lithofaciès
principal. M (mudstones), W (wackestones), P (packstones), G (grainstones). Légende des bioclastes Fig. 2.
inner moulds, mostly reworked; frequent loss of bodychambers
and truncations incompatible with the stratification.
In the nodular-brecciate facies, inner moulds of ammonites
are often fragmented and imbricate.
Finally, at the end of the Jurassic the generalized thermal
subsidence tends to deepen the paleomargin, while the huge
relief caused by the horst and graben system is smoothed.
Consequently, the area evolved to a basin in Lower Berriasian
(probably already from the uppermost Tithonian) with
deposition of periplatform limestones together with the local
sedimentation by planktonic microfossils (calpionellids, foraminifers,
radiolarians, as well as algae).
7. Conclusions
A multidisciplinary study has enabled greater precision in
updating the biostratigraphic framework and the paleoenvironmental
interpretation of the Jurassic succession at Sierra
Espuña. This area, which is one of the bigger, better exposed
and more fossiliferous Jurassic outcrops of the Malaguide
domain, can be considered a clue area to analyze the evolution
of the Internal Zones of the Betic Cordillera.
As a whole, ammonite biostratigraphic data from the Internal
Zones are scanty, and related only to the “Dorsal” and
Malaguide domain such as the Sierra Espuña area. Particularly
for this area, the previous biostratigraphic data, which
comes from the 1960s and 1970s, need to be updated and
revised. Thus, three previously studied and two new Jurassic
sections at Sierra Espuña were sampled, leading to a more
precise biostratigraphic ammonite framework. Ammonite
assemblages have enabled the recognition of the Domerian,
Lavinianum (Cornacaldense Subzone), Algovianum (Ragazzoni,
Bertrandi, Accuratum and Levidorsatum Subzones) and
Emaciatum (Solare and Elisa Subzones) Zones, the Lower
Toarcian, Polymorphum and Serpentinum Zones, the Middle
Toarcian, Bifrons and Gradata Zones, the Upper Toarcian,
Reynesi Zone, the uppermost Lower/Upper Bajocian, the
Lower Callovian (Bullatus and Gracilis Zones), the Middle
and Upper Oxfordian (Transversarium, Bifurcatus, Bimammatum
and Planula Zones) and the Lower and Upper Kimmeridgian
(Platynota, Strombecki, Divisum and Beckeri
Zones). Some of these zones and subzones are recognized, or
documented with reported faunas, for the first time.
As a whole, benthic assemblages dominated during the
Lower Jurassic, while benthonic/planktonic assemblages

developed within the Middle–Upper Jurassic. Ammonite
taphonomy reveals the abundance of reworked assemblages
with common truncation, imbrication and coating (by Fe–Mn
oxides) of inner moulds or shells. Nevertheless, only occasionally
during the Domerian–Lower Toarcian interval of
alternating yellowish marly/silty limestones the faunal condensation
mixed biostratigraphically recognizable horizons.
The assemblages of ammonite faunas analyzed from the
Domerian to the Kimmeridgian show a Mediterranean character.
Thus, the bio-chronostratigraphic zonal/subzonal
scheme was applied, with minimal changes, for Mediterranean
Province (Cariou and Hantzpergue, 1997).
The interpreted paleoenvironmental evolution of the Jurassic
Malaguide at Sierra Espuña appears to be similar, and
comparable in timing, to other perimediterranean Tethyan
paleomargins. It evolves as a passive margin, with development
of shallow carbonate platform, until the Domerian
(Lavinianum Zone), when the platform break-up took place,
starting the rifting stage. During this stage in the Dogger, the
horst–graven system begins and the area was drowned at considerable
depth. Then, from the Lower Callovian to the Middle
Oxfordian, the drifting stage started, emphasizing the horst–
graven system with development of condensed nodular limestones
in the raised sea-floor.
Acknowledgements
This research was economically co-financed by the research
Projects BTE2001-3020, BTE2001-3029, BTE2000-
0299 and BTE2003-01113 (Spanish Ministry of Science and
Technology) and Research Groups GR00-222, GV04B-629
(Generalitat Valenciana) and RNM-178 (Junta de Andalucía).
We are grateful to Professor A. Jiménez (University of
Granada) for Photograph assistance. We are also indebted to
Mr. D. Nesbitt for the revision of the English text.
References
Azema, J., 1960. Sur la stratigraphie du Jurassique des environs de Málaga
(Espagne). Comptes Rendus de l’Académie de Sciences de Paris 250,
2231–2233.
Azema, J., 1961. Étude géologique des abords de Málaga. Estudios Geológicos
17, 131–160.
Braga, J.C., 1983. Ammonites del Domeriense de la Zona Subbética (Cordilleras
Béticas, Sur de España). Tesis doctoral. Secretariado de Publicaciones,
Universidad de Granada.
Caracuel, J.E., Sandoval, J., Martín-Martín, M., Estévez, A., Martín-
Rojas, I., 2001. Datos preliminares del Jurásico del perfil de Malvariche
en Sierra Espuña (Complejo Maláguide; Zonas Internas Béticas). XVII
Jornadas de Sociedad Española de Paleontología, 271–278.
Cariou, E., 1980. L’étage Callovien dans le Centre-Ouest de la France. Thèse
de Doctorat ès-Sciences, Université de Poitiers.
Cariou, E., Hantzpergue, P., 1997. Groupe Français d’Études du Jurassique,
biostratigraphie du Jurassique ouest-européen et méditerranéen. Zonations
parallèles et distribution des invertébrés et microfossiles. Bulletin
des Centres de Recherches-Exploration Production d’Elf-Aquitaine 17,
1–400.
J.E. Caracuel et al. / Geobios 39 (2006) 25–42
Catalano, R., Lo Cicero, G., Sullo, A., 2002. Geology of Sicily: An Introduction.
In: Santantonio, M. (Ed.), General Field Trip Guidebook. VI
International Symposium Jurassic Stratigraphy (12–22 Septiembre
2002). Palermo, Sicily, pp. 1–320.
Colacicchi, R., Parisi, G., Zamparelli, V., 1999. Bioevents and integrated
biostratigraphy of the Triassic and Jurassic in Italy. Palaeopelagos, Special
Publication 3, 1–184.
Doyle, P., MacDonald, D.I.M., 1993. Belemnites Battlefields. Lethaia 26,
65–80.
Fallot, P., 1929. Esquisse géologique du Massif de la Sierra Espuña (Prov. de
Murcia). Boletín de la Real Sociedad Española de Historia Natural 29,
199–215.
Fallot, P., 1931–1934. Essais sur la répartition des terrains secondaires et
tertiaires dans le domaine des Alpides espagnoles (Lias, Dogger, Jurassique
supérieur). Géologie de la Méditerranée occidentale 4, 1–118.
Fallot, P., 1945. Estudios geológicos en la Zona Subbética entre Alicante y el
río Guadiana Menor. Monografías del Consejo Superior de Investigaciones
Científicas, Instituto Investigaciones Geológicas “. Lucas Mallada
5, 1–719.
Fernández-López, S.R., 2000. Temas de Tafonomía. Departamento de Paleontología,
Universidad Madrid.
Geel, T., 1973. The Geology of the Betic of Málaga, the Subbetic, and the
Zone between these two units in the Vélez Rubio area (SE Spain). GUA
Papers of Geology 5 (1), 1–179.
Geyer, O.F., Hinkelbein, K., 1971. Eisenoolithische Kondensations-
Horizonte im Lias der Sierra de Espuña (Provinz Murcia, Spanien).
Neues Jahrbuch für Geologie und Paläontologie, Monatshefte 1971 (10),
398–414.
Geyer, O.F., Hinkelbein, K., 1974. Las Oolitas ferruginosas del Jurásico de
la Sierra de Espuña (prov. de Murcia). Acta Geológica Hispánica 9,
102–106.
Guerrera, F., Martín-Algarra, A., Perrone, V., 1993. Late Oligocene–Miocene
syn-/late-orogenic successions in Western and Central Mediterranean
chains from Betic Cordillerra to Southern Apennines. Terra Nova 5,
525–544.
Kampchuur, W., Langerberg, C.W., Baena, J., Velando, F., García-
Monzón, F., Paquet, J., Rondeel, H.E., 1974. Mapa y Memoria Explicativa
del Mapa Geológico de España 1/50 000, Hoja de Coy (932).
Instituto Geológico y Minero de España, 1–32.
Maate, A., 1996. Estratigrafía y Evolución Paleogeográfica alpina del
Dominio Gomáride (Rif Interno, Marruecos). PhD Thesys Universidad
de Granada.
Mac Gillavry, H.J., Geel, T., Roep, T.B., Soediono, H., 1963. Further notes
on the geology of the Betic of Málaga, the Subbetic, and the Zone
Between these two Units, in the Region of Vélez Rubio (Southern
Spain). Geologische Rundschau 53, 233–259.
Martín-Martín, M., 1996. El Terciario del Dominio Malaguide en Sierra
Espuña (Cordillera Bética oriental, SE España): Estratigrafía y Evolución
Paleogeográfica. PhD Thesys Universidad de Granada.
Martín-Martín, M., Martín-Algarra, A., 1997. La estructura del área de
Sierra Espuña (Contacto Zonas Internas–Externas, sector oriental de la
Cordillera Bética). Estudios Geológicos 53, 237–248.
Martín-Rojas, I., Caracuel, J.E., Estévez, A., Martín-Martín, M., Sandoval,
J., 2002. Jurassic stratigraphy of the Malaguide Complex from Sierra
Espuña (Internal Betic Zone, SE Spain). In: Gaillard, C., Hantzpergue, P.
(Eds.), STRATI 2002, 3 e Congrès Français de Stratigraphie, Lyon
8-10 septembre 2002. Documents du Laboratoire de Géologie de Lyon
156, Lyon, pp. 156–157.
Navarro, A., Trigueros, E., 1963. Estudio Geológico del Borde Oriental de la
Sierra Espuña. Notas y Comunicaciones Instituto Geológico y Minero de
España 70, 205–210.
Paquet, J., 1962. Contribution à l’étude géologique de la Sierra Espuña
(Prov. de Murcia, Espagne). Annales de la Société Géologique de Nord
32, 9–17.
41

42 J.E. Caracuel et al. / Geobios 39 (2006) 25–42
Paquet, J., 1969. Étude géologique de l’Ouest de la province de Murcia
(Espagne). Mémoires du Bureau de Recherches Géologiques et
Minières, Paris 48, 1–270.
Peyre, N., Peyre, Y., 1960. Observaciones Geológicas Sobre Sierra Espuña
(Murcia). Notas y Comunicaciones Instituto Geológico y Minero de
España 59, 3–23.
Sandoval, J., Checa, A., 2002. Taphonomy of cephalopods concentrations in
the Jurassic of the Subbetic (southern Spain). In: De Renzi, M., et al.
(Eds.), Current Topics on Taphonomy and Fossilization. Collecció
Encontres 5. pp. 223–230.
Sartorio, D., Venturini, S., 1988. Southern Tethys Biofacies. Agip, S. Donato
Milanese.
Sequeiros, L., 1974. Paleobiogeografía del Calloviense y Oxfordiense en el
Sector Central de la Cordillera Bética. PhD Thesys, Universidad de
Granada.
Seyfried, H., 1978. Der subbetische Jura von Murcia (Südost-Spanien).
Geologisches Jahrbuch 29, 3–201.
Vera, J.A., 1988. Evolución de los sistemas de depósito en el margen Ibérico
de la Cordillera Bética. Revista de la Sociedad Geológica de España 1,
373–391.
Villasante, F.B., 1912. Sierra Espuña y otros criaderos de menor importancia.
In: Criaderos de hierro de España. Memorias del Instituto Geológico
y Minero de España 26, pp. 525–544.
Zempolich, W.G., 1993. The drowning Succession in Jurassic Carbonates of
the Venetian Alps, Italy: A Record of Supercontinent Breakup, Gradual
Eustatic Rise, and Eutrophication of Shallow-Water Environments.
AAPG Memoir 57, Carbonate Sequence Stratigraphy, 63–105.
Zieten, C.H., 1830–1834. Die Versteinerungen Württenberg, Stuttgart.