Palaeoenvironmental analysis based on rugose corals

Blackwell Publishing Ltd

and microfacies: a case study at La Cornuda section

(early Serpukhovian, Guadiato Area, SW Spain)
ALBERTO GMEZ-HERGUEDAS AND SERGIO RODRGUEZ

LETHAIA Gmez-Herguedas, A. & Rodrguez, S. 2009: Palaeoenvironmental analysis based on

rugose corals and microfacies: a case study at la cornuda section (early Serpukhovian,
Guadiato Area, SW Spain). Lethaia, Vol. 42, pp. 3954

Combined analysis of microfacies and rugose coral features provides a useful tool
for palaeoenvironmental studies in areas where outcrops are not appropriate for field
observations. A detailed study of Serpukhovian rugose corals from La Cornuda section
(Guadiato Area, SW Spain) by means of thin sections allowed the identification of
environments where they lived. All corals were collected in unit 1 of the section where
three different but intimately connected environments have been identified. Corals
developed mainly in small mounds built jointly by microbial communities, algae and
corals. Some corals also lived in calcareous shoals mainly composed of echinoderm
plates. Finally, some corals occur in Oncoidal limestone that represents a shallow ramp,
but they were mainly transported from shoals and mounds.  Environment, microbial
mounds, Mississippian, oncoids, Rugosa, Serpukhovian, shoals.

Alberto Gmez-Herguedas [a.gomez@geo.ucm.es], Sergio Rodrguez [sergrodr@geo.ucm.es],

Departamento y UEI de Paleontologa, Facultad de Ciencias Geolgicas e Instituto de
Geologa Econmica, UCM y CSIC, 28040 Madrid, Spain; manuscript received on 24/10/
07; manuscript accepted on 23/01/08.

Studies on palaeoenvironments are usually based
both on facies analysis of sedimentary rocks in the
field and on microfacies analysis with thin sections.
The microfacies analysis constitutes the main tool
in areas where outcrops are not appropriate for field
observations. We applied this method in the sparse
outcrops from the San Antonio-La Juliana Unit
(Guadiato Area, Czar & Rodrguez 1999, 2004).
Additionally, we applied detailed palaeoecological
and taphonomical studies on rugose corals. The
present paper shows the results of such a study. The
level of environmental information in this case is
high because of a detailed sampling of available
outcrops and preparation of a large number of thin
sections of corals and the enclosing host carbonate
sediment. Palaeoecological studies include auto-
ecological analysis and synecological analysis based
on the method of Neuman (1988), Vuillemin (1990),
Rodrguez (2001), Rodrguez & Gmez-Herguedas
(2002) and Somerville & Rodrguez (2007). Tapho-
nomical analysis of rugose corals followed the method
proposed by Rodrguez (2004).
The Guadiato Area is a large and complex area in
SW Spain (Fig. 1) that comprises Mississippian
siliciclastic and limestone rocks. It is divided into
three units (Czar & Rodrguez 1999): (i) Fresnedoso
Unit, composed mainly of Visan siliciclastic rocks;
(ii) Sierra del Castillo Unit, composed mainly of
Visan limestone rocks; and (iii) San Antonio-La
Juliana Unit, composed of Serpukhovian siliciclastic
and limestone rocks.
Studies on the Guadiato Area go back to the 19th
century (Yegros & Snchez 1850; Mallada 1880);
most palaeontological and stratigraphical studies
were focused on the Visan units. This is especially
marked for studies on rugose corals (Rodrguez 1985,
2004; Rodrguez & Falces 1996; Falces 1997, 1998;
Hernando 2000; Rodrguez-Curt 2000; Rodrguez
et al. 2001a, b, 2002, 2003, 2004, 2007; Somerville &
Rodrguez 2003, 2007; Rodrguez & Somerville 2007;
Rodrguez & Hernando 2005), but corals from the
Serpukhovian San Antonio-La Juliana Unit have
been only recently studied (Ramrez-Viu 1994;
Rodrguez et al. 1996; Falces & Rodrguez 2002;
Gmez-Herguedas 2003, 2006; Gmez-Herguedas &
Rodrguez 2003, 2005, 2006, 2007).
La Cornuda section
The La Cornuda Section was first measured by
P. Czar, S. Rodrguez and A. Calvo in 1994. Czar
(1998) described the section in detail and analysed
the age and palaeoenvironment of each unit based
on algae, foraminifera and microfacies. He identified
the foraminifer zones 17 and 18 of Mamet (1974),
included in the Pendleian stage (Lower Serpukhovian).
The section is located about 3 km northwest from

DOI 10.1111/j.1502-3931.2008.00106.x 2008 The Authors, Journal compilation 2008 The Lethaia Foundation
40 A. Gmez-Herguedas & S. Rodrguez LETHAIA 42 (2009)

Fig. 1. Location of the studied outcrop: A. Iberian Peninsula. B. Carboniferous outcrops in the boundary of Ossa-Morena and
Central-Iberian zones. 1. Berlanga, 2. Matachel, 3. El Casar, 4. Campillo de Llerena, 5. Benajarafe, 6. Guadiato-Guadalmellato, 7. Guadalbarbo,
8. Pedroches. C. Geological map of the northwestern part of the Guadiato Area with location of La Cornuda section. (Modified from
Czar and Rodrguez 2004).

Espiel, near La Cornuda Farm. The coordinates at
the base of the section are 5630 west and 381240
north. The total sequence is about 500 m in thickness,
but only a few limestones, calcareous sandstones and
conglomerates crop out (Fig. 2); tilled fields cover
most beds. Czar (1998) and Czar & Rodrguez
(2004) regarded the whole sequence as originated in
a shallow-water platform where the bioclastic
limestone, calcareous sandstone and conglomerate
represent mainly shoals and the mostly covered units
(lutites, limolites and marls, 90% of the section)
represent the surrounding lagoonal and shallow-
water platform facies.
Our study has been focused on the Unit 1, the
oldest limestone bed in the section, because it is the
richest in corals and shows several facies in the field,
providing complex information that allows the vali-
dation of the method. It is 1 to 3 m thick, showing
discontinuous outcrops. We have analysed 58 coral
specimens including solitary and colonial corals with
more than 130 thin sections. Fifty-seven specimens are
embedded in the rock and thin sections show micro-
facies in what they were sedimented. We also
studied 20 thin sections of limestones in different
parts of the unit in order to check other components
of the rock.
Three main facies have been identified:
1 The most common facies is bioclastic limestone
(50%) composed of crinoid, echinoid, bryozoan
and brachiopod fragments. Rugose corals (both
entire and broken) and chaetetids sponges are
common. The microfacies is echinoderm grain-
stonepackstone. Most echinoderms are broken
and rounded. Other minority components are
bivalves, trilobites, gastropods, foraminifers, ung-
darellaceans, Girvanella, lithoclasts, ooids and
cortoids. Sparite is more abundant than micrite.
Selection of bioclasts is poor; most bioclasts are
small (mode is smaller than 0.5 mm in size, but
some brachiopods and corals are entire or almost)
(Fig. 3A).
LETHAIA 42 (2009)
Fig. 2. Stratigraphical section at La Cornuda with location of Unit
used for the case study.
2 The bioclastic limestone passes laterally to bio-
constructed beds containing abundant rugose
corals. Microfacies is micropeloidal mudstone
wackestone that contains common rugose corals,
mostly in growth position and rarely reworked
(both solitary and colonial). The algae Falsocalci-
folium punctatum (Maslov) is locally abundant
and it shows foliose structure and flabellum-
shaped cups (Czar & Vachard 2004). Tabulate
corals are present too, but they are less common
than rugosans. Fragments of brachiopods, bivalves,
bryozoans, crinoids, gastropods and foraminifers
occur in the wackestone portions of the rock.
Stromatactoid cavities and micropeloidal textures
are common in this microfacies (Fig. 3B).
3 Oncoidal packstone. Oncoids are mainly composed
of Girvanella (calcimicrobe) and Sparaphralysia
Palaeoenvironment form rugosans and microfacies 41
Fig. 3. Microfacies types at Unit 1. A. Echinoderm grainstone
packstone (COR/1-130M1). B. Micropeloidal mudstone-wackestone
showing cavities and the problematic alga Falsocalcifolium
punctatum (Maslov) (COR/1-64M2). C. Oncoidal packstone
(COR/1-131M1). Scale bars = 2 mm.
(problematicum) and reach 5 cm in diameter
(Czar et al. 2003a). Nucleii of oncoids are usually
broken brachiopods, gastropods, and tabulate and
rugose corals. Sediment around oncoids is mainly
micritic and contains common echinoderm plates,
bryozoans and foraminifers. All corals in this
microfacies are reworked (Fig. 3C).
42 A. Gmez-Herguedas & S. Rodrguez
Palaeoecological analysis
Our palaeoecological study comprises two different
aspects.
1 A palaeoautoecological analysis focused on the
morphology of the corals and on surrounding
bioclasts and sediment in order to evaluate the
relationships of those corals with other organisms
and with the environment.
2 A palaeosynecological study, focused on the coral
assemblages in each sub-environment in order to
evaluate the structure of the ecosystem.
Palaeoautoecology
Our palaeoautoecological analysis has been carried
out following the methods proposed by Vuillemin
(1990), Scrutton (1998) and Rodrguez & Gmez-
Herguedas (2002). The information on the ecology
of the studied corals comes from external and
internal features of corals. The relationships of
corals with their environment have been studied by
means of an analysis of external shape, attachment
processes, crusts, colonizations and bioerosion
phenomena. Additional information on the coral
life-environment and the burial-environment was
obtained with the study of the level of biodegradation,
skeletal fragmentation, wear surfaces, etc.
External shape of corals
Colonial corals. Nineteen specimens (32.7% of the
total number) are colonial; 5.2% are massive speci-
mens, 12.1% are fasciculate and 15.5% show incipient
colonialism (common calicular offsets, but do not
develop large colonies).
Colonial corals lived usually in shallow, clean waters.
Consequently, the significant percentage of colonial
corals indicates a shallow-water environment. The
dominance of fasciculate and protocolonial corals
(Diphyphyllum fasciculatum, Diphyphyllum gracile,
Guadiatia pseudocoloniale Gmez-Herguedas &
Rodrguez, 2005 and unidentified lithostrotionid)
and the scarcity of massive corals indicate that the
environment was not rough, because the former
cannot tolerate very agitated waters. In contrast to
the fasciculate corals that occur in mudstone
wackestone, the only massive coral (Lithostrotion
maccoyanum) has been located in grainstone; it
seems to indicate that it lived in a different environ-
ment. But the micritic calicular filling in this species
indicate that the studied specimen is reworked and
LETHAIA 42 (2009)
it lived in the same environment as the fasciculate
species (see below). Therefore, all colonial species
lived in micropeloidal mudstones.
Solitary corals. They constitute the 67.2% of speci-
mens (39 in total) at La Cornuda. The external shape
provides detailed information on the coral life-
strategies (Neuman 1988; Rodrguez 2001) and on
the substrate (Rodrguez & Gmez-Herguedas 2002).
Ceratoid and turbinate or trochoid corals, straight or
slightly curved, are abundant in La Cornuda. These
external shapes indicate a liberosessile strategy. The
corals lived on the sediment surface in soft or firm
substrates, but they needed large bioclasts for attach-
ment of planulae. On the other hand, curved ceratoid
or trochoid specimens are very uncommon in La
Cornuda and rarely show thickenings in cardinal
quadrants that were used for stability on soft bottom
sediment. It indicates a partly buried life position in
soft bottom (Neuman 1988; Rodrguez et al. 1997;
Rodrguez 2001) where some clasts are available for
attachment of planulae. No significant differences
have been observed in the external shape of solitary
corals from each microfacies. Thus, the substrate
in all of them was firm to soft, but enough bioclasts
were available in all of them for attachment of
planulae.
Attachment processes
The main types of attachment structures in La Cornuda
are radiciform processes and talons (Rodrguez 2004).
Only one axophyllid specimen from La Cornuda
shows radiciform processes (Fig. 4A). It is located
in the oncoidal microfacies, rounded by echinoderm
plates and very close to the bioclastic limestone.
Consequently, it could live close to or within
bioclastic grainstonepackstone. Most commonly,
specimens of aulophyllids show talons (Fig. 4B). It
demonstrates that the substrate could be soft or firm,
but common clasts were available for attachment
or reinforcement of growth. Absence of attachment
processes in most specimens indicates that the
substrate should be firm but not soft. All specimens
having talons occur in the microbial microfacies or
come from it. Thus, it proves that the environment
provided common bioclasts for attachment in a
mainly muddy substrate.
Skeletal anomalies produced during the
coral life
Skeletal anomalies give much information on the
interaction of rugose corals and the environment
(including other organisms). Falces (1998) described
LETHAIA 42 (2009) Palaeoenvironment form rugosans and microfacies 43

Fig. 4. Attachment structures (arrowed). A. Radiciform process in axophyllid coral (COR/1-24, unidentified axophyllid sp.3). B. Talon
in aulophyllid coral (COR/1-68, Guadiatia pseudocoloniale). Scale bars = 1 mm.

Fig. 5. Skeletal anomalies. A. Rejuvenescence (arrowed) in longitudinal section of aulophyllid coral (COR/1-62, Guadiatia pseudocoloniale).
B. Wall deformation (arrowed) in transverse section of aulophyllid coral (details in the text) (COR/1-61, G. pseudocoloniale). Scale
bars = 1 mm.

five types of anomalies in rugose corals: rejuvenescence,

pathological deformation, anomalous growth of skeletal
structures, necrosis and syngenetic fractures.
Rejuvenescence occurs when environmental con-
ditions are not favourable and corals need to restrict
their metabolic requirements; then, they reduce their
size (Vuillemin 1990). When environmental condi-
tions are irregular, rejuvenescences became common.
They are rare in La Cornuda; thus, the environment
in La Cornuda was quite stable, but the presence of
scarce examples (only three specimens, Fig. 5A) indi-
cates that the environment suffered sporadic variations.
Only one case of skeletal anomaly in the wall has
been identified (Fig. 5B). It could be caused by some
kind of pathology or an external aggression, not
related with general environmental features.
External wall
External wall of corals is directly related to the
environment. Consequently, the features of the wall
could be controlled by some environmental factors.
However, a systematic analysis of walls in La Cornuda
corals shows that the wall morphology is directly
related to genetic but not environmental factors: (i)
cyathopsids and lithostrotionids (27.6%) show simple,
mainly smooth walls; (ii) aulophyllids (29.3%) possess
simple to slightly festooned walls; and (iii) axophyllids
(37.9%) show festooned to undulated walls.
However, thickness of walls seems to be related
to the environment. Those specimens that occur in
grainstone or packstone microfacies show thicker walls
than those occurring in mudstones or wackestones.
44 A. Gmez-Herguedas & S. Rodrguez
Consequently, thick walls seem to be related with
high-energy environments, as could be expected.
Dissepiments
The presence of dissepiments in rugose corals is a
conspicuous feature that has notable relationships
with environmental requirements; undissepimented
corals (Cyathaxonia fauna) are small, simple and typic-
ally lived in quiet, deep or turbid environments (Hill
1938; Sando 1980; Kullmann 1997). On the other hand,
dissepimented corals can reach larger size and lived in
clean, shallow waters and probably were related to
some kind of symbiotic algae (Vuillemin 1990).
Most corals from La Cornuda have dissepiments and
only one specimen of laccophyllid should be included
in the Cyathaxonia fauna. Consequently, the assem-
blage from La Cornuda lived in clean and shallow
waters. It agrees with observations on the whole com-
munity with abundant photosynthetic organisms and
the common encrusting algae and cyanobacteria.
Vuillemin (1990) stated that the type of dissepiments
is partly controlled by environmental factors. So,
this author claims that regular dissepiments are well
adapted to high-energy environment but lonsdaleoid
dissepiments are fragile and occur in low-energy
environments. Our observations in La Cornuda do
not agree with such hypothesis, because corals having
lonsdaleoid dissepiments (axophyllids, Melanophyllum?)
have thick walls and they are common in microfacies
produced in high-energy environments (26%).
Septa
Thickness and abundance of septa (that is to say,
skeletal density) are directly related to the energy-
level of environment if they are not located in precise
quadrants of the coral (Vuillemin 1990; Rodrguez
2001). Conspicuous thickenings of septa are not
common in La Cornuda. Only three specimens
show general thickening of septa.
Axial structure
Axial structures produce a high density of elements
in the axial zones of corals, giving them more resist-
ance to fragmentation and compression. So, they are
useful in high-energy environments. Vuillemin (1990)
stated that corals having axial structure (aulophyllina,
lonsadaleiina, lithostrotionina) were better adapted
to turbulent, shallow environments and that corals
lacking an axial structure should live in quiet water
environments. The assemblages from La Cornuda show
coexistence of rugose corals with and without axial
structures in microfacies produced in turbulent environ-
LETHAIA 42 (2009)
ments, as previously observed by Somerville & Rodrguez
(2007), indicating that absence of axial structure does
not imply low-energy environment. However, it is true
that corals missing axial structure, such as cyathopsids
are more common in muddy, quiet environments.
Tabulae
The tabularium is an important part of the skeleton
in rugose corals. Other elements such as dissepiments
may be absent in some rugosans, but tabulae occur
almost in all rugose coral genera. Being an internal
structure, tabulae should not be closely controlled by
environmental factors. However, density of tabulae
(number of tabulae/centimetre) has been regarded
as related to the growth rate (Vuillemin 1990). The
cited author stated that the time for secretion of
tabulae was regular for each species; so, changes in
the growth rate imply differences in the density of
tabulae. When rugose corals grew quickly, tabulae
became widely spaced. When rugose corals grew
slowly, they became very densely packed. The conse-
quence is that we can identify periods of favourable/
unfavourable environment for the development of
corals. Moreover, alternation of bands with densely
packed/sparse tabulae indicates a strong seasonality
in the environment related to temperature, sedimen-
tation rate or water turbidity.
Most if not all specimens from La Cornuda show
quite low density of tabulae. In addition, there is no
banding in the tabulae density. It implies that envi-
ronment was favourable for quick growth and there
were little seasonal changes; it points to a subtropical
clime and little influx of siliciclastic sediment
throughout each year.
Palaeosynecology
Encrustation and colonization
Encrusting and colonizing organisms (epifauna) are
common in marine environments and particularly in
shallow water. Epifauna leave clear imprints on the
surface of external skeletons such as that of rugose
corals. They give precise information on an important
part of the community where corals lived. Conse-
quently, the study of those encrustations and colon-
izations provide valuable data on the environment.
Two kinds of epifauna can be distinguished (Falces
1998): epibionts that colonize the skeleton while they
live in areas abandoned by the polyp, and epiliths that
colonize the skeletons after the coral is dead. Distinguish-
ing epibionts from epiliths is difficult when colonization
took place on the external wall, but epiliths are easily
LETHAIA 42 (2009) Palaeoenvironment form rugosans and microfacies 45

Fig. 6. Colonization on corals. A. Alternation of Girvanella and Sparaphralysia encrustations on axophyllid wall (COR/1-15,
unidentified axophyllid sp. 3). B. Successive encrustations of bryozoans and Girvanella on diphyphyllid wall (COR/1-3, Diphyphyllum?
sp.). C. Post-mortem encrustation of bryozoan on axophyllid wall and calice (COR/1-59, unidentified axophyllid sp. 3). D. Auloporoid
corallites attached to aulophyllid coral (COR/1-45, Guadiatia pseudocoloniale) and cyanobacteria crust. Scale bars = 1 mm.

identifiable when they took place on the calices or
covering post-mortem alterations of the skeleton.
Encrustations and colonizations are very common
in La Cornuda, affecting about 85% of specimens.
Colonizers are diverse:
Cyanobacteria and calcareous algae. They are the
most common encrusting organisms, affecting more
than 65% of specimens. They usually occur as partial
crusts on lateral surfaces of the corals, but some
specimens became totally covered by several layers of
cyanobacteria that may alternate with other encrusters
(e.g. bryozoans or chaetetids). The main colonizers
are Girvanella and Sparaphralysia, but there are also
crusts of Fasciella, codiaceans and indeterminate algae
(Czar 1998). Girvanella is the most common encrust-
ing organism and it is represented by three different
species. It can occur alone but more commonly com-
bined with Sparaphralysia (Fig. 6A) (Czar et al. 2003a).
Algae and cyanobacteria colonize the coral walls in
most cases. We identified some cases of encrustations
during the life of the coral. They are characterized
by thickness reduction towards the upper part of
the coral and absence of crusts in the calice. Other
encrustations took place after the death of the polyp
because they occur both, on the wall and on the calice
without conspicuous variation in thickness.
Some specimens are totally covered by several
layers of Girvanella, Sparaphralysia and other encrust-
ing organisms forming large cyanoliths similar to
those described by Czar et al. (2003a).
Some specimens are rounded by micropeloidal
texture that indicates microbial activity. Usually corals
from La Cornuda grew in favourable conditions
when microbial communities were developed around
them and even built small mounds together, but in
some cases they show irregularities in the wall possibly
caused by relation with such communities.
Bryozoans. Encrusting bryozoans are also common
in La Cornuda affecting about 40% of specimens.
They occupy wide areas of the wall but rarely is the
encrustation complete. When bryozoans cover the sur-
face of the coral walls cyanobacteria and algae tend
to be absent; if present, the encrustation of bryozoans
is commonly earlier (Fig. 6B). The explanation is that
bryozoans cannot encrust easily on the irregular and
probably soft surface provided by cyanobacteria and
algae. Bryozoans in La Cornuda colonized the coral walls
before and after the death of the polyp (Fig. 6C).
46 A. Gmez-Herguedas & S. Rodrguez LETHAIA 42 (2009)

Other encrusting organisms. Tabulate corals (syring-

oporoids), chaetetids and foraminifers are less com-
mon than other colonizers in La Cornuda but they
occur encrusting corals directly or encrusting layers
of cyanobacteria (Fig. 6D).
From all previous data it is clear that colonization
and incrustation took place during the life of the
corals (mainly) and after they died, before burial. But
incrustation continued even if coral skeletons were
remobilized after burial, indicating a high activity of
encrusting organisms.
Encrustation show different patterns in each facies.
Thus, all coral specimens in echinoderm grainstone
packstone are encrusted and in most cases there are two
layers of different encrusting organisms, but encrusta-
tions never reach thick development and rarely do
they cover the whole specimen. Cyanobacteria encrust
75% of coral specimens and they are combined with
algae (Sparaphralysia), chaetetids, bryozoans and
syringoporoids (from more to less abundant). Specimens
from mudstonewackestone show also high level of
encrustation, but lower than in other facies (80%).
Encrustations are also multiple in most cases, but
the covered surface area commonly used to be small.
Girvanella is the most common encrusting organism
(65.6%), but in this case bryozoans are also abundant
(56.2%). Encrusting organisms are more diverse than
in other facies: Sparaphralysia, other algae, foraminifers
and syringoporoids. However, all these encrusting
organisms occur rarely (less than 13%).
Obviously, all specimens occurring in the oncoidal
packstone are also encrusted. Encrustations here are
complete and very thick, but encrusting organisms
are less diverse. All oncoids are composed of Girvanella
and Sparaphralysia with lower participation of
bryozoans (40% of specimens) and other algae (30%).
Corals are commonly encrusted by different organ-
isms, but they also are attached to other organisms
(skeletons or not). They used a great variety of ele-
Fig. 7. Coral colonizations. A. Aulophyllid corallite attached to
ments for attachment: bivalves, brachiopods, masses an algal mat and subsequently covered by a bryozoan (COR/1-10,
of Girvanella and Sparaphralysia (Fig. 7A), chaetetids Guadiatia pseudocoloniale). B. Chaetetid-coral interaction
(Fig. 7B), ooids or algae (Fig. 7C). (COR/1-14, Axophyllum aff. pseudokirsopianum). C. Lithostro-
tionid corallite attached to a stacheininae alga (centre arrow) and
to an ooid nucleated on a small foraminifer (bottom left arrow)
(COR/1-41, lithostrotionid indet.). Scale bars = 1 mm.
Bioerosion
Some organisms bore the mineralized skeletons of
other organisms by means of mechanical systems, endolithic algae, and (ii) mesoscopic borings ellipsoidal
activity of organic acids, etc. These organisms pro- in shape similar to Gastrochaenolites (Fig. 8A).
duce borings that are conspicuous in the coral skele- When a living coral is bored, the polyp produce
ton. Bioerosion takes place mainly when skeletons changes in the skeleton in order to repair the dam-
are exposed for a long time before burial, indicating aged parts. No reparations have been found in the
a low sedimentation rate. bored corals from La Cornuda; so we assume that
Borings are scarce in La Cornuda, affecting only they were bored after the death of polyp.
about 20% of specimens. Two types of borings have Bioerosion and encrustations (cyanobacteria,
been identified: (i) microscopic borings produced by calcareous algae and bryozoans) may occur in the
LETHAIA 42 (2009)
Fig. 8. Bioerosion structures. A. Boring of Gastrochaenolites-type on axophyllid wall (COR/1-23, unidentified axophyllid sp. 3).
B. Borings cutting axophyllid wall (left) and axophyllid wall and algal crust (right) (COR/1-21 unidentified axophyllid sp. 2). Scale
bars = 1 mm.
same coral specimens at La Cornuda. When it
happens, it is easy to identify the order of the phe-
nomena. In the few examples from this section,
borings may cut encrustations, but encrustations
rarely cover borings. Thus, bioerosion may occur later
or simultaneously with encrustations, but probably
very rarely or not before it (Fig. 8B). The scarcity of
borings may be explained by a quite quick sedimen-
tation (but not enough for avoid them totally), which
also explain the scarcity of post-mortem encrustations.
Only 13 specimens show borings. The distribution
in facies is: six specimens in microbial mudstone
wackestone, two specimens in echinoderm grain-
stonepackstone and five specimens in oncoidal
packstone. Some specimens from oncoids seem to
come from mudstones; thus the boring activity was
mainly developed in the microbial facies (mounds).
Taphonomical analysis
For the taphonomical analysis of La Cornuda corals,
we followed the protocol proposed by Rodrguez
(2004), but only biostratinomic procceses were taken
into account for the identification of the environment.
Biodegradation
Organic matter that constitutes the soft parts of
organisms is quickly decomposed by activity of
aerobic bacteria. Only if the sedimentation rate is
very high (blocking the activity of bacteria) and/or
Palaeoenvironment form rugosans and microfacies 47
the environment is anaerobic will the organic decay
process not go to completion and produce sulphide
mineralization. In the case of corals, such mineraliza-
tion can only be located in calices, where the polyps
lived. After a detailed investigation, we can state that
no traces of sulphide minerals such as pyrite or
marcasite occur in La Cornuda. Therefore, all
environments were typically aerobic.
Sedimentary filling
Analysis of sedimentary filling of fossils is very useful
for identification of reworking processes. But coral
growth is very peculiar in the organic world. They
use a great quantity of material and energy building
their skeleton and leave much empty space in the
abandoned areas. However, that space cannot be
filled by sediments because it is closed to the exterior
by walls, dissepiments and tabulae. The skeleton of
corals is filled with surrounding sediment only if it is
broken. Even in this case, the filling is partial because
cementation of inner spaces is quick (Rodrguez
2004). The only space of the coral skeleton that is
completely filled by surrounding sediment is the
calice. Nevertheless, the analysis of sedimentary
filling of corals has provided interesting results.
Calices from La Cornuda show usually the same
textures as that surrounding sediment. In some cases,
the abundance and size of bioclasts are lower in
calices because large particles cannot penetrate in
them because the presence of septa and other struc-
tures there. We found only one case of sedimentary
48 A. Gmez-Herguedas & S. Rodrguez
Fig. 9. Reworking processes in Lithostrotion maccoyanum (COR/1-3). A. The coral was first buried in micritic sediment proved by the
micritic filling of calices (M). Subsequently it suffered subaerial exposure that produced a layer of iron oxide (F) and finally it was buried
in grainstone (G). B. This section shows an additional encrustation by bryozoans (B) subsequent to the first burial and previous to the
subaerial exposure. Scale bars = 1 mm.
filling that does not fit with the surrounding sediment.
In this case the sedimentary filling is mudstone
wackestone and the surrounding sediment is
packstone or grainstone. The specimen is a massive
colony of Lithostrotion (Fig. 9AB). Calices are covered
with micritic sediment containing very scarce bioclasts;
the specimen is partly fragmented and fractures are
filled with sediment having grainstone texture and
encrusting bryozoans cover the grainstone filling.
Additional layers of grainstone cover the bryozoans
that show a thin layer of oxides. These features show
a complex taphonomic story:
The coral lived in a mostly quiet-water environ-
ment where sedimentation of calcareous mud was
dominant (probably a microbial mound, see discus-
sion). It was buried by muddy sediment and calices
were filled. A sporadic but strong event of high
energy displaced the colony to another environment
where continuous waves prevented the sedimentation
of mud and partly fragmented and eroded the coral
(probably a calcareous sand shoal, see discussion);
eventually it suffered sub-aerial exposure. It became
buried again. A new event of high energy displaced
the coral from sediment and bryozoans colonized
some surfaces. A new burial in calcareous sand shoal
was definitive.
Transport evidences
Existence of transport evidences in the coral skeleton
indicates that the coral was moved. Knowledge on
remobilization of corals is an important tool for
locating the origin of the palaeoecological informa-
tion provided by the corals structures, because we
can identify if such information is valid for the
sediment where the coral was buried. Two main
LETHAIA 42 (2009)
types of transport evidences have been identified in
La Cornuda.
Abraded surfaces
This feature indicates a pulling transport. They are
very scarce in la Cornuda, affecting only to 17% of
specimens (Fig. 10A). Only one specimen comes from
bioclastic grainstonespackstones, five specimens come
from mudstoneswackestones and four specimens
come from oncoidal packstones. Thus, the surfaces
of abrasion were produced mainly in situ (in the
mounds) and during transport from mounds to
deeper-water environments.
Fragmentation
Pre-burial fragmentation in corals may be produced
by transport or by impact of transported bioclasts
against the fixed corals. Most specimens from La
Cornuda are fragmented in diverse degrees (70%).
In many instances, the fractures have been pro-
duced before cementation of interskeletal spaces,
producing a filling of the holes that are close to the
fracture zone with surrounding sediment, the
fractures are small and skeletal elements are mostly
preserved (Fig. 10B). In this case, the partial preser-
vation of broken structures indicates that transport
was not long. In other instances fractures are larger
and some skeletal structures are missed (Fig. 10C),
indicating a longer turbulent event.
Field observations indicate that about 1/4 of
broken corals were located in life position or almost.
These fractures were produced by impact of trans-
ported bioclasts or by bioerosion. On the contrary, most
fractures are clearly due to transport in turbulent
LETHAIA 42 (2009)
Fig. 10. Transport evidences. A. Abrasion surface prior to
encrustation (top centre); diphyphyllid corallite (COR/1-36,
Diphyphyllum? sp.). B. Peripheral fragmentation in cyathopsid
coral (COR/1-35, Lublinophyllum?). C. Intense fragmentation
of an axophyllid coral (COR/1-55, unidentified axophyllid sp. 3).
Scale bars = 1 mm.
regime. Consequently, at least 50% of specimens
from La Cornuda were transported. However, frag-
mentation is not extensive indicating that transport
was short.
Palaeoenvironment form rugosans and microfacies 49
Fragmentations affect both, colonial and solitary
corals, and are conspicuous even in massive corals.
Assemblages
As stated above, a detailed analysis of sediment sur-
rounding and filling each specimen provided the
identification of three main microfacies related to the
corals from La Cornuda: (i) echinoderm grainstone
packstone, (ii) microbial mudstonewackestone,
and (iii) oncoidal packstone. The identification of
three different microfacies in the same lithological
unit allowed the recognition of three coral
assemblages:
1 Most corals were recorded in microbial mud-
stonewackestone despite it being less common
in the field than echinoderm grainstonepack-
stone. Moreover, about 50% of corals located in
this microfacies are in life position.
2 Many, but not all specimens recorded in echino-
derm grainstonepackstone are remobilized despite
the fact that it indicates turbulent conditions.
This is inferred of field observations and analysis
of fillings and relationships with other organisms
(e.g. as chaetetids, algae and bryozoans). In addi-
tion, some other specimens show a short distance
transport. Thus, some rugose corals lived in that
microfacies.
3 All specimens occurring in oncoidal packstone
are transported and some of them were moved
for a long time because they show thick oncoidal
binding. They are, by far, less common than in
the two first microfacies.
Some features in the studied thin sections show
that these microfacies were closely related:
1 Small bioclasts in the matrices of all microfacies
are similar (crinoids, brachiopods, bivalves,
bryozoans, gastropods, foraminifers).
2 Some thin sections show more than one micro-
facies; several sections show development of
micropeloidal mudstone on the surface of the
grainstone, indicating that microbial communities
could develop quickly when turbulence diminished.
In addition, some corals included in grainstone
show micropeloidal fillings or coatings.
3 Some corals partly coated by cyanobacteria and algae
are included in grainstones. Moreover, some oncoids
include fragments of grainstone in their nucleii.
The analysis of identified taxa and original micro-
facies is shown in Table 1. When those data are
50 A. Gmez-Herguedas & S. Rodrguez LETHAIA 42 (2009)

Table 1. Distribution of taxa in the different microfacies. Remobilized specimens have been located in the original microfacies.
M1) Echinoderm grainstone-packstone. M2) Micropeloidal mudstone-wackestone. M3) Oncoidal packstone.

Taxa Microfacies 1 Microfacies 2 Microfacies 3

Unidentified laccophyllid 1
Haplolasma lamelliferum 1
Lublinophyllum? sp. 1
Melanophyllum? sp. 1 1 1
Melanophyllum? sp. 2 1
Melanophyllum? sp. 3 1
Melanophyllum? sp. 4 1
Aulokoninckophyllum? sp. 1
Clisiophyllum benziregense 1
Dibunophyllum dobroljubovae
Arachnolasma? sp. 1
Amygdalophyllum cornudensis 1 3
Guadiatia pseudocoloniale 9
Lithostrotion maccoyanum 3
Diphyphyllum fasciculatum 1
Diphyphyllum gracile 2
Diphyphyllum? sp. 1 1
Unidentified lithostrotionid 1
Axophyllum aff. pseudokirsopianum 2
Unidentified axophyllid sp. 1 2 1
Unidentified axophyllid sp. 2 2
Unidentified axophyllid sp. 3 1? 10 4
Incertae sedis 1 1
Incertae sedis 2 1

analysed and compared with previous observations
on thin sections, we can reach some interpretations:
1 Most identified species lived in an environment
where mudstones were produced indicating
favourable life conditions.
2 Some taxa (Melanophyllum? sp. 1, Amygdalophyl-
lum cornudensis Gmez-Herguedas & Rodrguez,
2005) lived in two different environments,
because they occur in mudstoneswackestones
and in grainstonespackstones without clear signs
of remobilization.
3 Some other taxa occur only in echinoderm
grainstonespackstones (Melanophyllum? sp. 2,
Axophyllum aff. pseudokirsopianum, unidentified
axophyllid sp. 2).
4 Several taxa (Lublinophyllum? sp., Melanophyllum?
sp. 4, Aulokoninckophyllum? sp., Clisiophyllum
benziregense) do not provide any information,
because they are always transported and could
not be related to a precise microfacies.
5 One taxon (Dibunophyllum dobroljubovae) does
not provide any information because it does not
have sediment around it.
As stated above, transport of corals from La Cornuda
was generally short and it was produced during high
energy events such as storms (chaotic distribution of
clasts and erosive surfaces prove it). Thus, the corals
recorded in microfacies 3 that are not present in
microfacies 1 and 2 probably lived in those environ-
ments or near. Moreover, a percentage (18.7%) of
corals from microfacies 3 occur also in microfacies 2,
indicating a close relationship between environments
in which both facies were sedimented.
The coral assemblage in echinoderm grainstone
packstone is not very diverse. The species recorded
there that do not show evidences of long transport
are Melanophyllum? sp. 1, Melanophyllum? sp. 2,
Amygdalophyllum cornudensis Gmez-Herguedas &
Rodrguez, 2005, Axophyllum aff. pseudokirsopianum
and unidentified axophyllid sp. 2. All of them (eight
specimens) are solitary dissepimented and they
possess thick inner structures. Chaetetids occur also
in this assemblage and are directly related to corals,
growing on the corals and forming also the base for
coral growth.
On the other hand, the coral assemblage from
micropeloidal mudstonewackestone is highly diverse
and is composed both of solitary (dissepimented and
undissepimented) and colonial corals (incipient,
fasciculate and massive). Rugose corals are randomly
distributed but with an evident domination of solitary
dissepimented over incipient, fasciculate, massive
and solitary undissepimented forms. It is dominated
by two endemic genera, Guadiatia pseudocoloniale
Gmez-Herguedas & Rodrguez, 2005, and uniden-
tified axophyllid sp. 3. Other taxa are unidentified
laccophyllid, Haplolasma lamelliferum, Melanophyl-
lum? sp. 1, Melanophyllum? sp. 3, Arachnolasma? sp.,
LETHAIA 42 (2009) Palaeoenvironment form rugosans and microfacies 51

Amygdalophyllum cornudensis Gmez-Herguedas

& Rodrguez, 2005, Lithostrotion maccoyanum,
Diphyphyllum fasciculatum, Diphyphyllum gracile,
Diphyphyllum? sp., unidentified lithostrotionid,
unidentified axophyllid sp. 1 and rugosae Incertae sedis.
This assemblage (39 specimens) is a very specialized
one, with most taxa being endemic and not found in
any other locality. In addition, most colonial corals
show very small corallites.
The coral assemblage in oncoidal packstone (10
specimens) is less diverse: Lublinophyllum? sp.,
Melanophyllum? sp. 4, Aulokoninckophyllum? sp.,
Clisiophyllum benziregense, Diphyphyllum? sp. and
unidentified axophyllid sp. 1 and 3. It does not
constitute a part of a community because the corals
are transported. However, we found in this environ-
ment some taxa that are not present in the other two
environments. Perhaps some of them would live in
this unstable environment and probably in somewhat
deeper zones.

Discussion
Observations and inferences presented previously
show a complex sedimentary unit where three closely
related environments have been identified (Fig. 11A).
Environment 1
Fig. 11. A. General model of the sedimentary environment at
the unit 1 in La Cornuda section. B. Community structure in
calcareous shoals. 1. Melanophyllum? sp. 2. Amygdalophyllum corn-
udensis. 3. Axophyllum aff. pseudokirsopianum. 4. Unidentified
axophyllid sp. 2. 5. Chaetetid sponges. 6. Crinoids. 7. Fenestrate
bryozoans. 8. Gigantoproductid brachiopods.

Calcareous sand shoals. Sedimentation is represented

by echinoderm grainstonepackstone. It is an area
with constant level of moderate to high energy. That
energy is produced by waves that fragment bioclasts
producing a low variation in clasts size. However,
some organisms that live in turbulent, well-oxygen-
ated environments existed there (e.g. chaetetids).
Those that represent the last generations before the
final sedimentation are preserved entire or almost.
The assemblage include some rugose corals; some of
the recorded specimens are clearly reworked from
other areas (e.g. the Lithostrotion colony described
above) but most solitary corals show only short
remobilization (little abrasion) and lived in this envi-
ronment (see Table 1 and chapter on assemblages).
The presence of zones with micrite indicates that in
some areas the wave energy was not constant and
sedimentation could include some mud. These areas
would be the basal areas of the shoals, close to the
fair weather wave-base. The presence of abundant
crinoidal ossicles indicates that these invertebrates
flourished around shoals.
Unfortunately, rock exposures are very poor, but
irregular distribution of outcrops and microfacies
and presence of interbedded areas of packstone and
grainstone and even micropeloidal mudstones indi-
cates that shoals did not form a continuous sand bar,
but a shoal complex with erosive surfaces and closely
related areas where a microbial community could
flourish (Fig. 11AB).
Environment 2
Typical microfacies is micropeloidal mudstone
wackestone. Irregular stromatactoid cavities are also
common in this microfacies. It corresponds to quiet-
water zones below the fair weather wave-base or in
areas protected by shoals. The low level of turbulence
allows the development of microbial communities
that produce sedimentation of micropeloidal mud-
stone with laminar or homogeneous texture (Pratt
1995; Rodrguez-Martnez et al. 2003).
In the case of La Cornuda, the microbial community,
evidenced by presence of micropeloidal mudstone
and stromatactoid cavities, is combined with a flour-
ishing coral community composed of a diverse
assemblage (see Table 1 and chapter on assemblages).
52 A. Gmez-Herguedas & S. Rodrguez
Fig. 12. A. Community structure in microbialcoral mounds.
1. Guadiatia pseudocoloniale. 2. Unidentified axophyllid spp. 3.
Haplolasma lamelliferum. 4. Melanophyllum? spp. 5. Arachno-
lasma? sp. 6. Amygdalophyllum cornudensis. 7. Lithostrotion
maccoyanum. 8. Diphyphylum fasciculatum. 9. Diphyphyllum
gracile. 10. Unidentified lithostrotionid. 11. Syringoporoids. 12.
Falsocalcifolium punctatum. B. Community structure in oncoidal
ramp. All corals are reworked and occur in oncoid nucleii.
Algae (mainly Falsocalcifolium punctatum), cyano-
bacteria, tabulate corals, bryozoans and crinoids are
also common in this environment.
This environment could be located either behind
the calcareous sand shoals of environment 1 or in
small mounds in front of and/or below the shoals
(below the fair weather wave-base). The common
mudstone fragments and bioclasts in the shoals
coming from microbial mudstone areas indicate that
the latter should be windward from the former. Thus,
we illustrate this environment following the second
hypothesis (Figs 11A, 12A).
The small mounds should be below, but close to
the fair weather wave-base. Consequently, storms
affected them regularly, destroying them partly and
transporting some of the specimens to the shoals.
These storms also would transport clasts to the
mound producing areas richest in bioclasts
(wackestone areas).
These mounds have some features in common
with Visan mounds described in the Guadiato Area
LETHAIA 42 (2009)
(Czar et al. 2003b; Rodrguez-Martnez et al. 2003),
such as the textures, the fabrics and the presence of
stromatactoid cavities. However, there are conspicuous
differences, because rugose corals and algae are
absent or virtually absent in the Visan mounds,
occurring only on their tops. The main reason for
these differences may be either that some rugose
corals could acquire compatibility with microbial
communities during the Serpukhovian or that most
Visan mounds developed in deeper water where
most rugose corals could not subsist.
The high level of endemism in the coral assem-
blage from microbial mounds at La Cornuda may be
related to the isolation of Serpukhovian platforms,
caused by regression after the Upper Visan trans-
gression. In addition, the level of production of new
colonial taxa with low viability is high in microbial
mounds (Rodrguez & Somerville in press). The small
corallite size of most colonial corals is explained by
the main evolutionary lines of lithostrotionid corals
during the Visan (Poty 1993).
Environment 3
We regard it as a shallow ramp close to the calcareous
sand shoals and microbial mounds. Oncoidal pack-
stone has been sedimented in this environment. It is
not as well characterized as echinoderm grainstone
packstone and micropeloidal mudstonewackestone
because: (i) It is less represented in unit 1 from La
Cornuda; it occurs usually in marginal areas of the
rock and seems to be a transitional facies to the marls
of unit 2, that are usually covered. (ii) It seems to be
a zone of remobilization, where clasts derived from
other environments accumulated. All rugose corals
are transported and probably some of them lived in
this zone. Commonly, oncoids are small here, show-
ing few layers. It means that this environment is close
to the area where the corals lived (and other inverte-
brates in the nucleus of oncoids, such as brachiopods,
tabulate corals and gastropods). However, some
oncoids have many layers, suggesting a long time of
remobilization. The sediment around oncoids is an
echinoderm packstone; thus, the main component is
the same as in the grainstone. The presence of the
same main bioclastic components and the presence
of several genera or species in common with environ-
ments 1 and 2 indicate that this environment was in
close proximity to them (Figs 11A, 12B).
Conclusions
Palaeoecological and taphonomical study of rugose
corals combined with analysis of microfacies allows
LETHAIA 42 (2009)
precise identification of palaeoenvironments. As a
case study, we analysed the section of La Cornuda
(Serpukhovian, SW Spain) where outcrops are poor
and fragmentary.
We identified three environments in one single
lithological unit:
1 Calcareous sand shoals developed close to the fair
weather wave-base. The main microfacies from
this environment is echinoderm grainstone
packstone. Rugose corals are common but some
of them are transported from other environment.
Despite this, one massive colony was located
there, but only solitary dissepimented rugosans
lived there.
2 Microbial-coral communities developed in small
mounds close to the shoals (below the fair weather
wave-base) where rugose corals were common.
The microfacies include micropeloidal mudstone
containing diverse corals in life position, and
wackestone containing rare transported corals.
Solitary (dissepimented and undissepimented) and
colonial (incipient, fasciculate and massive) rugosans
are conspicuous components of the mounds.
3 Shallow-water platform facies associated with
environments 1 and 2. The microfacies is oncoidal
packstone. Oncoids are composed of Girvanella
and Sparaphralysia that may form thin or thick
coatings around rugose and tabulate corals,
brachiopods and gastropods. All corals in this
environment are transported.
Acknowledgements. Field and laboratory work have been sup-
ported by the Spanish Science Ministry Project BTE2003-02065
and CAM grant for the Research Group of the Complutense
University 910231. The authors are grateful to Dr Pedro Czar
for the identification of algae, to Carlos Alonso for development
and digitalization of photographs and to Ian Somerville and
an anonymous referee for helpful reviews that improved this
paper.
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