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A new Plattenkalk Konservat Lagerstätte in the

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A new Plattenkalk Konservat Lagerstätte in the Upper Cretaceous of Gara Sbaa,

south-eastern Morocco
David M. Martill a, *, Nizar Ibrahim b, Paulo M. Brito c, Lahssen Baider d, Samir Zhouri d, Robert Loveridge a,
Darren Naish a, Richard Hing a
a
Palaeobiology Research Group, School of Earth and Environmental Sciences, University of Portsmouth, Burnaby Road, Portsmouth, PO1 3QL, UK
b
School of Medicine and Medical Sciences, University College Dublin, Belfield, Dublin, Ireland
c
Universidade do Estado do Rio de Janeiro, Instituto de Biologia, Departamento de Zoologia, Rio de Janeiro, Brazil
d
Laboratoire de Géosciences, Université Hassan II, Faculté des Sciences Ain Chock, Casablanca, Morocco

a r t i c l e i n f o a b s t r a c t

Article history: Laminated, dolomitised marine limestones within a transgressive fluvio-lagoonal-carbonate platform
Received 7 August 2009 sequence at Gara Sbaa in the Kem Kem region of south eastern Morocco yield well preserved marine
Accepted in revised form 24 January 2011 fishes and crustaceans. A rarer terrestrial component includes delicate fern fronds, insects and
Available online 2 February 2011
a lizard. Sedimentological structures and stratigraphic context indicate initial shallow waters with in
a carbonate lagoon with benthic microbial mats followed by deepening. High precision dating has yet
Keywords:
to be accomplished, but a Late Cenomanian or Early Turonian age is indicated on the basis of faunal
Cretaceous
and sequence-stratigraphic considerations. The fish assemblage has affinities with mid-Cretaceous
Konservat Lagerstätte
Vertebrata
ichthyofaunas in South America and Lebanon, including taxa in common at generic level.
Arthropoda Ó 2011 Elsevier Ltd. All rights reserved.
Morocco

1. Introduction while undertaking fieldwork on the fluvial Kem Kem beds during
November and December 2008. The project, with personnel from
The eastern Moroccan Kem Kem plateau, its escarpment and University College Dublin, Casablanca University and the Univer-
basin floor is something of a palaeontological paradise. Situated sity of Portsmouth centred on studying the vertebrate palae-
close to the border with Algeria, the deeply dissected plateau is ontology and palaeonvironments of the Kem Kem beds, the results
formed of highly fossiliferous early Late Cretaceous limestone of which will be reported elsewhere.
overlying fluvioclastic sequences of probable Cenomanian age
that yield amphibians (Rage and Dutheil 2008) turtles (Tong and 2. Locality and logistical notes
Buffetaut, 1996), crocodyliforms (Buffetaut, 1994; Broin, 2002;
Larsson and Sues, 2007), pterosaurs (Mader and Kellner, 1999; The new fossil Lagerstätte is exposed on the top of a small mesa
Wellnhofer and Buffetaut, 1999; Ibrahim et al., 2010), non-avian known as Gara Sbaa (sometimes spelled Gara Sba or Gara Es Sba,
dinosaurs (Buffetaut, 1989; Sereno et al., 1996, 1998; Russell and but pronounced gara sbah) with an elevation of w970 m located
Paesler, 2003), possible birds (Riff et al., 2004) and highly abun- adjacent to the Kem Kem plateau some 30 km south of Tafraoute in
dant fish remains (Cavin and Dutheil, 1999; Cavin and Brito, 2001; south-eastern Morocco (coordinates 30
300 28.4100 N; 4
500 33.9300
Forey and Cavin, 2007). This sequence rests unconformably on W) (Figs. 1 and 2). The locality is accessible only by 4  4 vehicle,
Devonian and older basement of the Tafilalt region, which is motorbike or camel train and there are no permanent tracks to the
equally fossiliferous and has become famous for its abundant locality, although the main track linking Taouz with Zagora skirts
orthocones and incredibly diverse and spectacular trilobite Gara Sbaa Hill some 5 km to the north-west. The terrain between
assemblages. The preliminary report presented here describes Gara Sbaa and the main track is typical hamada, dissected by
a newly recognised fossil Konservat Lagerstätte that was explored shallow oueds with drifting sand in some of the oueds approaching
Gara Sbaa from the south and west. There is no water locally and
the nearest supplies are at ‘Auberge Kem Kem’ at GPS coordinates
* Corresponding author. N:30
38.213; W: 04
48.628. A mobile telephone signal can be
E-mail address: [Link]@[Link] (D.M. Martill). obtained between two small rounded hills adjacent to the west

0195-6671/$ e see front matter Ó 2011 Elsevier Ltd. All rights reserved.
doi:10.1016/[Link].2011.01.005
434 D.M. Martill et al. / Cretaceous Research 32 (2011) 433e446

Fig. 1. Locality map showing the outcrop of the Kem Kem sequence and the Cenomanian/Turonian limestone escarpment in south-eastern Morocco. A telephone signal can be
received at locality X.

flank of Gara Sbaa (Figs. 1e3) and diesel can be bought at Tafraoute Attention should be paid to the presence of horned desert vipers
(GPS coordinates N:30
40.139; W:04
40.770). Gara Sbaa is (Cerastes cerastes) on the lower slopes.
completely surrounded by vertical escarpments and steep slopes
clad with rocky talus (Figs. 2B and 3A). There is no track to the 3. Geology
summit where outcrops of the Lagerstätte are quarried for fossils,
thus all equipment has to be carried up steep, dangerous slopes in The Gara Sbaa fossil Lagerstätte overlies a sequence of fluvial
temperatures often exceeding 40 degrees in the summer months. siliciclastics rich in vertebrate remains, including some of the

Fig. 2. A. Satellite images of the Kem Kem plateau and Gara Sbaa Hill. The arrow indicates the direction view seen in B. Oblique view of Hara Sbaa Hill looking towards SSE. Images
from Google Earth.
 D.M. Martill et al. / Cretaceous Research 32 (2011) 433e446 435

Fig. 3. General terrain and nature of outcrop of Cretaceous strata at Gara Sbaa. A, Gara Sbaa Hill from the base on the south side looking north east. B, the limestones and marls
between the Kem Kem sequence continental deposits and the Gara Sbaa laminites. C, the Gara Sbaa laminated carbonates with well preserved fossils. D, local fossil collectors collect
fossils for the fossil shops in Erfoud. E, a fossil quarry at the summit of Gara Sbaa Hill.
436 D.M. Martill et al. / Cretaceous Research 32 (2011) 433e446

largest predatory dinosaurs in the world (Sereno et al., 1996; Dal involved in so-called Alpine deformation that produced the High
Sasso et al., 2005). These dominantly red-bed strata, commonly Atlas ranges (Brede et al. 1992). The outcrop of the Kem Kem beds
called the Continental Intercalaire (e.g. Lefranc and Guiraud, 1990) and the overlying carbonate sequence is extensive and can be
and termed the Kem Kem sequence by Sereno et al. (1996) lie traced from the Tindouf Basin in the west to the Errachidia-Bou-
unconformably on Palaeozoic strata folded during the Variscan denip basins in the East. However, despite the considerable extent
orogeny (Zouhri et al., 2008) (Fig. 4). The Kem Kem sequence is of the carbonates, with a linear surface outcrop in excess of 300 km,
comformably overlain by extensive marine carbonates deposited the Gara Sbaa deposit appears to be of extremely limited extent
during the extensive Cenomanian/Turonian marine transgression occurring only on the summit of Gara Sbaa and on an adjacent
that can be traced widely across this region of Africa (e.g. Baidder, escarpment. There appears to be no obvious structural control for
2007; Busson et al., 1999; Ettachfini and Andreu, 2004). The this restricted facies distribution.
strata are mostly flat lying, except in more northerly parts of the There are no calcareous fossils in the Gara Sbaa Lagerstätte
outcrop where, to the north and west of Errachidia they have been and, consequently, it has proved difficult to establish the precise

Fig. 4. Simplified stratigraphic scheme for the Gara Sbaa locality with the level of the Gara Sbaa plattenkalk indicated.
 D.M. Martill et al. / Cretaceous Research 32 (2011) 433e446 437

Fig. 5. Preliminary stratigraphic log for the upper part of the sequence at Gara Sbaa. Here we refer to the laminites as the Gara Sbaa Member. It’s base has a sharp contact with the
underlying marls (see Fig. 7) and contrasts both lithologically and faunistically with underlying strata. Its upper limit is not seen at Gara Sbaa.
438 D.M. Martill et al. / Cretaceous Research 32 (2011) 433e446

age of the deposit. The underlying Kem Kem beds, in this region age is correct, then laminated carbonate deposition of the Gara
resting on folded Palaeozoic strata, has been dated as Albian or Sbaa Lagerstätte may have been influenced by the Late Cen-
Cenomanian on account of its vertebrate remains. Sereno et al. omanian ocean anoxic event (Schlanger et al., 1987; Macleod et al.,
(1996) suggest Cenomanian on account of the fish assemblage 2008).
but Taquet and Russell (1998) and Russell (1996) suggest
a possible Albian age, an uncertainty reflecting a ‘slop’ of around
4. Sedimentology
15 million years. However, over most of the outcrop the verte-
brate-bearing part of the Kem Kem beds is overlain be a series of
The fossil-bearing parts of the Gara Sbaa laminites are restricted
variegated mudstones, thin sandstones and evaporites for which
to a few horizons within the finely laminated dolomitised limestone
there is no age determination. These mudstones pass up into
with laminae of just 1e2 mm in thickness. In the middle to upper
marls and shelly limestones and, at Gara Sbaa, into the fossilif-
part of the unit the laminae are planar with few interruptions, but
erous laminated limestone described here (Fig. 5). The marls and
toward the base some bedding planes exhibit ripple marks with
shelly limestones yield abundant examples of Ilymatogyra cf.
wave lengths of w50 mm (Fig. 6B). Some bedding planes are
africana, a bivalve that usually indicates a mid to early Late Cen-
crowded with linear fractures interpreted here as sineresis cracks.
omanian age (Ettachfini and Andreu, 2004; Dhondt and Jaillard,
No trace fossils were seen on bedding surfaces and no bioturbation
2005). In the Goulmima region of southern Morocco laterally
was observed to cut the fine-scale lamination. Thin sections reveal
equivalent carbonates of the Akrabou Formation have been dated
the laminae to be composed of fine grained dolomite rhombs with
as Late Cenomanian to Turonian on account of their marine
little evidence for grading between laminae (Fig. 6A, C, D). Some
benthic and planktonic foraminiferan assemblages (Ettachfini and
prominent laminae are slightly darker than other laminae and have
Andreu, 2004). It is highly likely that the Gara Sbaa Lagerstätte is
a whispy grey appearance under light microscopy suggesting the
of Late Cenomanian or Early Turonian age but because there are
presence of clay minerals (Fig. 6C). Scanning electron microscopy
no strata overlying the laminites, a minimum age cannot be
reveals these darker layers to comprise a mixture of dolomite
determined at this locality. If the Cenomanian/Turonian boundary
rhombohedra with overgrowths of authigenic smectite clays.

Fig. 6. Sedimentary structures in the Gara Sbaa Lagerstätte. A, fine-scale lamination as seen in a cut surface. B, low amplitude ripple marked surface from base of laminite sequence.
C, Clay-rich lamina seen in thin section. D, whispy mix of smectitic clays and dolomite microspar.
 D.M. Martill et al. / Cretaceous Research 32 (2011) 433e446 439

Particularly conspicuous are nodules of chert, concentrated in

bands. The chert is early diagenetic, forming mammilate and
vermiform shapes (Fig. 7A,C), although the nodules do not appear to
have nucleated around fossils.

5. Fossil assemblage

5.1. Preservation and taphonomy

All of the fossils in the Gara Sbaa laminites are preserved in

a flattened condition. The vertebrates are preserved as complete
individuals, with bones articulated and preserved as brown to
amber coloured elements. In at least one example, bone acted as
a nucleus for the development of a bright blue mineral, most likely
the hydrated iron phosphate vivianite, but the specimen, a small
lizard (see Fig. 12) was not available for analysis. Orange-brown
iron staining is associated with some of the fish specimens, and
may reflect oxidised relics of pyrite formed in soft tissues. Simi-
larly, plant remains are preserved as iron oxide films (see below).
Invertebrates such as isopods are preserved as external moulds,
although in larger arthropods (w30e50 mm), such as decapods,
there is some cuticle preserved. An early diagenetic carbonate or
phosphate phase is present lining cuticular surfaces, giving the
fossils a white colouration that contrasts with the cream colour of
the dolomite (Fig. 13). A single gastropod was found as a composite
mould.

5.2. Fish

Fish are the most frequent fossils in the Gara Sbaa assemblage
and include a variety of actinopterygians, but no elasmobranchs or
sarcopterygians. The fish assemblage comprises a macrosemiid
(Fig. 8A), Agoultichthys chattertoni Murray and Wilson (2009), the
aspidorhynchid Belonostomus sp. (Fig. 10A), a new species of the
pycnodontid Pycnodus (Fig. 8C), a new species of the ichthyo-
dectiform Cladocyclus (Fig. 8B), two dercetids, Saurorhamphus and
Rhynchodercetis (Fig. 10B and C), at least two Paraclupeidae, Dip-
lomystus and an Ellimmichthys-like form and (Fig. 9A and B), a basal
ostaryophisian (Fig. 9C) and a new species of acanthomorph
(Fig. 11). Most frequent appear to be small, generically indetermi-
nate teleosts (Fig. 9C) and the paraclupeids Ellimmichthys and
Diplomystus. Juveniles of the pycnodontid Pycnodus are very similar
to Pycnodus laveirensis from the Turonian of Laveiras, Portugal and
to another species, as yet not described, from Mexico (Brito et al. in
prep.).
The Gara Sbaa dercetids are also relatively small specimens, and
no fish larger than 300 mm standard length were recovered, hint-
ing at some size selection mechanism.
Before the recent description of the macrosemiid Agoultichthys
chattertoni by Murray and Wilson (2009), the temporal and
geographic range of Macrosemiidae was from the Late Triassic to
the Albian, with representatives in Europe and in Mexico. Murray
and Wilson (2009) explained the presence of this taxon in
Morocco by a transgression event in the Late Cenomanian allowing
the Tethyan fauna to expand throughout northern Africa to the
Middle East where it also occurs in Lebanon. Belonostomus has
a world-wide distribution and ranges from the Late Jurassic to the Fig. 7. Sedimentary structures in the Gara Sbaa Lagerstätte. A. Sharp contact of the
Late Cretaceous. During the Cenomanian-Turonian, Belonostomus Gara Sbaa laminites with the underlying marls. Notice the chert horizon developed at
the base. B, sineresis cracks from base of laminate sequence. C, mammilated chert,
appeared to be distributed along shallow coastal waters of the
a common occurrence in the laminites.
Cretaceous seaway of North America (Mexico) extending east-
wards to shallow parts of the Tethys region including Morocco,
Lebanon, Israel, Italy and Slovenia. The Gara Sbaa Belonostomus
appears to have affinities with a group of Belonostomus species
comprising B. crassirostris from the Cenomanian of Pietraroia, Italy
and the Jebel Tselfat, northern Morroco (Arambourg, 1954),
440 D.M. Martill et al. / Cretaceous Research 32 (2011) 433e446

Fig. 8. Fossil fish from the Gara Sbaa Lagerstätte. A, Macrocemiid provisionally idientified as Agoultichthys chattertoni Murray and Wilson (2009). B, Cladocyclus sp. nov. C, Pyc-
nodontid provisionally identified as a new species of Pycnodus. Scale bar in a,b 20 mm, scale bar in c 10 mm.
 D.M. Martill et al. / Cretaceous Research 32 (2011) 433e446 441

B. lesinaensis from Slovenia (see Cavin and Kolar-Jurkovsek, 2000), beds. Similarly, the Kem Kem ichthyodectid Cladocyclus pankowskii
and a new species from Lebanon (Belonostomus sp. 2, Forey et al., recently described by Forey and Cavin (2007) differs significantly
2003; Brito in prep.). from Cladocyclus gardneri from the Brazilian Santana Formation in
The Gara Sbaa fish assemblage contrasts markedly with that of the form of its lower jaw, and should probably be referred to a new
the underlying Kem Kem beds which includes selachians (Dutheil, genus. The Gara Sbaa ichthyodectid cannot be assigned to C. pan-
1996), dipnoans (Martin, 1984), coelacanths (Cavin and Forey, kowskii, but does appear to belong in Cladocyclus being close to the
2004), cladistians (Dutheil, 1999a) as well as lepisosteids (Cavin Brazilian C. gardneri, although considerably smaller (Brito et al. in
and Brito, 2001), ichthyodectids (Forey and Cavin, 2007) and prep.). The presence of Cladocyclus in Morocco increases the
other actinopterygians distinct from those of Gara Sbaa (Dutheil, geographical distribution of this taxon, which occurs commonly in
1999b). It is notable that the fish taxa descreibed from the Kem the Aptian/Albian of north east Brazilian basins (e.g., Araripe,
Kem fluvial sandstones are dominantly found as isolated bones Parnaíba, Sergipe-Alagoas basins). Cladocyclus seems to be
and teeth of a few large taxa (e.g. Ceratodus, Onchopristis numidus, a typical southern Tethys species and the distribution of this taxon
and M. cf. gigas), whereas the laminated ?lacustrine silts of the in South America during the Lower Cretaceous is easily explained
Kem Kem sequence tend to yield smaller, more complete examples by the presence of widespread epicontinental seas extending
of different taxa (e.g. Serenoichthys kemkemensis Dutheil 1999), southward from the shallow waters of the Caribbean Tethys onto
differences that probably reflect both environmental and tapho- the South American continent (Brito, 1997; Maisey, 2000; Brito
nomic factors. The small acanthomorph from Gara Sbaa (Fig. 11) et al. 2008).
differs significantly from Spinocaudichthys described by Filleul and Diplomystus is known from the Cretaceous of Lebanon, and
Dutheil (2001) from the upper unit of the underlying Kem Kem tertiary of North America and China. The “Ellimmichthys-like” par-
aclupeid is very similar to Ellimmichthys longicostatus from Brazil,
and Ellimmichthys goodi from West Africa.
The dercetids have a circum-Tethyan distribution. Sauro-
rhamphus has two nominal species, both from the Lower Cen-
omanian: Saurorhamphus freyeri from Trieste-Komen (Goody,
1969) and S. judeaensis from Ein-Yabrud, Jerusalem, Israel (Chalifa,
1985). The genus Rhynchodercetis has six nominal species:
R. yovanovitchi from the Lower Cenomanian of Jebel Tselfat,
Morocco (Arambourg, 1954; Goody, 1969), (with R. cf. yovanovitchi
from the Middle Cenomanian of Namoura; Forey et al., 2003); R.
gortanii from the Lower Cenomanian of Trieste (Goody, 1969), R.
hakelensis from the Middle Cenomanian of Hakel, Lebanon (Pictet
and Humbert, 1866; Goody, 1969; Forey et al., 2003), R. serpentinus
from the Cenomanian Hakel and Hajula (Forey et al., 2003); R.
gracilis from the mid Cenomanian of Namoura (Chalifa, 1989; Forey
et al., 2003) and R. regio from the Turonian Agua Nueva Formation
of Mexico (Blanco and Alvarado-Ortega, 2006).
The Gara Sbaa ostaryophisian is a Lusitanichthys-like fish. Lusi-
tanichthys is known from the Cenomanian of Morocco (Cavin, 1999)
and from the Turonian of Laveiras, Portugal (Gayet, 1985).

5.3. Tetrapods

No tetrapods were collected during our field study, but a small

reptile (Fig. 12) was obtained by one of the collectors we
encountered at the site. This specimen was sold to a fossil dealer
and its present whereabouts is not known. The specimen appears
to represent an unidentified lizard (Evans pers. com. 2008), but
the preservation does not allow the recognition of diagnostic
features.

5.4. Invertebrates

Although invertebrates, mainly pectenacean and ostreacean

bivalves, are abundant in the non-laminated carbonate strata
immediately underlying the laminites, they appear to be absent
from the laminites themselves. We encountered only arthropods,
with a small isopod occurring most frequently (Fig. 13B). All of the
arthropods were articulated, complete or near complete
exoskeletons, and probably represented dead individuals, rather
than exuviae. A single example of a hymenopteran insect
(Fig. 13A) was discovered that is recognisable as an aculeate wasp
Fig. 9. Fossil fish from the Gara Sbaa Lagerstäätte. A, clupeid Diplomystus sp. B,
and preliminarily identified as a chrysidoid (cuckoo wasps) with
Ellimmichthys sp. C, unidentified ostaryophysian close to Lusitanichthys. Scale similarities to Bethylidae and amesigine chrysidids (Heads pers.
bars ¼ 10 mm. com. 2009).
442 D.M. Martill et al. / Cretaceous Research 32 (2011) 433e446

Fig. 10. Fossil fish from the Gara Sbaa Lagerstätte. A, Belonostomus sp. B, Saurorhamphus sp. C, Rhynchodercetis sp. Scale bar in A, 50 mm, in b,c ¼ 10 mm.
 D.M. Martill et al. / Cretaceous Research 32 (2011) 433e446 443

6. Discussion

The Gara Sbaa fossil locality represents a shallow marine plat-

tenkalk Konservat Lagerstätte deposited during the latest Cen-
omanian marine transgression, perhaps under the influence of
widespread oceanic anoxia. Its fossil assemblage is dominated by
marine fishes with rarer input from nearby terrestrial sources.
Sedimentological evidence indicates that the deposit was
formed as the transgression deepened, and that current swept
bottom waters gave way to quiet, probably anoxic conditions,
resulting in fine-scale sedimentary lamination and a lack of bio-
turbation. Diagenetic processes resulted in early production of
pyrite, later oxidised to goethite, with at least some preservation of
organic material such as insect and other arthropod cuticles. Some
late stage processes resulted in at least partial loss of cuticle,
except where early diagenetic mineral phases coated its surface.
The pale cream colour of the dolomitic laminites is probably
a consequence of deep weathering of the Kem Kem carbonate
plateau during the Cenozoic. The limited areal extent of the Gara
Sbaa Lagerstätte may reflect a gentle gradient into a deeper
depression within the Kem Kem carbonate platform, but a more
detailed analysis of the lateral variation and extent of this
remarkably rich deposit is required.
The fish assemblage is diverse, but typified by small species or
juveniles and, so far, has only yielded actinopterygians. There are
a number of important fish-bearing horizons within the Cretaceous
of south east Morocco, reviewed briefly by Cavin et al. (2001) and in
detail by Cavin and Dutheil (1999).
The fossil assemblage at Gara Sbaa differs greatly in both
composition and preservation from the underlying Kem Kem
sequence sandstone found widely in the region, but bears some
preservational similarities with the fossils from the Douira locality
Fig. 11. Part and counterpart of new acanthomorph from the Gara Sbaa Lagerstätte.
near Erfoud (note there are localities yielding fossil fish at Daoura
Scale bar 20 mm.
and Douira, and these should not be confused). At Douira articu-
lated, flattened freshwater fishes occur in laminated silts at the
A brachyuran crab (Fig. 13C) was also collected. It lacks the rear boundary between the lower, coarser sandstone unit of the Kem
most and the anterior two pairs of appendages, and the abdomen is Kem and the upper, more argillaceous unit. At Douara, near to the
everted with a convex lateral margins suggesting a female. The confluence of the Rheris and Ziz rivers, articulated, flattened fishes
rounded carapace is suggestive of a portunid crab. occur at a stratigraphic level that may be equivalent to that of Gara
Sbaa. Unfortunately, no field analysis of this locality has been
described (Cavin, 1999; Cavin et al., 2001).
5.5. Flora The Gara Sbaa fossil assemblage also differs from the Early
Turonian locality of Goulmima in the northern part of the Creta-
A few fragments of fern fronds (Fig. 14) were encountered with ceous basin where a marine fish assemblage occurs in early
additional examples seen in fossil dealers’ storehouses in Erfoud on diagenetic carbonate concretions similar to those of the Santana
a similar matrix. All are preserved as orange/brown goethite films Formation of Brazil (Cavin, 1997). Thus, although the Gara Sbaa
similar to those encountered in plants from the Crato Formation of locality shows some similarities with other fish localities within the
Brazil (Mohr et al., 2008). basin, its unique stratigraphic position and its distinctive, well

Fig. 12. Small, indeterminate lizard from the Gara Sbaa Lagerstätte. This is the only known tetrapod from this horizon and its present whereabouts is unknown. Some of its bones
were incrusted with a mineral resembling vivianite. Scale not known.
444 D.M. Martill et al. / Cretaceous Research 32 (2011) 433e446

Fig. 13. Invertebrates and flora from the Gara Sbaa Lagerstätte. A, Insecta, Hymenoptera gen. et sp. nov., probably close to aculeate (cuckoo) wasps; B, Crustacea, Isopoda;
C, Crustacea, Decapoda, brachyurid crab cf Marocarcinus pasinii Guinot et al., 2008. Scale bars: a,b ¼ 5 mm, c ¼ 10 mm.
 D.M. Martill et al. / Cretaceous Research 32 (2011) 433e446 445

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Cavin, L., Dutheil, D.B., 1999. A new Cenomanian ichthyofauna from southeastern
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Cavin, L., Forey, P.L., 2004. New mawsoniid coelacanth (Sacropterygii: Actinistia)
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Cavin, L., Kolar-Jurkovsek, T., 2000. Stratigraphic succession of the upper cretaceous
Fig. 14. Portion of fern frond from the Gara Sbaa Lagerstätte. The mode of preservation fish assemblages of Krasa (Slovenia). Geologija 43 (2), 165e195.
is remarkably similar to that of flora from the Aptian Crato Formation of Brazil. Cavin, L., Tong, H., Boudad, L., Meister, C., Piuz, A., Tabouelle, J., Aarab, M., Amiot, R.,
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preserved fossil assemblage clearly indicate that further analysis is Chalifa, Y., 1985. Saurorhamphus judeaensis (Salmoniformes: Enchodontidae), a new
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Chalifa, Y., 1989. Two new species of longirostrine fishes from the early Cenomanian
(Late Cretaceous) of Ein-Yabrud, Israel, with comments on the phylogeny of the
For further reading Dercetidae. Journal of Vertebrate Paleontology 9, 314e328.
Dal Sasso, C., Maganuco, S., Buffetaut, E., Mendez, E.M.A., 2005. New information on
After submission of this paper Cavin et al. (2010) published the skull of the enigmatic theropod Spinosaurus with remarks on its size and
affinities. Journal of Vertebrate Paleontology 25, 888e896.
a paper in which the Gara Sbaa locality is discussed in the context of Dhondt, A.V., Jaillard, E., 2005. Cretaceous bivalves from Ecuador and northern Peru.
other Moroccan Early Cretaceous vertebrate assemblages, referring Journal of South American Earth Sciences 19, 325e342.
to the Gara Sbaa assemblage as the Agoulti assemblage. Dutheil, D.B., 1996. Nouvelles découvertes d’élasmobranches dans le Crétacé
supérieur du Sud marocain, implications stratigraphiques. In: Vénec Peyré, M.-
T., Tassy, P. (Eds.), Deuxième Congrès national de Paléontologie. Paléontologie,
Histoire récente et Perspective a l’aube de l’an 2000, p. 28. Paris.
Acknowledgements Dutheil, D.B., 1999a. The first articulated fossil cladistian: Serenoichthys kemke-
mensis, gen. et sp. nov., from the Cretaceous of Morocco. Journal of Vertebrate
We are grateful to many people who supported our expedition Paleontology 19, 243e246.
Dutheil, D.B., 1999b. An overview of the freshwaterfish fauna from the Kem Kem
to Morocco. Financial support from the University of Portsmouth beds (Late Cretaceous: Cenomanian) of south eastern Morocco. In: Arratia, G.,
for DMM and RFL is gratefully acknowledged as is an AD ASTRA Schultze, H.-P. (Eds.), Mesozoic Fishes 2-Systematics and Fossil Record. Dr F.
scholarship awarded to NI. Thanks to Dr Sam Heads for determining Pfeil, München, pp. 553e563.
Ettachfini, El M., Andreu, B., 2004. Le Cénomanien et le Turonien de la plat-forme
the insect, Geoff Long for thin sections, Susan Evans for examining
Préafricaine du Maroc. Cretaceous Research 25, 277e302.
pictures of the lizard and the good folk at Google Earth. Two Filleul, A., Dutheil, D.B., 2001. Spinocaudichthys oumtkoutensis, a freshwater acan-
anonymous referees provided important commentary on the thomorph from the Cenomanian of Morocco. Journal of Vertebrate Paleontology
21, 774e780.
submitted version of this paper.
Forey, P.L., Lu, Y., Paterson, C., Davies, C.E., 2003. Fossil fishes from the Cenomanian
(Upper Cretaceous) of Namoura, Lebanon. Jouranl of Systematic Palaeontology
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