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Late Cretaceous continental and marine vertebrate assemblages of the Lano Quarry (Basque-Cantabrian Region, Iberian Peninsula): an update.

1. Introduction

The occurrence of fossil vertebrates in the Late Cretaceous of the Lano quarry is known since 1984, when the palaeontologist Xabier Orue-Etxebarria discovered bone remains in the quarry (Astibia et al., 1999a). Surface prospections along 1985 and 1986 led to the discovery of remains of turtles, crocodylians, and dinosaurs (Astibia et al., 1986, 1987; Sanz, 1986). Field research in Lano has been carried out by members of the Universidad del Pais Vasco/Euskal Herriko Unibertsitatea (UPV/EHU) in collaboration with researchers of the Universidad Autonoma de Madrid (UAM), the Centre National de la Recherche Scientifique (CNRS), the Universite Pierre et Marie Curie (Paris VI), the Universite de Montpellier II and the Museo de Ciencias Naturales de Alava/ Arabako Natur Zientzien Museoa (MCNA, Vitoria-Gasteiz). Excavations made from 1987 to 1997 yielded a very large collection of vertebrate bones and teeth, including thousands of microfossils. Many of them come from three fossiliferous levels of continental origin (L1A, L1B and L2 from the S1U3 unit of Astibia et al., 1987). Other specimens, mostly isolated teeth of marine vertebrates, have been found in two levels of an overlying unit (S2U1 of Astibia et al., 1987). Astibia et al. (1990) provided a preliminary list of the Lano vertebrates, including actinopterygians, lissamphibians, squamates, turtles, crocodyliforms, dinosaurs, pterosaurs, and mammals from the continental deposits, and selachians from the marine beds. Subsequently, a number of new vertebrate taxa have been erected on the basis of the Lano fossils, including mammals (Gheerbrant and Astibia, 1994), snakes (Rage, 1996), turtles (Lapparent de Broin and Murelaga, 1996) and crocodyliforms (Buscalioni et al., 1997). In 1999, the vertebrate associations of the Lano quarry were the subject of a monographic volume (Astibia et al., 1999b). In this work, information about the local and regional geology of the quarry and adjacent areas was given, as well as data related to the palynomorphs and taphonomy (Astibia et al., 1999b and references therein). The volume also included the systematic description of the vertebrates, including a new sauropod dinosaur from Lano (Sanz et al., 1999) and four new selachian species from Albaina (Cappetta and Corral, 1999). Other papers dealt with the petrological and geochemical features of the Lano vertebrate fossils (Elorza et al., 1999; Lecuyer et al., 2003 and Corrigendum), as well as the taphonomic features of the terrestrial and freshwater vertebrate assemblages (Pereda Suberbiola et al., 2000). Recent papers on Lano deal with the description of a new species of the mammal Lainodon (Gheerbrant and Astibia, 2012), the revision of the titanosaurian sauropod Lirainosaurus (Diez Diaz et al., 2011, 2012, 2013a, 2013b; Diez Diaz, 2013) and the dortokid turtle Dortoka (Perez-Garcia et al., 2012a), and the description of the theropod teeth assemblage (Torices, 2007; Torices et al., in press), among others. In the sublittoral beds of Albaina, the first dinosaur and turtle remains have recently been described (Pereda-Suberbiola et al., in press).

The aim of this paper is to give a synthetic work of the palaeontological studies made in the Lano quarry also updating the faunal list of the terrestrial and freshwater vertebrates from Lano and the marine vertebrates from Albaina.

Institutional abbreviation: MCNA, Museo de Ciencias Naturales de Alava/Arabako Natur Zientzien Museoa, Vitoria-Gasteiz (Spain).

2. Geographical and geological context

The disused quarry of Lano is located between the small villages of Lano and Albaina in the Condado de Trevino (an Alava's enclave administered by the province of Burgos), about 30 km south the city of Vitoria-Gasteiz to the north of the Iberian Peninsula (Fig. 1).

The quarry that was largely exploited for both foundry and glass sand by Eusebio Echave S.A. (ECHASA) comprises two working faces separated by a stream (the Arroyo de Granado). In the palaeontological literature, the western working front has been referred as the Albaina site while the eastern one was named Lano site (Fig. 2). The continental fossil outcrops (Lano site) are nowadays rather overgrown and therefore collection of new specimens is more difficult with the passage of time. On the contrary, the occasional collapse of a quarry face and rockfalls in Albaina site still produce new finds, but collection must be cautiously done. As a result of the cut, a section from Late Campanian to Danian has been exposed, revealing one of the best and more diverse vertebrate fossil sites in Western Europe.

Geologically, the area lies within the southern limb of the South-Cantabrian Synclinorium, a large structure mainly composed of Upper Cretaceous and Paleogene deposits on the southeastern part of the Basque-Cantabrian Region (Baceta et al., 1999). Locally, the Lano quarry is located on the southern limb of the Miranda-Trevino syncline and includes a continental to shallow marine succession.

The Late Campanian marks a period of overall regression in the southern part of the Basque-Cantabrian Region (i.e., Navarro-Cantabrian Trough), causing land uplift, and development of fossil-rich deltaic environments and episodes of non-deposition in some marginal areas. As a result, both terrestrial and freshwater vertebrate fossils occur within an alluvial system composed mainly of fluvial sands and silts (Gomez-Alday, 1999; Pereda Suberbiola et al., 2000) in two successive beds named as L1A and L1B. Another silty fossiliferous bed named L2 occurs at the eastern end of the Lano quarry, and is apparently older in age by its correlated stratigraphic position. The sedimentary structures observed within this unit are consistent with channel areas within an exten sive braided river system. The main fossil accumulations of Lano are associated with diagenetic ferruginous crusts, and the bones are often coated by a millimetre-sized layer of sandstone with iron-oxide matrix (Elorza et al., 1999). Based on stratigraphic correlations, these fossiliferous beds are regarded as Late Campanian to Early Maastrichtian in age (Baceta et al., 1999; lateral equivalent to the upper part of Depositional cycle DC-11 sensu Floquet, 1998; see Berreteaga, 2008) (Fig. 3).

The regressive depositional trend, exceptionally observed in the Lano-Albaina area during the Campanian and Early Maastrichtian, was later followed by transgressive sedimentation with marine strata in both limbs of the MirandaTrevino syncline. The upper part of the studied series, which reflects this episode, is chiefly consisting of over ten-metre thick marine rocks (a mixed carbonate-siliciclastic succession) cropping out in the upper levels of the quarry. Friable marine sandstones and mudstones are connected to yellow microvertebrate-rich calcarenites containing teeth and bone remains of selachians, actinopterygians, and mosasaurids by means of a lag (with occasionally vertebrate remains) that marks the cycle boundary CB-13 (sensu Floquet, 1998). This feature is clearly shown in the western part of quarry (Albaina site) by means of a major intra-Maastrichtian unconformity (IMU boundary) (Baceta, 1996; Baceta et al., 1999). Invertebrate macrofossils in these calcarenites are scarce, in part because of its diagenetic history which has turned them to decalcified limestones with moldic porosity, but nevertheless layers of accumulated orbitoidids and mollusc steinkerns occur. The lack of diagnostic fossils makes difficult an ac curate stratigraphic assignment, but according to the selachian content a Late, but not latest, Maastrichtian age is given (Cappetta and Corral, 1999). This unnamed relatively soft formation is considered to be equivalent to the DC-13 depositional cycle that was represented by the Torme Formation in the Villarcayo area of Burgos province (North-Castilian Platform; see Floquet, 1991; Berretega, 2008). The overlying Fresnedo Formation consists of dolomitised carbonates, easily recognized in the area as ruiniform reliefs. This lithologic succession represents a new depositional sequence DC14, bounded by cycle boundary CB-14 (sensu Floquet, 1998).

3. Material and methods

Fossil remains from the L1A, L1B and L2 fossiliferous levels of the Lano quarry were collected by detailed surface digging. Eight excavation campaigns were made at Lano from 1987 to 1997. During the 1995 field season, the L1A level was mapped following a metre-square grid system. Both the L1A and L1B levels were mapped during the excavations of 1997. In addition, microfossils were collected by screen-washing of the sediments from the L1A level. About 8 tons of sediments were washed. Three meshes were used during screen-washing, with wire openings of 2, 0.7 and 0.5 mm, respectively.

Taphonomic features of the fluvial vertebrate assemblage of L1A were analyzed in order to interpret the taphonomic history of the fossil association. This analysis involved the study of a large number of variables within a well-defined sedimentological context (see Astibia et al., 1999c; Pereda Suberbiola et al., 2000). The microfossils were studied separately as they could have had a different taphonomic history.

Marine vertebrate fossils from Albaina come from the higher part of quarry face, which makes difficult direct collecting. Large isolated specimens (mostly teeth) occurred most commonly as scattered individual specimens on rock surfaces, and were directly hand-collected or by breaking off calcarenite rocks and disagregating them with a dilute solution of acetic acid. Vertebrate microfossils were obtained by screen-washing and picked up from the residue (Cappetta and Corral, 1999). Specimens were further preparated and curated at the MCNA laboratory.

The fossil vertebrate collection from the Lano quarry consists of thousands of bones and teeth, including abundant microfossils. The specimens are provisionally housed in the MCNA of Vitoria-Gasteiz (see Astibia et al., 1999b and articles therein for a detailed list of the material).

4. Taphonomy, geochemical features of the vertebrate fossils, and palaeoecology

The vertebrate fossil association of Lano, especially the L1A sample, is a heterogeneous assemblage of elements ranging from isolated bones and teeth to partially articulated skeletons (Astibia et al., 1999c; Pereda Suberbiola et al., 2000). The estimated body size ranges from less than 100 g for lissamphibians and mammals to about 3-4 tons for the largest dinosaur. The modal category of body size and weight is smaller than 1 m and 5 kg, respectively. Adult individuals are much more frequent than juveniles. Several states of skeletal articulation are represented, including articulated or associated specimens in close proximity, but most of the bones are disarticulated and dispersed. Dermal bones and isolated teeth are the most common elements in the assemblage. The spatial density in the L1A level is relatively high (about 80 elements per square metre). The bones, especially the long ones, are somewhat preferentially oriented (parallel to the fluvial current), and with a variable dip. The bone distribution is non-uniform, with a relatively large coefficient of variation in bones per square metre. A large percentage of elements are broken, and two-thirds of bones are splintered. The weathering and abrasion ranges are variable according to the taxon. There is no evidence of predatory activity or chemical alteration; only fungi marks have been observed on the surface of turtle plates (Astibia et al., 1999c; Pereda Suberbiola et al., 2000).

When compared with hypothetical taphonomic examples of vertebrate accumulations (see Behrensmeyer, 1991), the features of the Lano assemblage are not compatible with flood-related monospecific mass accumulations. It looks like a fluvial attritional model of sorted remains of both small and large vertebrates. The presence of at least two modes of fossil accumulations in L1A is worthy of consideration. On one hand, the dispersed and unassociated elements were probably accumulated by the fluvial system over a large interval of time (attritional accumulation). On the other hand, the articulated and associated bones were added to the bone assemblage, probably as a result of carcasses floating into the site (Pereda Suberbiola et al., 2000). This implies that both parautochthonous and allochthonous elements (sensu FernAndez Lopez, 2000) are probably mixed together in the fossil association of Lano; palaeoecologically, there are demic elements, consisting of aquatic (bony fish, amphibians) or semi-aquatic elements (crocodyliforms, chelonians) and also ademic, terrestrial ones (solemydid turtles, dinosaurs, mammals). The dinosaur bones are commonly fractured and show a greater degree of abrasion than those of freshwater vertebrates, indicating that they may be allochthonous (Pereda Suberbiola et al., 2000). Remains of actinopterygians, amphibians, pleurodiran turtles, and eusuchian crocodyliforms are interpreted as being para-autochthonous (i.e., fluvial specific elements).

The fossil association of Lano lies between the taphonomic modes for attritional vertebrate assemblages in fluvial channels proposed by Behrensmeyer (1988). Both lithologic and taphonomic features indicate that it tends more towards the channel-fill mode than towards the channel-lag mode (Astibia et al., 1999c; Pereda Suberbiola et al., 2000). Moreover, sedimentological features observed at Lano suggest a gradual abandonment of the channel, with occasional periods of reactivation (Gomez-Alday, 1999).

Petrologically, the vertebrate fossil remains from Lano are composed of well-crystallized francolite (carbonate fluorapatite). The replacement of biogenic hydroxyapatite by francolite is related to diagenetic changes (Elorza et al., 1999). The crystallinity index of the Lano fossil bones (CI = 0.2-0.3) is lower than that of modern bones. Compared to fresh bones, the francolite has higher concentrations of trace elements and rare earth elements (EREE 500 to 900 ppm). The geochemical composition of the Lano vertebrate fossils shows a homogeneous REE trend (Elorza et al., 1999; Lecuyer et al., 2003; Berreteaga, 2008). This suggests that the diagenetic processes were homogeneous. There is no evidence of mixing with previously diagenetically altered, reworked bones.

The diagenetic processes that lead to the Lano bone features have been interpreted as follows: 1) relatively rapid burial; 2) development of matrix and cement; 3) lithostatic compaction and fracturing of the bones (post-fossilization bone modification); 4) cementation of cavities and skeletal fractures (partial or total filling); 5) epidiagenesis and final fracturing (Pereda-Suberbiola et al., 1992, 2000; Elorza et al., 1999).

The main bone accumulations of Lano are associated with ferruginous surfaces. Moreover, the vertebrate remains are usually covered with nodule-like iron oxides, which are mainly composed of goethite and small detrital quartz grains with a minor percentage of clays (illite-kaolinite). The nature of the iron oxides that cover the bones is similar to that of the ferruginous structures. The development of the crusts and globule irons could have resulted from a hydromorphic process because of seasonal variations of the phreatic water (Astibia et al., 1999c; Elorza et al., 1999). This feature and the relative percentage of illite and kaolinite suggest a climate with dry and wet seasonal periods.

The stable isotope compositions of the Lano vertebrate fossils provide indirect evidence of a warm climate (subtropical to tropical) during the Campanian-Maastrichtian transition, in agreement with faunal associations (see Lecuyer et al., 2003 and Corrigendum; Berreteaga, 2008).

The small vertebrates with aquatic habits are dominant in the Lano association (Pereda Suberbiola et al., 2000). The presence of pelomedusoid turtles and crocodyliforms is consistent with an intertropical, warm climate. This interpretation agrees with the palynological assemblage found in organic matter rich beds of the alluvial system (see Fig. 3): the undergrowth was composed of ferns, gymnosperm (mainly pines and cypress) and angiosperm plants, which are indicative of a humid, temperate to subtropical climate (NunezBetelu, 1999).

With regard to the marine assemblage of Albaina, most of the fossils are isolated teeth that occur as bioclasts in a matrix. The total number exceeds a thousand of vertebrate elements. The taphonomic factors involved in the genesis of the Albaina outcrops are not established yet; then, no attempt of palaeoecological interpretation has been done (Poyato-Ariza et al., 1999; Cappetta and Corral, 1999). The fact that in the same layer selachian teeth with a pristine conservation are found together with others showing signs of abrasion suggests that the latter remained in a nearshore energetic environment for a long time before the burial. In a few cases, teeth are so highly eroded that it is difficult to identify them. Some teeth are broken off with matrix-filled cracks because of compaction by mechanical processes. Finally, the apex of some well-preserved teeth may be splintered with sharp edges, while it is smooth-rounded in other cases; these features are probably due to damage during feeding (Cappetta and Corral, 1999).

5. The vertebrate fossil assemblage from the continental beds of Lano

At least 37 continental vertebrate taxa are known in the Lano 1 and 2 sites, including actinopterygians, lissamphibians, lepidosaurs, turtles, crocodyliforms, dinosaurs, pterosaurs, and mammals (see Table 1). More than half of the taxa present in the L1A association have been recognized from microfossils. Seven genera and ten species have been erected to date on the basis of the Lano material (Table 2).

5.1. Actinopterygians

Ray-finned fish remains from Lano include mainly ganoid scales, vertebrae and some bones and teeth. The occurrence of opisthocoelous vertebral centra indicates the presence of Lepisosteiformes, or gars, to which the ganoid scales are associated and a supracleithrum with a patch of ganoid on its external side. Cavin (1999) referred the lepisosteid remains to the genus Atractosteus on the basis of the study of Wiley (1976). A recent review of fossil and Recent Lepisosteiformes by Grande (2010) led to reevaluate osteological diagnostic characters of the group. Consequently, the Lano material is here referred to an indeterminate Lepisosteidae. Gar remains are not uncommon in the Late Cretaceous deposits of Europe, but generally not complete enough for being identified at the generic and specific level (see Cavin, 1999 and references).

Teleosteans are also present in the Lano assemblage with isolated, thin and subcircular teeth attributed to an indeterminate Phyllodontidae by Cavin (1999), but referable with caution to Phyllodus by comparison with the material from Albaina (Poyato-Ariza et al., 1999). A fragment of tooth plate was referred to ?Palaeolabrus, but this identification should be considered with extreme caution. The occurrence of Phyllodontidae and Palaeolabridae in the Late Cretaceous of Europe was first recorded in Lano (Cavin, 1999), then Phyllodontidae have been recovered in two other sites from the Maastrichtian of southern France and Spain (Laurent et al., 1999; Berreteaga et al., 2011).

5.2. Lissamphibians

Lissamphibian remains are relatively abundant in Lano and consist of about 200 isolated bones. The Lano assemblage is one of the richest and most diverse among those from the Late Cretaceous of Europe (Duffaud and Rage, 1999; Duffaud, 2000; Folie and Codrea, 2005). Unfortunately, the available material does not enable identification below the family level. The Lano association comprises at least five taxa: one indeterminate albanerpetontid Allocaudata (represented by incomplete dentaries, axis, and fragmentary humeri), one indeterminate Caudata (atlas, trunk vertebrae, humerus), and at least three anurans: one indeterminate Discoglossidae (maxillaries, angulars, vertebrae, urostyle, humeri, and ilia), one indeterminate Palaeobatrachidae (angulars, presacral vertebra, synsacrums, humeri, and ilia), and at least one indeterminate anuran (maxillaries, humeri and ilia that could belong to one or several species) (Duffaud and Rage, 1999; Duffaud, 2000). The palaeobatrachid remains are dominant relative to other taxa (about 70% of the lissamphibian assemblage). Interestingly, Lano has yielded one of the oldest Salamandridae (Duffaud and Rage, 1999; Duffaud, 2000). Moreover, the palaeobatrachid of Lano ranks among the oldest doubtless representatives of the family (Duffaud, 2000; Wuttke et al., 2012).

5.3. Lepidosaurs

Although the squamate remains from Lano are not really numerous (about 50 specimens), the assemblage is one of the richest and most diverse from the Late Cretaceous of Europe (Rage, 1999; Folie and Codrea, 2005). Lano has produced at least eight species: six "lacertilians" (a group that is considered to be paraphyletic) and two madtsoiid snakes still known only from Lano: Madtsoia laurasiae and Herensugea caristiorum (Rage, 1996, 1999). Recently, M. laurasiae has been removed from Madtsoia to Menarana, a genus known elsewhere only from the Maastrichtian of Madagascar (LaDuke et al., 2010).

"Lacertilians" of Lano include a non-acrodontan iguanian, i.e. Iguanidae sensu lato (represented by maxillary and other fragments with teeth), an indeterminate scincomorphan (maxillary or dentary fragments with teeth) and at least three other indeterminate species (based on isolated teeth) (Rage, 1999). One isolated tooth may belong to Paramacellodidae, a group of scincomorphans known in the Late Cretaceous of Europe (Folie and Codrea, 2005). Lano was the first locality to document the presence of Iguanidae s.l. in the Cretaceous of Europe (Rage, 1999). Several vertebrae from Lano have tentatively been referred to Amphisbaenia (Astibia et al., 1990). If the identification is correct, it is the oldest representative of this group in Europe. Nevertheless, this assignment remains uncertain as a referral to Anguidae cannot be ruled out. Then, it would be the oldest anguid recorded in Europe (Rage, 1999; Blain et al., 2010; Auge, 2012).

Several isolated vertebrae from Lano have been referred to indeterminate lacertilians (Rage, 1999). Two of them display a peculiar morphology, with a pachyostotic centrum and zygosphene-zygantrum articulations. This material may belong to a non-ophidian pythonomorph (i.e., varanoid; A. Houssaye, pers. comm.).

Snakes represent about half of the squamate remains from Lano. The two madtsoiid taxa known in the locality (Fig. 4) rank among the very rare members of this Gondwanan family in the Laurasian continents (Rage, 1996; LaDuke et al., 2010; Vasile et al., 2013). Menarana laurasiae is a large snake (estimated length 2.5 m, 7 cm in mid-body diameter; see LaDuke et al., 2010). Herensugea caristiorum is smaller in size (approximately 1 m in total length). It was probably a fossorial or secretive form (Rage, 1999). Finally, there is no evidence of "aniliid" snakes at Lano. The preliminary identification of Aniliidae (Astibia et al., 1990) was based on fragmentary vertebrae that actually belong to the madtsoiid Herensugea (Rage, 1999).

Recently, Apesteguia (2012) has suggested the presence of an eilenodontine sphenodontian in Lano on the basis of a fragmentary maxilla or dentary with acrodont dentition. However, the original description of this piece considered the tooth attachment as being pleurodont, so it was referred to an indeterminate lacertilian (Rage, 1999: Fig. 5).

5.4. Turtles

Turtle bones are the most dominant elements in the macrofossil assemblage of Lano. More than a thousand specimens have been found in Lano. Most of them correspond to disjointed plates and fragments of plates. However, some partial fragments of shells as well as isolated vertebrae and pelvic bones have also been collected (Lapparent de Broin and Murelaga, 1999). More than a hundred individuals are probably represented in the assemblage. The Lano turtles are currently attributed to three different species (Fig. 5). One of them is the solemydid Solemys vermiculata (stem Testudines). The other two taxa are members of Pan-Pleurodira: the dortokid Dortoka vasconica and the bothremydid Polysternon atlanticum (Lapparent de Broin and Murelaga, 1996, 1999; Perez-Garcia, 2009, 2012).

Nearly half of the turtle specimens found in Lano belong to Dortoka vasconica. It is the smallest turtle taxon recognized in the site, the estimated maximum length of the adults in the site, the estimated maximum length of the adults being less than 20 cm. Dortoka vasconica is characterized by the presence of a medial strong ornamentation of tubercles and minute pits, crests and ridges on the neurals and the medial region of the costal plates (Lapparent de Broin et al., 2004). The presence of a pair of large, persistent fontanelles on its carapace, with an autapomorphic configuration, together with its highly vascularised bone histology, indicates that it was an aquatic freshwater turtle (Fig. 5E; Perez-Garcia et al., 2012a). Dortoka is a representative of Dortokidae, an endemic European lineage of Pan-Pleurodira. Due to the absence of cranial material, its systematic position is a debated topic. Dortokids are generally considered as the sister taxon of Eupleurodira or the sister taxon of Pelomedusoides (Lapparent de Broin and Murelaga, 1999; Gaffney et al., 2006).

Bothremydids are the most abundant and diverse group of turtles from the Late Cretaceous of southern Europe (Lapparent de Broin and Murelaga, 1999; Gaffney et al., 2006; Perez-Garcia et al., 2012b, 2013). The carapace of Polysternon atlanticum (up to 32 cm long) is smaller than that of the other known species of this genus, Polysternon provinciale. Its outer surface shows "pelomedusoid ornamentation", composed of small dichotomous discontinuous grooves that may anastomose forming small and almost flat polygons (Lapparent de Broin and Murelaga, 1999). Polysternon is identified as a member of Foxemydina, a group to which belong all the bothremydids identified in the Cretaceous of Europe (Elochelys, Polysternon, Foxemys, Iberoccitane mys), except the Portuguese taxon Rosasia (Gaffney et al., 2006; Perez-Garcia et al., 2013). It has been interpreted that Polysternon probably lived in the deepest aquatic part of the fluvial environment of Lano (Lapparent de Broin and Murelaga, 1999).

Solemys vermiculata is the largest turtle in Lano: in the larger known adults, the length of the carapace reach about 70 cm. Solemys is decorated by granulations, close or separated, and by vermiculations (Lapparent de Broin and Murelaga, 1999). The study of both cranial and postcranial material of Solemydidae allows its identification as belonging to the stem group of Testudines (Joyce et al., 2011; Anquetin, 2012). The presence of osteoderms, known in the limbs of several representatives of Solemydidae, suggests terrestrial habitat (Lapparent de Broin and Murelaga, 1999; Joyce et al., 2011). This hypothesis has recently been confirmed by histological studies (Scheyer et al., 2015)

5.5. Crocodyliforms

Crocodyliform remains are abundant in Lano. Both cranial and postcranial bones have been found, most of them as disarticulated elements. Isolated teeth are by far the most abundant, including several hundred of specimens from smaller and larger individuals. On the basis of isolated skull bones (e.g., maxillae), mandibular remains and teeth from Lano, Buscalioni et al. (1997) erected Acynodon iberoccitanus and Musturzabalsuchus buffetauti (Fig. 6); these taxa were assigned to the Eusuchia as members of Alligatoroidea. In addition, Buscalioni et al. (1999) recognized two other taxa in Lano: an unnamed eusuchian, probably belonging to Al lodaposuchus, and Ischyrochampsa-like taxon with uncertain affinities. Isolated postcranial elements of Lano were provisionally regarded as indeterminate. At least procoelous vertebrae belong to eusuchians.

Acynodon is a brevirostrine eusuchian of small size (about 1 m long). The genus is characterized by a wide, short and flat snout and heterodont dentition without caniniform teeth. The dentition comprises a gradation of spatulated anterior teeth to globular (tribodont) posterior ones (Buscalioni et al., 1999; Martin, 2007). The absence of caniniform teeth coupled with the presence of enlarged molariform teeth suggests that Acynodon probably fed on slowly moving hard-shelled preys (Delfino et al., 2008). The phylogenetic analyses placed Acynodon as a basal member of Globidonta within Alligatoroidea (Buscalioni et al., 1999, Martin, 2007, Delfino et al., 2008). The description of new complete skulls from France and Italy (Martin, 2007; Delfino et al., 2008) as Acynodon promoted the revision of the European material to verify its alligatoroidean origin by other authors since an alternative phylogenetic hypothesis putatively placed this peculiar eusuchian within Hylaeochampsidae (Brochu et al., 2012).

Musturzabalsuchus is larger in size than Acynodon. The rostrum of Musturzabalsuchus is unique in having a maxilla with a conspicuous curvilinear lateral contour and a mid-constriction (Buscalioni et al., 1997, 1999). M. buffetauti is considered to be a basal member of Alligatoroidea (Buscalioni et al., 1999; NarvAez and Ortega, 2011). However, the status of this taxon remains uncertain and its relationships with other basal eusuchians from the Late Cretaceous of Europe, such as Allodaposuchus andMassaliasuchus, are an open question (Martin and Delfino, 2010).

The presence of a third eusuchian taxon in Lano distinct from Acynodon and Musturzabalsuchus is based on an isolated basicranial bone (Buscalioni et al., 1999). This material was referred to Allodaposuchus precedens in the revision made by Buscalioni et al. (2001). The genus Allodaposuchus is a major component of the Late Cretaceous crocodyliform faunas of Europe (Martin and Delfino, 2010), and it shows a wide spectrum of variation. Generally placed as the sister taxon of the Crocodylia crown group, it still needs a more comprehensive revision for a precise phylogenetic standing (Buscalioni et al., 2001, 2011; Pol et al., 2009; Martin, 2010; Puertolas et al., 2014).

Finally, isolated large teeth from Lano have been compared with those ofIschyrochampsa meridionalis (Buscalioni et al., 1999). This taxon was originally described as a member of Trematochampsidae (Vasse, 1995), a wastebasket taxon of ziphodont crocodyliforms. Ischyrochampsa has been regarded as a member of Neosuchia (Buscalioni et al., 2003), but its relationships remain currently unclear.

5.6. Dinosaurs

Lano has yielded numerous dinosaur remains. Titanosaurian sauropods are the best represented dinosaurs in number of specimens, followed by theropods (mostly teeth), ankylosaurs, and ornithopods. In number of species, theropods are the most diverse. At least ten distinct species have been identified in Lano; only the sauropod Lirainosaurus astibiae has been defined to date on the basis of the Lano material (Tables 1, 3).


The Lano theropods have not yet been described in detail: the material consists of several vertebrae and limb bones, and more than a hundred isolated teeth. As far as known, Lano is the richest site in theropod teeth from the Late Cretaceous of Europe (see Osi et al., 2010). Torices (2007) identified six different morphotypes, which could correspond to five species of small theropods and a sixth species of large size. All the small-sized theropod teeth (crown tooth height less than 17.5 mm) correspond to coelurosaurians: Coelurosauria indet., cf. Richardoestesia sp., a tiny maniraptoran that may be a new taxon, and two dromaeosaurids: cf. Pyroraptor olympus (originally described in the Trets locality of Provence; see Allain and Taquet, 2000), and Dromaeosauridae indet. (Torices, 2007; Torices et al., in press). In addition, two tooth morphotypes of large size (crown height up to 62 mm) could correspond to juvenile and adult individuals of the same taxon. These teeth are tentatively assigned to Theropoda indet. (Torices et al., in press). It has been discussed if these teeth could belong to neoceratosaurians but they lack ornamentation in the tooth enamel characteristic of this group and in the statistical analyses they showed stronger affinities to Tyrannosauridae. Due to these statistically uncertain affinities they are identified only as indeterminate theropods (Torices et al., in press). With regard to postcranial material, a pair of femora found at Lano has been compared to that of the abelisauroid Tarascosaurus from the Campanian of Provence (Le Loeuff and Buffetaut, 1991; Le Loeuff, 1992). Finally, an isolated tibiotarsus exhibits bird-like features and a partial sacrum first provisionally regarded as pterosaurian actually belongs to a large ground bird probably related to Gargantuavis (Buffetaut et al., 2006; and work currently in progress).


All the sauropod remains from Lano are referred to the titanosaur Lirainosaurus astibiae. This taxon was originally described on the basis of a skull fragment, isolated teeth, several vertebrae (e.g., holotypic anterior caudal vertebra; Fig. 7), appendicular bones and osteoderms (Sanz et al., 1999). New material, including basicranial, axial and appendicular elements, provides further information about the skeletal anatomy of Lirainosaurus (Diez Diaz et al., 2011, 2012, 2013a, 2013b, 2013c; Diez Diaz, 2013). Besides Lano, remains of Lirainosaurus have also described in other Iberian sites (Company et al., 2009; Ortega and Perez-Garcia, 2009; Diez Diaz, 2013). This taxon is diagnosed by autopomorphic features observed in the basicranium, vertebrae, and appendicular bones. More than a hundred teeth of Lirainosaurus have been collected in Lano; this sample is one of the largest known for titanosaurs. Tooth differences in size, morphology, ornamentation and microwear are regarded as ontogenetic changes; a switch in the diet and food processing between the juvenile and adult individuals has been hypothesized (Diez Diaz et al., 2012). Lirainosaurus was a small and slender titanosaur (estimated body size up to 6 m and 2 to 4 tn of body mass for the largest individuals; Diez Diaz, 2013; Diez Diaz et al., 2013b). It is considered to be a derived lithostrotian member of Saltasauridae. L. astibiae is the only titanosaurian species erected to date from the Late Cretaceous sites of the Iberian Peninsula.


Armoured dinosaurs are represented in Lano by a partial skeleton (synsacrum, pelvis and hindlimb bones) and scattered elements, including maxilla and lower jaw remains, isolated teeth, vertebrae and ribs, limb bones, and osteoderms (Pereda Suberbiola, 1993a, 1999). These remains have been assigned to the ankylosaur Struthiosaurus on the basis of features observed in the lower jaw, synsacrum, pelvis, and dermal armour (Pereda Suberbiola et al., 1995; Garcia and Pereda Suberbiola, 2003). Due to minor differences relative to known species of Struthiosaurus from the Campanian-Maastrichtian of Europe, the Lano material is tentatively referred to as Struthiosaurus sp. Struthiosaurus was a dwarf ankylosaur, with adults having a body length less than 3 m (Pereda Suberbiola and Galton, 2009). This dinosaur is a member of Nodosauridae (Pereda Suberbiola, 1993b; Thompson et al., 2012), the only known clade of ankylosaurs from the latest Cretaceous of Europe.


Ornithopod remains are scarce at Lano, but at least two taxa are represented. Isolated teeth, vertebrae and limb bones (humerus, femora) have been referred to the basal iguanodontian Rhabdodontidae Rhabdodon sp. (Pereda Suberbiola and Sanz, 1999). Rhabdodontids are a group of basal ornithopods endemic to Europe (Weishampel et al., 2003; Osi et al., 2012). In addition, Lano has yielded a single tooth belonging to an indeterminate hadrosauroid (Pereda Suberbiola et al., 2003). Consequently, this latter finding testifies of the co-existence of rhabdodontids and hadrosauroids in the latest Cretaceous of the Iberian Peninsula. Based on an incomplete ilium, the presence of a third ornithopod taxon in Lano is worthy of consideration (this material remains currently undescribed).

5.7. Pterosaurs

Lano is one of the most productive pterosaur sites from the Late Cretaceous in Europe. Pterosaur remains consist of an edentulous jaw fragment, elements of the vertebral column (cervical vertebrae, notarium), and limb bones (wing bones, femora) belonging to several individuals (Astibia et al., 1990; Buffetaut, 1999). Some of these bones have not yet been described in detail. Metacarpal (length 346 mm) and several phalanges of the fourth finger suggest a minimum wingspan of 3 to 3.5 m, which means they were large but not gigantic pterosaurs. The material obtained was referred to an indeterminate azhdarchid (Astibia et al., 1990), and later regarded as cf. Azhdarcho sp. because of close resemblances with Azhdarcho lancicollis from the Turonian of Uzbekistan in Central Asia, mainly on the basis of jaw morphology and general size (Buffetaut, 1999). Azhdarchid remains are relatively common in the latest Cretaceous continental sites of Europe (Company et al., 1999; Barrett et al., 2008; Buffetaut, 2008; Averianov, 2014) and new taxa have recently been described (Osi et al., 2005; Vremir et al., 2013 and references). Pending a full description of this material, the Lano pterosaur is provisionally referred to Azhdarchidae indet.

5.8. Mammals

The mammal assemblage of Lano is the richest known in the latest Cretaceous of southern Europe. It consists of 20 isolated teeth, i.e. molars, premolars, including deciduous teeth, and an incisor (Gheerbrant and Astibia, 1994, 1999, 2012). The faunule is composed exclusively of therian forms, and more specifically of zhelestid eutherians. Zhelestids are dentally ungulate-like mammals, but were recently excluded from Placentalia (Wible et al., 2009) and are instead considered as herbivorous stem eutherians of Cretaceous Asian origin (Gheerbrant and Astibia, 1999; Archibald and Averianov, 2012).

The zhelestids from Lano and other European sites are included in the subfamily Lainodontinae Gheerbrant and Astibia, 2012, which documents a modest mammal radiation of five or six species restricted to Europe. Three different species of the lainodontine genus Lainodon have been documented in Lano (Fig. 8): L. orueetxebarriai (Gheerbrant and Astibia, 1994, 1999), L. ragei and a still unnamed species (Gheerbrant and Astibia, 2012). The zhelestid mammals are characterized by a crushing-grinding dietary adaptation related to an early herbivorous trend among eutherians. The Lainodontinae (Zhelestidae) is the most diverse and dominant taxon in the Late Cretaceous mammalian faunas of Western Europe, by contrast to Central European sites that include only kogaionid multituberculates (Kielan-Jaworowska et al., 2004; Gheerbrant and Astibia, 2012). The Zhelestidae family has been treated as paraphyletic, but recent phylogenetic work supports its monophyly (Archibald and Averianov, 2012).

6. The vertebrate fossil assemblage from the shallow marine beds of Albaina

A rich vertebrate assemblage has also been recovered from some calcarenites, and in less quantity from interbedded friable sandstones, which form the uppermost beds of the depositional cycle DC-13. This shallow marine assemblage has yielded at least thirty seven species including sharks and rays, pycnodontiforms, teleosteans, mosasaurids, and plesiosaurs (Table 3). Most of the material consists of isolated teeth but other skeletal remains occur as indicated below. In addition, new vertebrate fossils collected in the Albaina beds consist of partial turtle plates and a fragmentary dinosaur femur (Pereda Suberbiola et al., in press).

6.1. Selachians

Cartilaginous fishes are represented by isolated teeth, indeterminate selachian dermal denticles, thorns and tail spines. The association recovered from the Albaina site, which was studied by Cappetta and Corral (1999), is particularly rich in shark and ray teeth and has yielded so far 15 genera, including two new genera, and 19 species of selachians, four of which are new rhinobatoid ray species (Fig. 9; see also Table 3).

Shark taxa consists of lamniforms (the anacoracids Squalicorax pristodontus and S. kaupi; the otodontid Cretolamna appendiculata, the serratolamnid Serratolamna serrata; the odontaspidids Carcharias heathi, Carcharias aff. gracilis and Odontaspis bronni), orectolobiforms (the ginglymostomatids Plitacoscyllium lehneri and the hemyscyllid Chiloscyllium sp.), and carcharhiniforms (the triakid Palaeogaleus faujasi). Rays include rajiforms (the rhinobatids Rhinobatos echavei and R. ibericus and the rhinobatoids incertae familiae Ataktobatis variabilis and Vascobatis albaitensis, all of them first defined in Albaina; the sclerorhynchids Dalpiazia stromeri and Ganopristis leptodon) and myliobatiforms species (the dasyatoid Coupatezia fallax, the rhombodontids Rhombodus andriesi and R. binkhorsti) (Cappetta and Corral, 1999; see Table 2 for the faunal list found in the locality).

Cappetta and Corral (1999) described the occurrence of Plicatoscyllium minutum (Forir, 1887) in Albaina. However, a closer study of the original material of Plicatoscyllium lehneri (Leriche, 1938) allowed to confirm it as a valid species, not a senior synonym of the former, and therefore the assignment of the Ginglymostomatidae teeth found in Albaina to Plicatoscyllium lehneri.

The selachian association of Albaina indicates a Maastrichtian age, and more precisely a Late--but not latest--Maastrichtian age according to the presence of Rhombodus andriesi and Odontaspis bronni species. In sum, the Albaina site is the most productive Late Cretaceous locality of fossil selachian remains in the Iberian Peninsula and among the most diverse Late Cretaceous sites of southwestern Europe.

6.2. Actinopterygians

The Albaina beds have yielded a relatively diverse actinopterygian ichthyofauna (Poyato-Ariza et al., 1999). Most of the remains consist of isolated teeth and tooth plates, although a few vertebral centra and fragments of fin spines are also known. The total number of specimens is about 800.

Poyato-Ariza et al. (1999) provided a conservative approach to the identification of the Albaina actinopterygian remains based on morphotypes rather than strict taxonomical identification. Twelve tooth morphotypes were distinguished, which may correspond to a maximum diversity of 12 taxa, including pycnodontiforms and teleosteans. Five morphotypes may correspond to pycnodonts: Pycnodontiformes indet. (pharyngeal teeth), Pycnodontoidei indet. A (flattened, ellipsoidal-like tooth), Pycnodontoidei indet. B (incisiform teeth), cf. Anomoeodus sp. (flattened, comma-shaped tooth), and cf. Paramicrodon sp. (vomerian tooth plate). Because of the heterodonty in pycnodonts, two or more of these morphotypes may coexist in the same taxon. The teleostean tooth morphotypes of Albaina may correspond to: cf. Elopiformes indet. (vertebral centra); the phyllodontids Phyllodus sp. (parasphenoid tooth plate and isolated teeth) and Paralbula sp. (basibranchial tooth plate and isolated teeth); the aulopiform Enchodus sp. (two morphotypes of caniniform teeth, which may or may not correspond to different species); and Acanthomorpha indet. (basal fragments of fin spines). The pharyngeal teeth tentatively previously assessed to cf. Stephanodus sp. fit into the morphologic variability of the pharyngeal teeth of pycnodonts and other groups (e.g., Thies, 1989; Poyato-Ariza and Wenz, 2002, 2005), so they probably do not represent a distinct taxon and are better considered Neopterygii indet. The most abundant taxon is by far Paralbula sp. (about 70% of the actinopterygian teeth), followed by Enchodus spp. (20%).

The actinopterygian association of Albaina is similar to that found in the nearby site of Quintanilla la Ojada (Burgos province) (Berreteaga et al., 2011), probably due to taphonomic processes that selectively destroyed less durable remains. As in other Iberian coeval localities, the associations of Albaina and Quintanilla la Ojada show a mixture of some teleosteans plus diversified relict, non-teleostean forms (such as pycnodontiforms). This is a nice example of the replacement process that would turn the actinopterygian composition of aquatic ecosystems during the Late Cretaceous in this part of the world into the modern, teleostean-dominated ichthyofaunas (Poyato-Ariza et al., 1999; Berreteaga et al., 2011; PoyatoAriza and Martin-Abad, 2013).

6.3. Mosasaurids

Mosasaurid material from Albaina consists primarily of a collection of isolated teeth and vertebrae. At least five distinct taxa have been recognized on the basis of tooth features: Mosasaurus hoffmanni, Mosasaurus sp., Prognathodon sectorius, Prognathodon solvayi, Prognathodon sp., and Platecarpus cf. ictericus (Bardet et al., 1997, 1999, 2013). As in other latest Cretaceous mosasaurid associations, Mosasaurinae taxa are dominant relative to Russellosaurina ones. Albaina has yielded the most diverse mosasaurid assemblage found to date in the Iberian Peninsula (Bardet et al., 2008, 2013).

6.4. Plesiosaurs

Plesiosaurs are represented by a single elasmosaurid tooth (Bardet et al., 1999). This is the only record of Elasmosauri dae in the Late Cretaceous of the Iberian Peninsula (Bardet et al., 2008), and one of the scarce mentions of this clade in the Maastrichtian of Europe (Vincent et al., 2011).

6.5 Turtles

The material consists of two partial turtle plates, one of them belonging to a member of Bothremydidae (Pleurodira) and the other to an indeterminate taxon, probably corresponding to a Pan-Cryptodira (Pereda-Suberbiola et al., in press).

6.6. Dinosaurs

The first dinosaur fossil found in the sublittoral beds of Albaina is a fragmentary ornithopod femur, which can be interpreted as the result of the passive transport of a floating carcasse from the mainland. It is one of the few hadrosauroid remains from the Late Maastrichtian of Europe found in marine environments, and the first one described from sublittoral deposits in the Iberian Peninsula (Pereda-Suberbiola et al., in press).

7. Significance of the vertebrate associations from the Lano quarry.

The Lano quarry is one of the most noteworthy Late Cretaceous vertebrate localities of Europe, taking into account both fossil richness and taxonomic diversity. Considering the terrestrial and freshwater vertebrates, at least 37 species are known in the Late Campanian to Early Maastrichtian beds, making it the most diverse association hitherto known for this age in Europe (Astibia et al., 1990, 1999b). Of the nearly 40 tetrapod families (or higher-level taxa) recorded in the Santonian-Maastrichtian of Europe (Pereda Suberbiola, 2009), at least 23 are known in Lano. Lano enlarges our knowledge of the latest Cretaceous faunas: it includes one of the oldest records of both Salamandridae and Palaeobatrachidae lissamphibians in the world (Duffaud and Rage, 1999), and one of the oldest records of amphisbaenians or anguids in Europe (Rage, 1999). The squamate association is one of the richest and most diverse known from the Cretaceous of Europe (Rage, 1999, 2013; LaDuke et al., 2010). Besides, the Lano fluvial beds have yielded fossil remains leading to the erection of a number of reptilian taxa that are widely represented in the latest Cretaceous localities of Europe, such as the turtles Solemys and Dortoka (Lapparent de Broin and Murelaga, 1999; Perez-Garcia et al., 2012a) and the crocodilians Acynodon andMusturzabalsuchus (Buscalioni et al., 1999; Martin and Delfino, 2010). With regard to the dinosaurs, the non-avian theropod association of Lano is the most diversified of the Campanian-Maastrichtian of Europe (Torices et al., in press); and the sauropod Lirainosaurus is currently the best known titanosaur of Europe (Diez Diaz et al., 2011, 2012, 2013a, 2103b; Diez Diaz, 2013). Finally, the mammalian faunule of Lano is one of the richest hitherto discovered in Europe and provides data on the evolution of early eutherians (Gheerbrant and Astibia, 2012).

The vertebrate fossil association of Lano is considered to be a typical continental faunal assemblage from the Late Campanian-Early Maastrichtian of the Ibero-Armorican Domain (SW Europe). Other relevant sites of the same age are: Chera in Valencia (Company et al., 1999) and Lo Hueco in Cuenca (Ortega et al., 2008, 2015), in the Iberian Peninsula; Bellevue in Campagne-sur-Aude (Aude), Cruzy (Herault), La Boucharde in Trets, Velaux-Bastide Neuve (both in Bouchesdu-Rhone), and Fox-Amphoux (Var), in Languedoc and Provence (Le Loeuff, 1992; Garcia et al., 2000, 2010; Laurent et al., 2001; Allain and Pereda Suberbiola, 2003; Buffetaut, 2005). These vertebrate fossil assemblages are mainly dominated by titanosaurian sauropods, rhabdodontid ornithopods, nodosaurid ankylosaurs and dromaeosaurid theropods among dinosaurs, alligatoroid crocodyliforms, bothremydid and solemydid turtles, and zhelestid mammals. Other common components of the faunas are lepisosteid actinopterygians, the dortokid turtle Dortoka, the basal eusuchian Allodaposuchus and closely related forms, abelisauroid theropods and azhdarchid pterosaurs. A number of taxa that have been recorded in other Ibero-Armorican sites are apparently absent in Lano, i.e., sparid fishes, batrachosauroid urodeles, sebecosuchian-like crocodilians, and enantiornithine birds (see Le Loeuff, 1991; Buffetaut et al., 1997; Pereda Suberbiola, 2009; Csiki et al., 2015).

The continental vertebrate faunas from the CampanianMaastrichtian of southwestern Europe show biogeographical affinities with those of the Laurasian landmasses, either from Palaeolaurasia (sensu Russell, 1993, i.e., Central Asia with North America and probably Europe), Euramerica, Asia or endemic to Europe, but also contain Gondwanan elements (Astibia et al., 1990; Le Loeuff, 1991; Rage, 2002; Pereda Suberbiola, 2009). The picture that emerges is that of continental faunas that evolved isolated in the European archipelago as a result of vicariance during the Late Cretaceous. The isolation from other landmasses may have facilitated the survival of relict (vicariant) taxa in Europe until Campanian-Maastrichtian times. Moreover, dispersal events between Asia (and, tentatively, between North America) and Europe have been documented during the Late Cretaceous (see Pereda Suberbiola, 2009; Weishampel et al., 2010; Prieto-Marquez et al., 2013; Csiki et al., 2015 and references therein). The faunal exchanges that are hypothesized to occur between the Gondwanan landmasses and Europe during the Late Cretaceous are still matters of debate (Gheerbrant and Rage, 2006).

With regard to the marine vertebrate association of Albaina, which is composed of selachians, actinopterygians, mosasaurids, and plesiosaurs, as noted earlier, at least 37 species are recorded from these Late Maastrichtian shallow sublittoral beds. As a result, this is the most diverse assemblage of marine vertebrates found so far in the latest Cretaceous of the Ibero-Armorican Realm (southern Europe). The selachian association of Albaina is original in showing a mixture of both southern Tethyan (Moroccan phosphatic basins) and northern European boreal (Belgian and German basins) species, but also containing some particular taxa, i.e. rhinobatoids (Cappetta and Corral, 1999; Corral, in prep.). The mosasaurid assemblage of Albaina is typical of the Northern Tethys margin palaeoprovince located around palaeolatitudes 30[degrees]-40[degrees] N (Bardet, 2012). Recently, dinosaur and turtle remains have also been found in the Late Maastrichtian sublittoral beds of Albaina (Pereda-Suberbiola et al., in press). Turtle fossils belong to a member of Pleurodira (Bothremydidae) and probably to an indeterminate Pan-Cryptodira. The occurrence of dinosaurs in this shallow marine environment can be interpreted as the result of a passive transport from the mainland.


We would like to thank all of the people involved in the excavation and study of the fossil vertebrates from the Lano quarry, especially Mr. Juan Echave (ECHASA) and Mr. Jesus Alonso (Museo de Ciencias Naturales de Alava/Arabako Natur Zientzien Museoa, Vitoria-Gasteiz). We are grateful to the referees Drs. Attila Osi (Budapest) and David B. Weishampel (Baltimore) for their constructive comments on the manuscript. Field research in the Lano quarry has been supported by financial aids of the Ministerio de Economia y Competitividad (MINECO projects CGL2010-18851/BTE and CGL2013-47521-P), Ministerio de Ciencia e Innovacion (CGL2007-64061/BTE), Ministerio de Ciencia y Tecnologia (CGL2004-02338/BTE, BTE2001-0185-C02-01, BOS20001369), Ministerio de Educacion y Ciencia (Accion Integrada hispano-francesa 201-B), Diputacion Foral de Alava/Arabako Foru Aldundia (93/23, 95/A23), Gobierno Vasco/Eusko Jaurlaritza (IT834-13, IT-320-10, IT-361-07, GV121.310-4/87), Universidad del Pais Vasco/Euskal Herriko Unibertsitatea (9/ UPV00121.310-15303/2003), Centre National de la Recherche Scientifique (CNRS, France), Institut National des Sciences de l'Univers (INSU), The Dinosaur Society (research grant 1997) and the National Geographic Research (grant #6597-99). This work is part of a palaeontological collaboration (Convenio especifico de colaboracion) between the Universidad del Pais Vasco/EHU, the Centre National de la Recherche Scientifique (CNRS, France) and the Museum National d'Histoire Naturelle (MNHN, Paris). Contribution ISEM n[degrees] 2015-049.


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X. Pereda-Suberbiola (1) *, J.C. Corral (2), H. Astibia (1), A. Badiola (1), N. Bardet (3), A. Berreteaga (1), E. Buffetaut (4), A.D. Buscalioni (5), H. Cappetta (6), L. Cavin (7), V. Diez Diaz (1), E. Gheerbrant (3), X. Murelaga (1), F Ortega (8), A. Perez-Garcia (9), F. Poyato-Ariza (5), J.-C. Rage (3), J.L. Sanz (5), A. Torices (8,10)

(1) Universidad del Pais Vasco/Euskal Herriko Unibertsitatea, Facultad de Ciencia y Tecnologia, Departamento de Estratigrafia y Paleontologia, Apartado 644, 48080 Bilbao, Spain.

(2) Arabako Natur Zientzien Museoa/Museo de Ciencias Naturales de Alava, Siervas de Jesus, 01001 Vitoria-Gasteiz, Spain.

(3) Sorbonne Universites, CR2P CNRS-MNHN-UPMCParis 6, Departement Histoire de la Terre, Museum National d'Histoire Naturelle, CP 38, 57 rue Cuvier, 75005 Paris, France.

(4) CNRS UMR 8538, Laboratoire de Geologie de l 'Ecole Normale Superieure, 24 rue Lhomond, 75231 Paris Cedex 05, France.

(5) Unidad de Paleontologia, Departamento de Biologia, c/Darwin 2, Universidad Autonoma de Madrid, Cantoblanco, 18049 Madrid, Spain.

(6) Institut des Sciences de l'Evolution (UM2, CNRS, IRD), Universite Montpellier 2, Place Eugene Bataillon, 34095 Montpellier Cedex 5, France.

(7) Museum d'Histoire Naturelle, CP 6434, 1211 Geneva 6, Switzerland.

(8) Grupo de Biologia Evolutiva, Facultad de Ciencias, UNED, Senda del Rey, 9, 28040 Madrid, Spain,

(9) Centro de Geologia, Faculdade de Ciencias, Universidade de Lisboa, Edificio C6, Campo Grande, 1749-016Lisbon, Portugal.

(10) Department of Biological Sciences, Faculty of Science, CW405 Biological Sciences Centre, University of Alberta, Edmonton, Alberta, Canada T6G 2E9.

e-mail addresses: (X.P.-S, * corresponding author); (J.C.C.); humberto.astibia@ehu. eus (H.A.); (A.Ba.); (N.B.); (A.Be.), (E.B.); angela. (A.D.B.); (H.C.); (L.C.); (V.D.D.); (E.G.); (X.M.); (F.O.); (A.P.-G.); francisco. (FJ.P.-A.); (J.C.R.); (J.L.S.); (A.T.)

Received: 23 December 2013 / Accepted: 18 December 2014 / Available online: 25 March 2015

Table 1. Updated list of continental vertebrates from Lano (Late
Campanian-Early Maastrichtian).

                                                L1A   L1B   L2

       Lepisosteidae indet.                     X     X
       Phyllodontidae indet.                    X
       ?Palaeolabrus sp.                        X
       Albanerpetonidae indet.                  X
       Salamandridae indet.                     X
       Discoglossidae indet.                    X
       Palaeobatrachidae indet.                 X
       Anura indet. (one or several species)    X
       Iguanidae indet.                         X
       Scincomorpha indet.                      X
       Lacertilia indet. (3 species)            X
       Amphisbaenia or Anguidae indet.          X
       Menarana laurasiae                       X     X     X
       Herensugea caristiorum                   X
       ?Eilenodontinae indet.                   X?
  Stem Testudines
       Solemys vermiculata                      X     X
       Dortoka vasconica                        X     X
       Polysternon atlanticum                   X           X
       Bothremydidae indet.                                 X
       ?Ischyrochampsa sp.                      X           X
       Allodaposuchus sp.                       X
       Acynodon iberoccitanus                   X     X     X
       Musturzabalsuchus buffetauti             X     X     X
       cf. Tarascosaurus sp.                    X     X
       Coelurosauria indet.                     X
       cf. Richardoestesia sp.                  X
       Maniraptora indet.                       X
       Paraves indet.                           X
       cf. Pyroraptor olympus                   X
       cf. Dromaeosauridae indet.               X
       Lirainosaurus astibiae                   X     X     X
       Ornithopoda indet.                       X
       Rhabdodon sp.                            X     X     X
       Hadrosauroidea indet.                    X
       Struthiosaurus sp.                       X     X     X
       Azhdarchidae indet.                      X     X
       Lainodon orueetxebarriai                 X
       Lainodon ragei                           X
       Lainodon sp.                             X

Table 2. List of vertebrate taxa erected on the basis of material found
in the Late Cretaceous of the Lano quarry.

HIGHER TAXA                          GENERA

LANO (continental unit, Late Campanian to Early Maastrichtian)

Dortokidae Lapparent de Broin and    Dortoka Lapparent de Broin and
Murelaga, 1996                       Murelaga, 1996

Solemydidae Lapparent de Broin and   Solemys Lapparent de Broin and
Murelaga, 1996                       Murelaga, 1996

                                     Herensugea Rage, 1996

                                     Acynodon Buscalioni, Ortega and
                                     Vasse, 1997

                                     Musturzabalsuchus Buscalioni,
                                     Ortega and Vasse, 1997

                                     Lirainosaurus Sanz, Powell, Le
                                     Loeuff, Martinez and Pereda
                                     Suberbiola, 1999

Lainodontinae Gheerbrant and         Lainodon Gheerbrant and Astibia,
Astibia, 2012                        1994

ALBAINA (marine unit, Late
Maastrichtian) Rays                  Ataktobatis Cappetta and Corral,

                                     Vascobatis Cappetta and Corral,

HIGHER TAXA                          SPECIES

LANO (continental unit, Late Campanian to Early Maastrichtian)

Dortokidae Lapparent de Broin and    Dortoka vasconica Lapparent de
Murelaga, 1996                       Broin and Murelaga, 1996
                                     Polysternon atlanticum Lapparent
                                     de Broin and Murelaga, 1996

Solemydidae Lapparent de Broin and   Solemys vermiculata Lapparent de
Murelaga, 1996                       Broin and Murelaga, 1996

                                     Herensugea caristiorum Rage, 1996
                                     Menarana laurasiae (Rage, 1996)
                                     Acynodon iberoccitanus Buscalioni,
                                     Ortega and Vasse, 1997

                                     Musturzabalsuchus buffetauti
                                     Buscalioni, Ortega and Vasse, 1997

                                     Lirainosaurus astibiae Sanz,
                                     Powell, Le Loeuff, Martinez and
                                     Pereda Suberbiola, 1999

Lainodontinae Gheerbrant and         Lainodon orueetxebarriai
Astibia, 2012                        Gheerbrant and Astibia, 1994

                                     Lainodon ragei Gheerbrant and
                                     Astibia, 2012

ALBAINA (marine unit, Late
Maastrichtian) Rays                  Ataktobatis variabilis Cappetta
                                     and Corral, 1999

                                     Vascobatis albaitensis Cappetta
                                     and Corral, 1999

                                     Rhinobatos echavei Cappetta and
                                     Corral, 1999

                                     Rhinobatos ibericus Cappetta and
                                     Corral, 1999

Table 3. Updated list of vertebrates from the shallow marine beds of
Albaina (Late Maastrichtian).


                       Squalicorax pristodontus
                       Squalicorax kaupi
                       Cretolamna appendiculata
                       Serratolamna serrata
                       Carcharias heathi
                       Carcharias aff. gracilis
                       Odontaspis bronni
                       Plicatoscyllium lehneri
                       Chiloscyllium sp.
                       Palaeogaleus faujasi
                       Rhinobatos echavei
                       Rhinobatos ibericus
                  Rhinobatoidei incertae familiae
                       Ataktobatis variabilis
                       Vascobatis albaitensis
                       Dalpiazia stromeri
                       Ganopristis leptodon
                  Dasyatoidea incertae familiae
                       Coupatezia fallax
                       Rhombodus binkhorsti
                       Rhombodus andriesi
      Cf. Pan-Cryptodyra indet.
        cf. Polysternon atlanticum
                        Pycnodontiformes indet.
                        Pycnodontoidei indet. A
                        Pycnodontoidei indet. B
                        cf. Anomoeodus sp.
                        cf. Paramicrodon sp.
                        cf. Elopiformes indet.
                          Phyllodus sp.
                          Paralbula sp.
                          Enchodus sp. type A
                          Enchodus sp. type B
                          Acanthomorpha indet.
               Neopterygii indet. (pharyngeal teeth;
                       formerly cf. Stephanodus sp.)
                       Mosasaurus hoffmanni
                       Mosasaurus sp.
                       Prognathodon sectorius
                       Prognathodon solvayi
                       Prognathodon sp.
                       Platecarpus cf. ictericus
                       Elasmosauridae indet.
        Hadrosauroidea indet.
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Title Annotation:texto en ingles
Author:Pereda-Suberbiola, X.; Corral, J.C.; Astibia, H.; Badiola, A.; Bardet, N.; Berreteaga, A.; Buffetaut
Publication:Journal of Iberian Geology
Date:Jan 1, 2015
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