Journal of Iberian Geology https://doi.org/10.1007/s41513-023-00220-y RESEARCH PAPER A new vertebrate assemblage from the matute formation of the Cameros Basin (Ágreda, Spain): implications for the diversity during the jurassic/cretaceous boundary E. Puértolas-Pascual1 · M. Aurell1 · D. D. Bermúdez-Rochas2 · J. I. Canudo1 · A. E. Fernandes3,4 · A. Galobart5 · M. Moreno-Azanza1,6 · A. Pérez-García7 · D. Castanera8 Received: 15 March 2023 / Accepted: 13 September 2023 © The Author(s) 2023 Abstract The Ribota site (Ágreda, Soria, Spain) is a new locality in the Matute Formation (Tithonian–Berriasian) composed of several carbonate layers, outstandingly rich in macrovertebrate remains. Fossils show an unusual replacement of the original bioapatite by quartz, and are found as positive reliefs protruding from lacustrine limestone beds. This type of conservation has allowed the identification of around one hundred vertebrate bone accumulations in an outcrop of more than 10 hectares. Osteichthyans (articulated partial skeletons, cranial material, and isolated postcranial bones and scales), crocodylomorphs (disarticulated cranial material, isolated teeth, vertebrae and osteoderms), turtles (partial carapaces and plastra, but also isolated plates) and pterosaurs (cranial and appendicular elements) have been identified. Around 80 speci- mens have been collected and a preliminary study of part of the collection (35 specimens) has allowed the identification of at least 5 different taxa: Halecomorphi indet., Neoginglymodi indet., Goniopholididae indet., Testudinata indet., and Pterodactyloidea indet. This new site represents one of the few sites from this time interval preserved in a fully lacustrine environment, so these vertebrate assemblages are unique and composed of different animals that presumably lived around and within the lake. They are dominated by aquatic and amphibian vertebrates and was formed by attrition in this lacus- trine environment, possibly far from the lake shoreline. These macrovertebrate assemblages provide new data about the diversity in the faunal ecosystems from the Jurassic/Cretaceous transition of the Iberian Basin Rift System. Keywords Osteichthyes · Crocodylomorpha · Testudinata · Pterosauria · Tera Group · Tithonian–Berriasian Resumen El yacimiento de Ribota (Ágreda, Soria, España) es una nueva localidad en la Formación Matute (Titoniense-Berriasiense) compuesta por varios estratos de calizas excepcionalmente ricos en restos de macrovertebrados. Los fósiles muestran una sustitución inusual del bioapatito original por cuarzo, y se encuentran como relieves positivos sobresaliendo de la superficie de las calizas lacustres. Este tipo de conservación ha permitido la preservación e identificación de alrededor de cien acumulaciones de huesos de vertebrados en una superficie de más de 10 hectáreas. Se han identificado peces osteíctios (esqueletos parciales articulados, material craneal y escamas y huesos postcraneales aislados), cocodrilomorfos (material craneal desarticulado, dientes aislados, vértebras y osteodermos), tortugas (caparazones y plastrones pasciales, pero también placas aisladas) y pterosaurios (elementos craneales y apendiculares). Se han recuperado alrededor de 80 especímenes y un estudio preliminar de parte de la colección (35 especímenes) ha permitido la identificación de al menos 5 taxones diferentes: Halecomorphi indet., Neoginglymodi indet., Goniopholididae indet., Testudinata indet. y Pterodac- tyloidea indet. Este nuevo yacimiento representa uno de los pocos yacimientos de este intervalo temporal conservados en un ambiente completamente lacustre, por lo que estas asociaciones de vertebrados son únicas y están compuestas por Extended author information available on the last page of the article 13 Journal of Iberian Geology diferentes animales que presumiblemente vivieron alrededor y dentro del lago. Están dominadas por vertebrados acuáticos y anfibios, y se formaron por atrición en este entorno lacustre, posiblemente dentro del lago y lejos de la orilla. Estas asociaciones de macrovertebrados proporcionan nuevos datos sobre la diversidad faunística en los ecosistemas presentes durante la transición Jurásico/Cretácico del Sistema de Rift de la Cuenca Ibérica. Palabras clave Osteichthyes · Crocodylomorpha · Testudinata · Pterosauria · Grupo Tera · Titoniense–Berriasiense most probably formed during the Tithonian–Berriasian 1 Introduction transition (Aurell et al., 2021). This scarce vertebrate fossil content contrasts with the relatively abundant osteological The sedimentary successions deposited during the Jurassic/ record (mostly sauropod dinosaurs) with similar age, found Cretaceous transition in the eastern Cameros basin (north in the western Cameros basin (mainly in Burgos Province) central Spain) are well known for their large number of (e.g., Torcida Fernández-Baldor et al., 2020), in other areas tracksites. The diverse ichnofauna of the Berriasian depos- of the Iberian Range, such as the Maestrazgo and South Ibe- its (Oncala Group) of the Huérteles Formation in the North rian basins (northwest of Valencia and southwest of Teruel of Soria province stands out. This region has such an ich- provinces) (e.g., Ruiz-Omeñaca et al., 2004; Aurell et al., nological richness, (containing more than 7,000 vertebrate 2016; Campos-Soto et al., 2019), or in the northern Aquita- footprints produced mainly by dinosaurs, but also from ine Basin in SW France (Allain et al., 2022). pterosaurs, crocodylomorphs and turtles [e.g., Fuentes Vid- Here we present the new palaeontological site of Ribota arte et al., 2005; Hernández Medrano et al., 2006; Castanera (Ágreda, Soria province, Spain) (Fig. 1), which contains one et al., 2018]), that it gave rise to the creation of the touristic of the most abundant vertebrate fossil assemblages of the project “Ichnite Route of Soria” (Barco et al., 2013; Cas- whole Cameros basin. Until now, hundreds of bone remains tanera et al., 2018), where a selection of dinosaur tracksites (Figs. 2, 3, 4, 5, 6, 7 and 8) that correspond to osteichthyan are prepared for tourist visits. This contrasts with the very fishes (Fig. 5), crocodylomorphs (Fig. 6), turtles (Fig. 7) scarce osteological fossil record found in the geological and pterosaurs (Fig. 8) have been identified, and some of units that span around the Jurassic/Cretaceous (Tithonian– them, including all the remains described in this work, Berriasian) transition of the Tera Group (Agreda, Magaña were recovered from the field. This paper aims to present and Matute Fms) in this sector of the Cameros basin. a first approach to the genesis of this site (Figs. 3 and 4) One of the Tithonian–Berriasian geological units from and its palaeontological assemblage, in order to reconstruct the eastern Cameros basin that has yielded some fossil bones the ecosystem of this area of the Cameros basin around the is the Matute Fm (Salomon, 1982a, b;mez Fernández and Jurassic/Cretaceous boundary. Meléndez, 1994). The latter authors indicate that remains of fishes and reptiles are relatively frequent in this unit, how- ever only two notable specimens have been described in this 2 Geographic and geological context formation. One corresponds to an articulated ginglymodian actinopterygian fish described as Camerichthys lunae Ber- The Ribota fossil site is located 5 km northeast from the múdez-Rochas & Poyato-Ariza, 2015, from the locality of village of Ágreda (Soria Province, Castilla y León, north- San Andrés de San Pedro (northeast of Soria province). The central Spain), a town located about 10 km northwest of the second is the freshwater aquatic stem turtle Pleurosternon Moncayo, the highest mountain of the Iberian Chain. The moncayensis Pérez-García et al., 2022 (represented by a par- site outcrops in a sparsely vegetated area adjacent to the tial skull and several postcranial remains of a pleurosternid), north of a meander of the Val River, about 300 m northwest recovered from the surroundings of Ágreda (east of Soria of the “Pozo de las Truchas” waterfall. province), and only two kilometers away from the site here Geologically, Ribota is located within the Cameros basin. studied. In addition, also within the eastern Cameros basin This basin is found in the northwestern part of the Iberian in the town of Tera (north of Soria), some dinosaur remains Basin Rift System (Fig. 1A), which was formed due to of similar age from the Magaña Fm have also been reported extensional tectonics during the latest Jurassic–Early Creta- (Canudo et al., 2010), and an articulated fish assigned to ceous rifting stage. This rifting has been related to the open- Lepidotes sp. has been described from the Berriasian of ing of the Western Tethys Sea and the Northern Atlantic the Valdeprado Fm (Pascual-Arribas et al., 2007). South of Ocean (see Salas et al., 2001; Mas et al., 2002, 2004, 2011; Soria, in the Torrelapaja subbasin, the vertebrate fossil site Aurell et al., 2019, 2021 and references therein). The Ribota of La Atalaya (Bijuesca Fm) includes large sized sauropod site is located in the easternmost part of the eastern Cameros and crocodylomorph remains preserved in palustrine facies, basin (Fig. 1B, C). 13 Journal of Iberian Geology Fig. 1 A) Geographic and geo- logical location of the Ribota site (Soria, Castilla y León, Spain) within the Iberian Peninsula; B) geological location of the site within the Cameros basin, modi- fied from Rodríguez-Barreiro et al. (2022); C) detailed geological map of the area and its strati- graphic framework, modified Gómez Fernández and Meléndez (1994) The depocenter of the Cameros basin is filled with more The late Tithonian–earliest Berriasian age assumed here for than 8,000 m of Upper Jurassic–Lower Cretaceous lacus- the Matute Fm (Fig. 1C; see also Pérez-García et al., 2022) trine/coastal carbonates and evaporites alternated with is based on the charophyte and ostracod content described fluvial siliciclastic deposits (Salas et al., 2001; Mas et al., in previous works (Gómez Fernández and Meléndez 1994; 2004). Tischer (1996) divided this succession into five Tejero & Fernández-Gianotti, 2004; Schudack & Schudack, groups (Tera, Oncala, Urbión, Enciso, and Oliván). Suc- 2009). cessive genetic units were recognised, and the Tera Group The Ribota site is located in the lower part of the lacus- was divided into three alloformations (Gómez-Fernández & trine carbonates of the Matute Fm (Fig. 1C), within a car- Meléndez, 1994), that were subsequently used as lithostrati- bonate-dominated interval outcropping over a large area of graphic units (e.g., Mas et al., 2004; Clemente, 2010; Mar- about 10 hectares (Fig. 2A). Ribota is a rich locality where tín-Chivelet el al., 2019). The lower Ágreda Fm consists of isolated bones are found in almost every square meter of the shallow marine, alluvial, lacustrine to palustrine deposits; outcrop. Based on the closeness of several specific points the middle Magaña Fm is characterised by fluvial facies, and with the higher bone concentrations, Ribota can be divided the upper Matute Fm (also known as the Sierra Matute Fm) into three main areas (north, south, and east, Fig. 2A). is dominated by lacustrine deposits and includes the Ribota The north and south areas are stratigraphically equivalent site studied in this work (Fig. 1C). This unit was originally (almost the same strata), whereas the east area corresponds interpreted as early Berriasian in age (Salomon, 1982a, b). to slightly lower strata located about 150 m east from the The Berriasian age assignment for this unit, based on biostra- other areas, but in the same formation and similar lime- tigraphy, was also supported by Martín-Closas and Alonso stone facies. The division into 3 areas is provisional until a Millán (1998), Salas et al. (2001), Mas and Salas (2002) more exhaustive stratigraphic study is carried out. The most or Clemente (2010). However, other authors considered the important fossiliferous accumulations are in the south area Matute Fm as Tithonian in age (Mas et al., 1993; Gómez- within the same stratigraphic level. Except for some artic- Fernández & Meléndez, 1994), although the top of the unit ulated partial skeletons of fishes and turtles (exclusively could even reach the beginning of the Berriasian (Gómez- represented by shell elements), most of the remains appear Fernández, 1992; Meléndez & Gómez-Fernández, 2000). scattered and disarticulated along the different outcrops in Accordingly, the precise age of this unit remains uncertain. the area. 13 Journal of Iberian Geology Fig. 2 A) Google Earth view of the Ribota site showing the different small outcrops with ver- tebrate remains; B) photograph and general view of the first site with vertebrate remains found in the east area, point “Fig. 1B” in the satellite photo in (A); C) fossiliferous point with a high concentration of disarticulated fish bones and a detail zoom of these remains (D) and a pyrite crystal pointed by a blue arrow; E) some pyrite crystals found in Ribota area The studied geological samples are very fossiliferous taxonomic identification (Table 1). The fossils, together with wackestone limestones, whereas the most common fossils their surrounding rock matrix, were delimited in blocks cut are disarticulated ostracod valves, followed by vertebrate with a radial saw and extracted with a chisel and hammer. remains. All the samples were collected in the southern zone Each collected sample has been labelled with the acronym of Ribota, except for the RB72 sample that was recovered RB (from Ribota) and the subsequent field number (RB1 in the eastern area (Fig. 2A). All fossiliferous remains show to RB79), and they have been deposited (fossil numbers, a variable degree of recrystallization that might be related nº 2023/6/RB1 to 2023/6/RB79) in the Museo Numantino, to the low-grade hydrothermal metamorphism that affected provincial de Soria (Soria, Spain). For simplicity, through- the Matute Formation (e.g., Barrenechea et al., 2001). The out the text, each specimen is just referred with the field invertebrate remains have been completely replaced by number (RB1 to RB79). silica (probably quartz), whereas the vertebrate remains Eight standard (30-micron thin) sections of selected bone partially retain their original apatite composition, being specimens within limestone matrix (one Testudinata RB5; partially replaced by quartz and calcite. Isolated calcite and two Neoginglymodi RB11 and RB26; and four Crocody- dolomite crystals can be found dispersed in a chlorite-domi- lomorpha RB6, RB52, RB72 and RB74) where prepared, nated matrix, where clay mineral crystals show no preferred in order to analyse the fossil diagenesis processes at the orientation. Secondary idiomorphic pyrite crystals of vari- Ribota site. The thin sections were examined and photo- able size, from sub-millimetric to centimetric, have grown graphed with an Olympus BX53M petrographic polarizing both in the matrix and inside the bone elements. These microscope equipped with a digital camera, housed at the pyrite crystals are partially oxidised, showing concentric IUCA Microscopy Lab at Universidad de Zaragoza. After rings of hematite and goethite, and present very small irreg- this preliminary analysis, samples where carbon-coated ular inclusions of relics of apatite and calcite, incorporated and analysed in a Merlin Carl Zeiss Field Emission Scan- into the crystal during growth. ning Electron Microscope housed at the Servicio de Apoyo a la Investigación (SAI) of the Universidad de Zaragoza. Chemical and textural information on the mineral phases, 3 Materials and methods both in the fossils and in the encasing matrix, was obtained using Backscattered electron images (BSE) and Energy Dis- To date, almost 80 vertebrate fossils have been recovered persive X-Ray Analysis (EDS) and were acquired at 15 kV in the area. The material has been collected under the per- and an IProbe of 600 pA, at a working distance of 5.5 mm, mit Exp. 37/2019-SO issued by the Dirección General de using a Cobalt standard for calibration of the semiquantita- Patrimonio Cultural of the Junta de Castilla y León. Here tive analysis. we study only those samples that allowed a preliminary 13 Journal of Iberian Geology Table 1 List of the studied mate- Osteichthyes rial showing the identification, Acronym Material Taxa Area taxonomy and location in the outcrop RB 11 articulated partial skeleton Neoginglymodi indet. South area RB13 dentary Halecomorphi indet. East area RB14 articulated partial skeleton Neoginglymodi indet. East area (exsitu) RB26 articulated partial skeleton Neoginglymodi indet. South area RB27 dentary Neoginglymodi indet. South area RB58 dermopalatine fragment Neoginglymodi indet. East area Crocodylomorpha Acronym Material Taxa Area RB6 diaphysis fragment Goniopholididae indet. South area RB17 disarticulated cranial remains Goniopholididae indet. North area RB21 caudal vertebra Goniopholididae indet. South area RB35 ventral osteoderm Goniopholididae indet. South area RB39 isolated tooth Goniopholididae indet. South area RB44 dorsal vertebra Goniopholididae indet. South area RB52 tooth Goniopholididae indet. South area RB57 cervical vertebra Goniopholididae indet. South area RB61 dorsal osteoderm Goniopholididae indet. South area RB62 ventral osteoderm Goniopholididae indet. South area RB66 caudal vertebra Goniopholididae indet. South area RB69 sacral vertebra Goniopholididae indet. East area RB72 osteoderm Goniopholididae indet. East area RB74-78 vertebrae Goniopholididae indet. South area Testudinata Acronym Material Taxa Area RB5 Shell fragment Testudinata indet. South area RB16 bridge peripheral Testudinata indet. South area RB18 costal Testudinata indet. North area RB20 hypoplastron Testudinata indet. South area RB24 costal Testudinata indet. South area Pterosauria Acronym Material Taxa Area RB19 mandibular fragment Pterodactyloidea indet. South area RB28 Femur Pterodactyloidea indet. South area RB29 humerus Pterodactyloidea indet. South area RB34 epiphyseal fragment Pterodactyloidea indet. South area RB37 Partial radius Pterodactyloidea indet. South area RB55 digit IV phalanx Pterodactyloidea indet. South area 4 Taphonomy than the rock, and showing a very well-preserved exter- nal morphology. However, when broken, the bone tissue No meaningful differences concerning the lithology, abun- appears highly recrystallised, presenting conchoidal frac- dance, and diversity of fossils, or taphonomic patterns have tures, bright white and grey colours, and barely retaining its been observed between the sampled areas. The prelimi- internal structure, making it almost indistinguishable from nary taphonomic analysis here reported, and supported by the encasing rock. field and thin section observations, considers the working Bone remains appear distributed throughout the outcrop, hypothesis that all of them represent a single bonebed (north without an appreciable distribution pattern, and no signifi- and south areas), or successive bonebeds (e.g., east area), cant variation in the concentration of fossils is observed accumulated by similar non-differentiable processes. Thus, along the thickness of the studied section, within its several all the specimens will be analysed as coming from a single bone-bearing levels. Nevertheless, some areas show a rela- locality. tive increase in fossils elements, specifically of fossil fish The fossil taphonomic history of the site is unusual and bones (Fig. 2B-D) and some accumulations of crocodylo- interesting. In the field, the fossils outcrop as dark brown to morph remains (Fig. 6F). Most fossils are bone fragments, black positive reliefs protruding from the limestone, harder but complete disarticulated elements are also frequent, and 13 Journal of Iberian Geology Fig. 3 Thin section of one scale of RB26 Neoginglymodi indet. (A) Fig. 4 A, C, D) RB6 Goniopholididae indet. thin section of the diaphy- polarized transmitted light showing the preserved lamination of the sis of an undetermined long bone. B, D, F) RB52 Goniopholididae histostructure of the scale, and diagenetic fractures infilled with quartz. indet. thin section of a tooth cut at the base of the crown. (A) polar- (B) cross polarized transmitted light image of the same view as (A), ized transmitted light image showing the preserved lamination of the showing that most of the bone has been replaced by quartz (brown histostructure of the cortical and trabecular bone. Note both biominer- and yellow interference colours), whereas only a small amount of the alized tissues preserve the fibrous aspect of the bioapatite, and haver- original fluorapatite is preserved in the bottom part of the scale (denim sian channels can be observed in the trabecular bone. (B) polarized blue interference colours). Note that two different quartz phases can transmitted light image. Dentine of the teeth still shows lamination, be observed in (A) but are indistinguishable in (B), one that is brown whereas enamel is not present. Note fractures caused by diagenetic in (A) and another that is completely transparent suggesting the pres- compression. (C) cross polarized transmitted light image. The general ence of an epitaxial growth of quartz after the bone replacement. White denim blue interference colour of the fluorapatite suggests the origi- arrows point to recrystallized ostracod shells. (C) EDS map showing nal composition of the bone is preserved, except for some relatively the elemental composition of the area in the white square in (A). Flu- small areas were quartz, with a white interference colour, has replaced orapatite in light blue whereas quartz is shown in purple. Dark blue the structure. Large calcite crystals can be observed in the medul- calcite infilled cracks can be observed in the top half of the map. Note lary cavity of the bone, together with indetermined silicate minerals the presence of small greenish pyrite crystal in the bottom part of the of bright interference colours. (D) cross polarized transmitted light image. (D) BSE image of the squared area in 3D. The encasing sedi- image, showing that the bone has been completely recrystallized by ment is a mixture of calcitic micrite and chlorite. Anatomical abbrevia- medium quartz crystals of brownish-yellowish interference colour. tions: Cal., Calcite; Chl., Chlorite; F-Ap., Fluorapatite; Hl., Halite; Py., Drusy infillings are seen filling the fractures, both radially and around Pyrite; Qz., Quartz the teeth. The pulp cavity is filled with a calcite and chlorite matrix, similar to the encasing sediment. (E) BSE image. Note the presence associated remains of crocodylomorphs and articulated of large areas preserving its original fluorapatite composition, inter- remains of Testudinata and Osteichthyes have also been stratified with quartz, which has started the replacing process. Some cracks are filled with Iron oxides. Large calcite crystals with relics of identified. No preferent orientation of long or flat bones has fluorapatite grow in the medullary cavity. (F) BSE image of the teeth, been observed on the field, and there is no apparent classi- completely replaced by quartz. A slightly lighter phase of quartz is fication by size or shape of the specimens. Individual bones filling the cracks, but no significative differences in composition were show various degrees of preservation, from bone splinters detected by EDS. The surface of the teeth, where the enamel should have been, has been replaced by chlorite. Anatomical abbreviations: and fragments, to complete delicate elements such as a Cal., Calcite; Chl., Chlorite; F-Ap., Fluorapatite; Fe-Ox., Iron oxides; pterosaur limb and cranial bones. No evidence of subaerial Hl., Halite; Py., Pyrite; Qz., Quartz exposure or weathering has been observed, including flaking or any presence of crusts or dissolutions. No plastic defor- bioapatite of the bones have been partially (Fig. 3 A, C, D) mations have been identified in any of the specimens, and or totally (Fig. 4B, D, F) replaced by silica. Petrographic fractures caused by bone-to-bone contacts are also absent. thin sections confirm that the SiO2 is actually crystalline The fossil diagenetic history of the osteological elements quartz, which is characterized by white and brown interfer- analysed is of special interest, as all elements are heav- ence colours and small to medium crystals with prismatic ily recrystallized. Concerning its geochemical composi- habits and blocky to undulating extinction, contrasting to the tion, BSE images and EDX analyses show that the original large, denim-blue fluorapatite crystals with fibrous habits. 13 Journal of Iberian Geology Contrary to what it is observed in hand samples, petrographic by a recrystallization of the opal into quartz. This process thin sections show that the internal structure of the bone tis- was also sufficiently slow to allow the survival of the bone sue is relatively well preserved, despite the extensive min- laminations during this replacement. During the peak of eralization. Bone growth lines can be observed, although low-grade metamorphism in the Cameros basin, the temper- are partially distorted. Occasional calcite and pyrite crystals atures may have reached 350–370ºC and pressures of up to have partially replaced the bone tissue (Fig. 3D) and are also 1 Kbar (e.g., Casquet et al., 1992, Barrenechea et al., 2001), present in the calcite matrix. Fractures, probably caused by which probably caused this unusual preservation of the fos- lithostatic compaction of the specimens, are filled with cal- sil bones. Further research is needed to explore the detailed cite, quartz (which may be enriched with clay minerals), and mechanisms of the fossildiagenetic pathways in the context occasionally iron oxides (Fig. 4E). The sediment infilling of low-grade metamorphism. the bone cavities, such as medullary cavities or vasculariza- tion in osteoderms, is mainly micrite, and has abundant neo formed calcite crystals, chlorites, other silicate minerals, 5 Vertebrate assemblage and euhedral pyrites. Some bones (especially those where the replacement by quartz has been completed) may show a In the outcrop, the most abundant remains correspond to chlorite coating. isolated and disarticulated fish bones and scales (Fig. 2C, No significative differences in mineralogical or chemical D), although some partial skeletons of indeterminate ostei- composition have been observed between the bones attrib- chthyans were also collected (Fig. 5). Other vertebrate uted to different taxa, suggesting that there is no taxonomic remains include crocodylomorph isolated teeth, osteo- control of the diagenetic processes. derms, postcranial and a few cranial bones (Fig. 6); Testu- Preliminary taphonomic analysis allows the character- dinata isolated plates and partial shells (Fig. 7); and a few ization of the assemblages as time-averaged, osteichthyan isolated cranial and appendicular pterosaur bones (Fig. 8). dominated, multitaxic bonebeds (especially the south area). Interestingly, not a single dinosaur bone has been identified The apparent lack of evidence of transport, with no pre- in the area so far. ferred orientation or sorting of elements, together with the range of degrees of articulation of the specimens suggest 5.1 Fishes that the assemblage formed by attrition and accumulation of bone fragments, and disarticulated and partially articulated The Late Jurassic to the Early Cretaceous actinopterygian skeletal remains in a lacustrine environment. fish faunas from continental or transitional environments Despite the relatively small sample analysed, preliminary of Europe are commonly characterized by the presence taphonomic and palaeoecological hypothesis about the com- of remains of ginglymodians and halecomorphs, together position of the assemblage can be postulated. The assem- with other groups, such as pycnodontiforms or basal tele- blage is dominated by obligate aquatic taxa (osteichthyan osts (e.g. Forey & Sweetman, 2011; Pouech et al., 2015; fishes), which represent the more articulated specimens. Ruiz-Omeñaca and Bermúdez-Rochas, 2010). The Early They are followed by amphibian organisms (Crocodylomor- Cretaceous of the Cameros Basin is no exception and the pha and Testudinata), which show partial articulation and presence of remains of these groups of fishes have also been some degree of association, suggesting that the environment recorded in the Enciso Group (Bermúdez-Rochas & Poyato- was an extensive lake with a permanent water level, and the Ariza, 2007), with also abundant ginglymodian remains in accumulation was produced far from the coast of the lake. the Urbion Group. This is also supported by the lack of fully terrestrial animals, Fish faunas in the Ribota site are represented by abun- outside of disarticulated elements of flying pterosaurs which dant osteichthyan isolated remains (teeth, ganoid scales and would have been incorporated to the assemblage after fall- bone fragments), and some incomplete articulated or semi- ing to their deaths inside the lake. articulated remains. A preliminary study of these remains The low degree metamorphism present in the Matute points to a low fish faunal diversity with specimens of two Fm has resulted in a very particular mode of preservation major groups of actinopterygians, namely: Halecomorphi of the fossil bones in the Ribota bonebeds. Opalized ver- and Neoginglymodi. tebrate bones are known in the literature (e.g., Pewkliang The Jurassic-Cretaceous boundary osteichthyan fishes et al., 2008), where the microstructure of the fossils is record is scarce in Spain (Poyato-Ariza et al., 1999) and the perfectly preserved at the level of the individual osteons. data that the Ribota site could provide is potentially inter- It seems that an analogue process may have occurred in esting in order to understand the distribution and evolution the Ribota bonebed, which allowed the preservation of of these fish faunas during the development of the Iberian the detailed histostructure of the bone, but it was followed Basin Rift System. 13 Journal of Iberian Geology Fig. 5 Fish remains from the Ribota fossil site. (A) left dentary RB13 of Halecomorphi indet. in lateral view; (B) possible right dentary RB27 of Neoginglymodi indet. in medial view; (C) pos- sible dermopalatine fragment RB58 of Neoginglymodi indet. in ventral view; (D) slab of an articulated partial skeleton RB14 of Neoginglymodi indet. in lat- eral view; (E) articulated partial skeleton RB26 of Neoginglymodi indet. in lateral view; F and G) Isolated scales in the field assigned to Neoginglymodi indet Osteichthyes Huxley, 1880. 5.1.2 Description Actinopterygii Cope, 1887. Neopterygii Regan, 1923. Neoginglymodian fishes’ remains are abundant in the Ribota Holostei Müller, 1844 (sensu Grande, 2010). Site, with the presence of scales, teeth, and bones. Most of Halecomorphi Cope, 1872 (sensu Grande & Bemis, them have been located isolated and scattered all over the 1998). site, but several articulated or semiarticulated remains have Halecomorphi indet. been recovered. Among them, there are some cranial and postcranial bones, at least three mandibular bones with teeth 5.1.1 Description attached, and some sets of scales in connection. Most of the specimens are partially covered by limestone and their Halecomorph fishes are represented by one isolated remain exposed surfaces are damaged by meteorization, so further (RB13) corresponding to a partial left dentary with at least preparation is required in order to study them properly. four incomplete teeth attached (Fig. 5A). The specimen was Two specimens (RB27 and RB58) are bone fragments recovered in the east area (Fig. 2A) and partially exposed in with several teeth in connection (Fig. 5B, C). These speci- lateral view, and as a result, the lateral surface of the den- mens are incomplete and still partially covered by lime- tary is quite damaged and abraded by erosion. Despite this, stone, so it is difficult to identify them. One specimen seems remains of the mandibular sensory canal can be observed. to correspond to a right dentary (RB27, Fig. 5B) and the This canal traverses the bone parallel to its ventral margin, other one could be a dermopalatine (RB58, Fig. 5C). Both with three large pores visible, where the anterior and poste- specimens show some degree of heterodonty, as the inner rior ones are larger than the middle one. The dentary is elon- teeth are more robust and bigger than the marginal ones. In gated, robust and shallow, with a total length of 4.75 cm and the dentary the posterior teeth are about twice the size of the a maximum height of approximately 1 cm at its posterior anterior ones. All teeth are rounded to oval in section and end. Teeth are only present in its anterior section. They are present a conspicuous papilla in the centre of their occlusal slightly conical and well-spaced, but poorly preserved (their surface. This character is also present in the teeth of Cam- tips are missing), without any presence of acrodine caps or erichthys lunae (Bermúdez-Rochas & Poyato-Ariza, 2015). potential carinate terminations. The two anterior teeth are The abundant scales recovered are ganoid type scales, slightly posteriorly curved. approximately rhomboidal in shape (RB14, RB26, Fig. 5D, Holostei Müller, 1844 (sensu Grande, 2010). E, F, G). There is no evidence of surface ornamentation or Ginglymodi Cope, 1872 (sensu Grande, 2010). serration in their posterior borders, but the ganoine surface Neoginglymodi López-Arbarello & Sferco, 2018. of exposed scales is heavily worn, and most of their posterior Neoginglymodi indet. borders are broken. Despite this, a variation in their general morphologies, that corresponds to the different body region that they occupied, can be observed. Based on this morphol- ogy, in the Ribota site there are scales corresponding to all 13 Journal of Iberian Geology the regions of the body. Among the morphologies observed, (Tykoski et al., 2002). The group survived to the present at least one scale corresponds to the dorsal ridge, and seems with the crown group Crocodylia as the clade that includes to bear a short spine (Fig. 5F), as it occurs in the dorsal all extant species and their closest fossil relatives. In terres- spines of C. lunae (Bermúdez-Rochas & Poyato-Ariza, trial to transitional environments from the Middle Jurassic 2015). The so-called peg-and-socket articulation (typical of to the Early Cretaceous of Europe, the most typical crocody- ganoid scales) is not visible in the remains exposed. How- lomorph fossil assemblage is dominated by the neosuchian ever, some specimens show a well-developed rostro-caudal clades Atoposauridae, Bernissartiidae, and Goniopholididae articulation, with the corresponding two anteriorly oriented (e.g., Brinkmann, 1989; Buscalioni et al., 2008; Schwarz- processes (Fig. 5G). Wings et al., 2009; Gasca et al., 2012; Puértolas-Pascual et al., 2015b; Tennant et al., 2016a; Guillaume et al., 2020). 5.1.3 Remarks However, deposits with crocodylomorphs from the Juras- sic–Cretaceous transition are still scarce, giving the Ribota Halecomorph fishes are common in the Jurassic and Cre- site greater interest. At this site, the most abundant reptiles taceous faunas of Spain (i.e., Wenz, 1995; Ruiz-Omeñaca correspond to crocodylomorphs remains (Fig. 6), which et al., 2006; Martín-Abad & Poyato-Ariza, 2017) and, as represent more than half of the collected bones. commented above, remains of this group, assigned to Ami- Crocodylomorpha Hay, 1930 (sensu Walker, 1970). iformes, have been previously cited in the Early Cretaceous Crocodyliformes Hay, 1930. of the Cameros basin (Bermúdez-Rochas & Poyato-Ariza, Mesoeucrocodylia Whetstone & Whybrow, 1983. 2007). Their finding in the Ribota site represents their first Neosuchia Gervais, 1871 (sensu Benton & Clark, 1988). record in the Matute Formation, and their oldest record in Neosuchia indet. the basin. Unfortunately, the preservation condition of the only specimen recovered does not allow a higher taxonomic 5.2.1 Description precision to be achieved at this time. The presence of isolated ganoid scales in the Matute Fm Most of these crocodylomorph remains correspond to iso- was recorded for the first time in the Virgen del Prado site lated vertebrae (Fig. 6A-E), however, some of them were (Inestrillas-Aguilar del Río Alhama, La Rioja, Spain) by recovered associated in an area of less than 1 m2 and pre- Moratalla (1993), and assigned to the genus Lepidotes at the senting similar sizes (Fig. 6F), so they could belong to the time. After the revised classification of the ginglymodian same individual. Since most of the vertebrae are partially fishes made by López-Arbarello (2012), the assessment of eroded and/or contained within the rock matrix, some details these kind of isolated ganoid scales to the genus Lepidotes have been hidden. However, their general morphology, can no longer be considered valid (see Bermúdez-Rochas shape of the vertebral centra, and the position of the ribs, & Poyato-Ariza, 2015). More than twenty years later, the diapophyses and parapophyses (when these are preserved) taxon Camerichthys lunae was erected in the San Andrés have allowed for assigning them to different regions of the de San Pedro Site (Soria), based on an articulated speci- vertebral column. Both in anterior and posterior views, the men with the skull and the anterodorsal portion of the body articular surfaces of the centra are sub-circular and gently preserved (Bermúdez-Rochas & Poyato-Ariza, 2015). The concave due to the presence of a shallow central depres- rostro-caudal articulation present in the Ribota site ganoid sion (amphicoelous condition), and they are almost as dor- scales is known in several ginglymodian genera, such as soventrally tall as they are mediolaterally wide. The ventral Callipurbeckia or Scheenstia, belonging to the orders Semi- and lateral surfaces of the centra are slightly concave, giv- onotiformes and Lepisosteiformes, respectively (see López- ing them the typical hourglass shape. At least one cervical Arbarello & Sferco, 2018). Therefore, although they present vertebra has been recovered (RB57; Fig. 6A). This vertebra similarities with C. lunae, the attribution of these scales is identified by the shape and position of the diapophyses to this taxon cannot be assured, and a higher taxonomic and parapophyses. The presence of parapophyses on the lat- assignment cannot be made until more complete material eral surfaces of the centrum (but in its dorsal half), would is recovered. indicate that it is one of the last posterior cervical vertebrae (e.g., Mook, 1921; Puértolas-Pascual et al., 2015a). The 5.2 Crocodylomorpha most abundant vertebrae correspond to the dorsal region of the trunk (RB44; Fig. 6B), and they have been identified The crocodylomorphs of the Late Jurassic and Early Cre- based on the presence of long transverse processes located taceous of Europe are dominated by the Neosuchia, a clade dorsally in the vertebral centrum and at the same height as of mesoeucrocodylian crocodyliformes, with their oldest the neural arch. A sacral vertebra has also been identified record in the Early Jurassic (Sinemurian) of North America (RB69; Fig. 6C). It has been recognised as a sacral element 13 Journal of Iberian Geology Fig. 6 Crocodylomorph remains from the Ribota fossil site. (A) cervical vertebra RB57 in anterior view; (B) dorsal vertebra RB44 in anteroposterior view; (C) sacral vertebra RB69 in anteroposterior view; (D) caudal vertebra RB21 in lateral view; (E) caudal vertebra RB66 in lateral view; (F) disarticu- lated vertebrae (RB74-78) and osteoderms highlighted in red that could correspond to the same individual; (G) dorsal osteoderm RB61 in dorsal view; (H) ventral osteoderm RB62 in ventral view; (I) ventral osteoderm RB35 in ventral view; (J) isolated tooth RB39; K) disarticulated cranial remains RB17. Anatomical abbreviations: al., alveolus; ce., centrum; cr., caudal rib; de., dentary; di., diapophysis; nc., neural canal; ns., neural spine; p.a., parapophysis; pfm., poste- rior foramen for the mandibular ramus of cranial nerve V; pre., prezygapophysis; tr., transver- sal process; sp., splenial; su., surangular. Unnumbered scale bars = 2 cm due to the presence of a very stout fused rib (also referred to juveniles fuse their caudal vertebrae very early in ontogeny as the transverse process) (Gomes de Souza, 2018) attached (Brochu, 1996; Ikejiri, 2012). to the lateral margin of the vertebral centrum (Mook, 1921). Neosuchia Gervais, 1871 (sensu Benton & Clark, 1988). Whether it belongs to the first or second sacral vertebra Goniopholididae Cope, 1875. cannot be confirmed with certainty since it is incomplete Goniopholididae indet. and partially covered by matrix. At least one caudal ver- tebra (RB21; Fig. 6D) from the anterior or middle region 5.2.2 Description of the tail has been identified. Although it is not complete, this caudal vertebra is characterized by the presence of The second most abundant crocodylomorph elements at an elongated vertebral centrum with a more quadrangular the site are osteoderms (Fig. 6G-I). Although most of them articular surface, and transverse processes (= fused caudal appear isolated and fragmented, making their identification ribs according to Gomes de Souza, 2018) arising from the difficult, some characteristics of the most complete ones dorsal region of the centrum rather than from the neural allow to assign them anatomically and taxonomically. At arch. In addition, the ventral region of this centrum appears least three positional types of osteoderms have been identi- to possess two articulation facets for the chevrons attached fied: dorsal, ventral, and appendicular. One of the largest to two subtle anteroposterior parallel ridges (Mook, 1921). and most complete osteoderms (RB61) has been interpreted Interestingly, some small isolated centra (RB66; Fig. 6E), as belonging to the dorsal region of the trunk. This dorsal (tentatively assigned to caudal vertebrae) were found dis- osteoderm (Fig. 6G) is twice as wide as it is long, with a articulated by the opened neurocentral suture. If these ver- smooth non-ornamented articular facet in the anterior mar- tebrae are confirmed to be caudal, they would indicate the gin, an anterolateral peg, and the lateral margin ventrally presence of juvenile individuals in addition to adults, since deflected with a high angle. These osteoderms are typi- cal of crocodylomorphs with biseriate armour (two dorsal 13 Journal of Iberian Geology rows) and a closed paravertebral bracing system (Salisbury are more informative, since the combination of dorsal osteo- & Frey, 2001; Puértolas-Pascual et al., 2015a; Puértolas- derms that are twice as wide as they are long, with peg and Pascual & Mateus, 2020). At least two ventral osteoderms groove articulation and a ventrally folded lateral margin, (RB62 and RB35) have been identified (Fig. 6H, I). They together with the presence of polygonal ventral osteoderms, are flat, equidimensional (as wide as they are long), polyg- points to the clade Goniopholididae (e.g., Wu et al., 1996; onal with pentagonal to hexagonal contours, with straight Salisbury & Frey, 2001; Puértolas-Pascual & Mateus, 2020). edges, and present the typical ornamentation of big circular Furthermore, the presence of generalist conical teeth is also pits evenly distributed over their ventral surface. Although typical of this clade of neosuchian crocodylomorphs, among they were isolated, their edges present some crenulation, others (Guillaume et al., 2020). Therefore, given the palaeo- indicating their sutures with adjacent osteoderms, which geographic/chronological context, the size of the bones, the would form a rather rigid ventral armour (Wu et al., 1996; morphology of the vertebrae and teeth, and the characteris- Salisbury & Frey, 2001). Some small, flat, elliptical to sub- tic shape of the osteoderms, most of these remains can be circular osteoderms have been interpreted as belonging to provisionally assigned to Goniopholididae indet. the appendicular region. This type of osteoderm can be con- fused with those of the neck region, however the latter usu- 5.3 Testudinata ally have a keel, at least in some goniopholidids and extant crocodylians (Wu et al., 1996). The most abundant and diverse lineage of turtles in the Neosuchia Gervais, 1871 (sensu Benton & Clark, 1988). Upper Jurassic levels of Europe is that of Thalassochely- Neosuchia indet. dia (Anquetin et al., 2017). Several representatives of the group of littoral thalassochelydians Plesiochelyidae have 5.2.3 Description been recognized in the Iberian Peninsula, as well as the freshwater Hylaeochelys, the latter being the only thalas- Among the cranial material recovered at the site, the most sochelydian surviving after the Jurassic–Cretaceous transi- abundant elements are isolated teeth (RB39; Fig. 6J). All tion (see Pérez-García, 2017; and references therein). The the teeth belong to the same generalist morphotype, consist- North American and European lineage of the paracrypto- ing of conical crowns, with smooth mesial and distal cari- diran turtles Pleurosternidae is recognized within both the nae, and ornamentation consisting of parallel longitudinal Late Jurassic and the Early Cretaceous. The Iberian record basiapical striae. Only a group of disarticulated (but associ- of this group of freshwater basal turtles (i.e., members of ated) cranial bones (RB17) have been recovered from the Testudinata not attributable to the crown Testudines) is cur- site (Fig. 6K). Although they are partially covered by rock rently identified as the most diverse in Europe, with sev- matrix, they can be recognized as part of a mandible. A den- eral Iberian representatives having been defined from the tary fragment, probably from the symphyseal region, with Kimmeridgian to the Albian levels (see Pérez-García et al., five circular dental alveoli (two of them preserving tooth 2022; and references therein). Another lineage identified in fragments) has been identified. In addition, these associ- Europe from both Upper Jurassic sites (where its record is ated bones also include an indeterminate cranial element very limited) and throughout the Cretaceous is that of the (probably a surangular fragment) and a splenial fragment in terrestrial stem turtles Helochelydridae. Although the Ibe- which two posterior perforations for the mandibular ramus rian Late Jurassic record includes other lineages of turtles, of cranial nerve V seem to be present. the current limited availability of material does not allow us to know which lineages they belong to. However, a diverse 5.2.4 Remarks fauna of freshwater eucryptodiran turtles is recognized in Europe, and especially in the Spanish record, after the Juras- Although the crocodylomorph material at the Ribota site is sic–Cretaceous transition, probably from the Berriasian, as very fragmentary and most elements are isolated, taking it a result of diachronic dispersals to this continent (see Pérez- as a whole has made it possible to make a series of taxo- García, 2017; and references therein). nomic inferences. Regarding the vertebrae, most of them Testudinata Klein, 1760. belong to individuals of medium to large size. In addition, Testudinata indet. the presence of amphicoelous vertebral centra points to the presence of non-eusuchian taxa. Considering the known 5.3.1 Description taxa in this type of palaeoenvironment, age and palaeobio- geographical region, these vertebrae most probably belong All turtle specimens so far identified in Ribota correspond to neosuchian crocodylomorphs (e.g., Guillaume et al., to shell elements (Fig. 7). They are represented by isolated 2020; Puértolas-Pascual & Mateus, 2020). The osteoderms plates, both from the carapace (Fig. 7A-C) and from the 13 Journal of Iberian Geology Fig. 7 Selection of turtle remains from the Ribota fossil site. A) visceral view of an almost com- plete costal RB18; B) anterior or posterior view of the proximal region of a costal RB24; C) bridge peripheral RB16, in dorsal view; D-D’) partial left hypoplastron RB20, in ventral view. Dotted lines in D’ indicate broken edges; continuous black lines correspond to the margins of the plate; and the border of the scutes are represented by thicker gray lines. Scute abbreviations: ab., abdominal; fe., femoral; inf., inframarginal; m., marginal plastron (Fig. 7D). Although most of these remains are not plates from Ribota cannot be attributed to Pleurosternidae complete, they generally show a well-preserved outer sur- because the taxon identified from there lacks mesoplas- face, with a rough pattern (Fig. 7E). The material currently tra, as well as an ornamental pattern composed by pits or found does not show evidence for the presence of more than tubercles, and by sulci on the margins of the plates (Pérez- one taxon in Ribota. García et al., 2022). The characters described here (i.e., In visceral view, the proximal end of the dorsal ribs is absence of mesoplastra, present of inframarginal scutes), well-developed (RB18 and RB24), being relatively robust are compatible with those of the members of the lineage (Fig. 7A, B). Both the ventro-medial margin of the bridge of Thalassochelydia, very abundant and diverse in the Late peripherals RB16 (Fig. 7C), and the lateral one of the only Jurassic levels of Europe, from the Oxfordian to the Titho- hypoplastron RB20 found (Fig. 6D), indicate that the con- nian (Anquetin at al., 2017; Pérez-García & Ortega, 2022). tact between the carapace and the plastron was at least par- However, most of its representatives (including all mem- tially ligamentous. The hypoplastron corresponded to that bers of Plesiochelyidae, a lineage exclusive to Europe and of a taxon with inframarginals, with at least two of these well-represented in the Iberian record) are recognized as scutes overlapping this plate. The anterior hypoplastral mar- littoral forms, which is not compatible with the sedimen- gin is not perpendicular to the axial plane (that plane being tary environment interpreted for the Matute Alloformation. recognized by the suture of this plate with the other hypo- The only freshwater member of Thalassochelydia currently plastron), but rather it is postero-medially directed. Thus, known is Hylaeochelys which, furthermore, is the only one this taxon lacked mesoplastra between the hyoplastra and that survived until the Early Cretaceous (Pérez-García et al., the hypoplastra. The abdominal-femoral sulcus is latero- 2023). The material analyzed here is not attributable to this posteriorly directed. genus represented in the Iberian Peninsula, considering the absence of a slightly fluted outer surface of the plates. The 5.3.2 Remarks limited availability of characters in the turtle material found at Ribota is also compatible with that of another lineage of As has been indicated in the Introduction, a turtle was freshwater forms recognized in Europe: Xinjiangchelyidae. described in Ágreda. It corresponded to a member of the Although this group has an Asian origin, it reached Europe Pleurosternidae (Paracryptodira), and was attributed to the during the Lower Cretaceous, at the Berriasian or Valangin- new species Pleurosternon moncayensis, exclusively repre- ian. It was a diverse lineage in the European Lower Creta- sented by its holotype (Pérez-García et al., 2022). That turtle ceous record and especially in that of the Iberian Peninsula, was defined by both cranial and shell remains. The turtle where several forms have been identified (see Pérez-García, 13 Journal of Iberian Geology 2017; and references therein). Therefore, although the cur- fragmentary fossil bone material (e.g. Fuentes Vidarte & rent limited information on the Ribota turtles does not allow Meijide Calvo, 1996, 2010; Buffetaut, 1999; Ruiz-Omeñaca us their precise systematic attribution, they could corre- et al., 2004; Vullo et al., 2009). Most known trackway spond to a single form, compatible with Thalassochelydia localities are found in the north and northeast of Spain, in and with Xinjiangchelyidae. The potential future discovery the provinces of Asturias, La Rioja, and Soria, with some of new remains in this fossil locality will be necessary to fossil bones also occurring in the latter two, and addition- perform a more precise systematic attribution. However, the ally in the provinces of Aragón, Valencia and Cuenca (e.g., so far identified remains in Ribota are relevant since they Holgado et al., 2011). Largely these fossils consist of post- allow the identification of two turtle lineages in Ágreda, as cranial remains of isolated bones (mainly wing elements) they are not compatible with Paracryptodira. and teeth, although cranial material has also been recovered, most notably in the holotype fossils of Prejanopterus cur- 5.4 Pterosauria virostris Fuentes Vidarte and Mejide Calvo, 2010 (Pereda- Suberbiola et al., 2012), Europejara olcadesorum (Vullo et The time period of the Jurassic–Cretaceous transition is a al., 2012), and Iberodactylus andreui (Holgado et al., 2019). transformative time for the pterosaur faunal composition, The Ribota pterosaur assemblage is thus far represented where non-pterodactyloids were already in the process of by six isolated elements: preliminary identifications des- global extinction, and the Pterodactyloidea had already ignate them as a humerus (RB29, Fig. 8A), a radius frag- began their diversification, radiating geographically and ment (RB37, Fig. 8B), a femur (RB28, Fig. 8C), a digit IV taxonomically throughout the remainder of the Cretaceous phalanx I (RB55, Fig. 8D), an epiphyseal fragment (RB34, (although the exact early evolutionary drivers behind this Fig. 8E), and a mandibular fragment (RB19, Fig. 8F). transition are still largely undetermined). The specific time Order Pterosauria Owen, 1842 (sensu Andres & Padian, periods of the Tithonian–Berriasian, occurring just after the 2020a). peak of this marked change, were therefore still typified by Suborder Pterodactyloidea Plieninger, 1901 (sensu residual rhamphorhynchids, but also largely occupied by Andres and Padian, 2020b). anurognathids, pterodactylids and ctenochasmatids (Paul, Pterodactyloidea indet. 2022). Today, the known Late Jurassic–Early Cretaceous ptero- 5.4.1 Description saur occurrences in Spain have mainly been represented by trackways (e.g., Lockley et al., 1995, 2008; Wright et al., RB29 is a mostly complete left humerus (Fig. 8A), visible in 1998; Hernández Medrano et al., 2006; Pascual Arribas et ventral view. Based on humerus scaling with P. curvirostris al., 2015; Piñuela, 2015; Pascual Arribas and Hernández- (the closest taxon geographically, and similar morphologi- Medrano, 2016), but also by disparate occurrences of cally) (Pereda-Suberbiola et al., 2012), the Ribota specimen Fig. 8 Pterosaur remains from the Ribota fossil site. (A) humerus RB29 in ventral view; (B) Partial radius RB37 in indeterminate view; (C) Femur RB28 in poste- rior view; (D) a digit IV phalanx RB55 in indeterminate view (note that they are two different pictures superimposed; E) epiph- yseal fragment RB34 in lateral view; F) mandibular fragment RB19 in dorsal view. Anatomical abbreviations: dpc., deltopectoral crest. Scale bars = 2 cm 13 Journal of Iberian Geology RB29 is about 30% smaller and would have achieved about capped by a mushroom-shaped constriction of the neck, and one-meter wingspan. The humerus is severely eroded along well differentiated from the femoral shaft by an 158° angle. both articular ends, and especially in the area of the delto- pectoral crest, which is seemingly placed distal to the ulnar 5.4.3 Remarks crest (a condition of pterodactyloids according to Unwin & Martill, 2018). At the proximal end, the ulnar crest faces Although likely belonging to different individuals, since anterolaterally, with a distinct concavity between it and they were isolated and separated by several meters, it is not the region of the deltopectoral crest. The humeral shaft is possible at the moment to determine if the pterosaur ele- straight, with an ovoid cross-section at its midpoint, and ments from Ribota belong to the same species, and since moderately expanded mediolaterally towards both articular there are no overlapping elements in the assemblage, more ends. sampling would be needed to ensure whether one or more Mandibular fragment RB19 (Fig. 8F) is one of the most than one species is here represented. The bones were identi- informative pterosaur bones in the Ribota assemblage, and fied as belonging to pterosaurs due to the hollowness of the is exposed dorsally and laterally, with sediment obscuring bones and a cortical thinness of between 0.3 and 1.6 mm its ventral surface. The fragment is very slender, with almost (e.g., Bennet, 2003). The preservation of these elements is parallel margins and a very thin midline rectilinear groove. excellent in terms of three-dimensionality with almost no The exact position of the fragment along the length of the taphonomic distortion, however significant exposure to the entire mandible is indeterminate (due to being broken off elements over time has led to extensive erosional damage both ends), but it is likely almost very near the anterior-most or breakage along the bone surfaces, rendering certain fea- end of the dentary. In the mandible fragment, a total of two tures indistinguishable. Thus, some elements are not cur- partial and eight complete alveoli have been preserved. Pos- rently identifiable beyond the level of Pterosauria indet., terior to the partial alveoli (one in each side), four complete such as a distal left radius fragment RB37 (Fig. 8B) (visible and well-developed raised alveoli are visible on each side, in anterior view, and missing large sections of cortical bone directed anterolaterally. The alveoli are all ovoid, elongated throughout the shaft and articular end), right digit IV pha- anteroposteriorly, although they vary in diameter, alternat- lanx I RB55 (Fig. 8D) (visible in ventral view, and although ing between larger and smaller along the tooth row. Each nearly complete, sustained the extensor tendon process pair of alveoli located laterally to each other also present being completely eroded away anterior to the ventral cot- asymmetry in size, with a larger alveolus in one side of the yle), and epiphyseal fragment RB34 (Fig. 8E) (which is to mandible and a smaller one on the opposite side. Further- too highly eroded to glean sufficient morphological infor- more, each alveolus is not aligned at the same anteroposte- mation, precluding any assignation). The remaining better- rior height as its lateral counterpart, giving an extra degree preserved elements, however, are more informative. of asymmetry to the dental arrangement. In transverse sec- The humeral shaft RB29 is straight (as in pterodactyloid tion, the broken ends of the symphysis show a distinctive pterosaurs), unlike the typical curved shaft of basal ptero- rounded-triangle cross-section, compressed laterally, with saurs (Unwin & Martill, 2018). Humerus RB29 is therefore the pointed apex facing ventrally. provisionally assigned to Pterodactyloidea indet. Regarding Suborder Pterodactyloidea Plieninger, 1901 (sensu the femur RB28, the 158° angle of its femoral head with Andres and Padian, 2020b). respect to the shaft, is consistent with the Ornithocheiridae Pteranodontoidea Kellner, 2003. (Unwin, 2003). Also, based on this femoral neck-shaft angle Pteranodontoidea indet. greater than 135°, it also corresponds to the Pteranodontoi- dea of Kellner (2003) in this analysis. 5.4.2 Description As previously stated, the cranial material is more infor- mative. Although it is significantly smaller in size, on first RB28 is a left femur (Fig. 8C) only visible from its posterior glance, RB19 shows distinct morphological similarities face, as it is still embedded in matrix. It is highly eroded with Prejanopterus curvirostris, particularly in the trian- on its articular ends, and in particular in the region of the gular cross-section of the symphyseal area, however the greater trochanter (which has been entirely eroded away) alveoli on P. curvirostris are organized quite differently and in the asymmetrical distal epiphyseal region. The shaft and more symmetrical than in RB19. P. curvirostris was is ovoid in cross-section, relatively straight anteroposte- assigned to the Pterodactylidae (Pereda-Suberbiola et al., riorly, and slightly bowed mediolaterally, also exhibiting 2012), however some authors have implied that it belongs a slight expansion in mediolateral width to the distal end. instead to the Lonchodectidae (Witton, 2003; Paul, 2022), The femoral head is strongly offset and proximally directed, a contentious group. However, the alveolar margins of the Ribota specimen do not appear as pronouncedly raised as 13 Journal of Iberian Geology most lonchodectids (whose alveoli usually are elevated different areas and is becoming a key area to understanding above the medial portion of the occlusal surface), nor is the how those faunal changes occurred. Significant sites have midline groove as pronounced as the sulcus present in most been found in areas such as the Lusitanian Basin in Portugal lonchodectids (Unwin, 2001; Averianov, 2020). Mandibular (e.g., Martin & Krebs, 2000; Malafaia et al., 2010; Mateus fragment RB19 also shows morphological similarities with et al., 2017, and references therein), the Asturian Basin Aetodactylus halli Myers, 2010, which was assigned to the (Piñuela, 2015, and references therein), and several areas Ornithocheiridae (as is Ribota femur RB28), particularly of the Iberian Basin Rift System (mainly South Iberian and in the labiolingually-compressed alveoli, but also differing Maestrazgo basins) in Spain (see Aurell et al., 2016, 2019; in RB19’s more lateral positioning of the alveoli along the Campos-Soto et al., 2017;, 2019, and references therein). It symphysis (a feature more commonly associated with the is also worth noting that adjacent to the Iberian Peninsula, Ctenochasmatidae [Unwin, 2003]). Therefore, mandibu- the Aquitaine Basin has also yielded rich and diverse Titho- lar fragment RB19 is here conservatively assigned to the nian and Berriasian vertebrate assemblages that show strong Pterodactyloidea (the suborder inclusive of all the afore- similarities with Iberian faunas (Buffetaut et al., 1989; mentioned groups), but due to the differences observed with Mazin et al., 2008; Vullo et al., 2014; Allain et al., 2022). these other taxa, could potentially be further representa- Among the Lusitanian localities with abundant osteo- tive of a new taxon. A more detailed study of the specimen logical vertebrate remains, Guimarota (Martin & Krebs, and the possibility of more material being recovered in the 2000) and Andrés (Malafaia et al., 2010) in Leiria (Portu- future are necessary to clarify its phylogenetic position. gal) stand out. These two localities are characterized by a Although the majority of elements in this preliminary highly diverse vertebrate fauna including fishes, lizards, assemblage of pterosaurs have overall been informatively crocodylomorphs, pterosaurs, and dinosaurs among others. limited by their physical preservation, the potential of both There are several Spanish areas with abundant evidence of the further preparation of these specimens, coupled with these groups of vertebrates within this time interval (Kim- future excavation efforts will doubtlessly yield a signifi- meridgian–Berriasian) such as the Asturian Basin (e.g., cant and valuable picture of the pterosaur palaeodiversity Lockley et al., 2008; Piñuela 2015) or the Cameros basin of the Ribota locality, as well as the overall landscape for (Hernández-Medrano et al., 2006; Castanera et al., 2018), the Pterosauria during the transitional Tithonian–Berriasian but their records are mainly tracksites, and bone remains time period worldwide. are comparatively scarce. In the Asturian basin, osteologi- cal remains from different groups of vertebrates have been reported and mainly represented by dinosaurs, pterosaurs, 6 Implications for the palaeobiodiversity in crocodylomorphs, turtles, and fishes (e.g., Ruiz-Omeñaca the Iberian ecosystems during the Jurassic/ and Bermúdez-Rochas, 2010; Ruiz-Omeñaca et al., 2006;, Cretaceous boundary 2012; Rauhut et al., 2018). In the eastern Cameros basin, dinosaurs, pterosaurs, crocodylomorphs, and turtles are rep- The Jurassic–Cretaceous transition is a significant geo- resented in the ichnological record (Hernández-Medrano logical period, and considerable faunal and environmental et al., 2006; Castanera et al., 2018), and osteological fos- changes occurred across the Tithonian–Berriasian transition sils are also represented by isolated remains from different (see Tennant et al., 2016b, 2017; Allain et al., 2022). Instead sites and from different vertebrate groups, such as fishes of representing a significant faunal turnover between the (Pascual-Arribas et al., 2007; Bermúdez-Rochas & Poyato- Tithonian and the Berriasian, these faunal changes are more Ariza, 2015), turtles (Pérez-García et al., 2022), dinosaurs related with environment shifts associated to a sea level (Canudo et al., 2010), and pterosaurs (Fuentes Vidarte & regression at the end of the Jurassic (Allain et al., 2022). Meijide Calvo, 1996. In the western Cameros basin and in Among the groups described in the present paper, Tennant the closely located Torrelapaja subasin, new sites with dif- et al. (2016b) indicate that marine osteichthyans and shal- ferent identified taxa (dinosaurs and crocodylomorphs) have low marine turtles show a considerable decrease across been recently discovered, but their study is currently under the boundary (see also Pérez-García, 2017 and references way (Torcida Fernández-Baldor et al., 2020; Aurell et al., therein) whereas freshwater and terrestrial turtles, and gonio- 2021). pholidid crocodylomorphs, seem to be unaffected. Ptero- Thus, the Ribota assemblage with a total minimum num- dactyloids increasingly flourished right after the boundary ber of 5 taxa (Halecomorphi indet., Neoginglymodi indet., (Butler et al., 2013; Tennant et al., 2016b). In this regard, Goniopholididae indet., Testudinata indet. and Pterodacty- the Iberian Peninsula has yielded many vertebrate localities loidea indet.) represents one of the highest macrovertebrate from the end of the Jurassic (Kimmeridgian–Tithonian) and accumulations in the eastern Cameros basin, providing new Jurassic–Cretaceous transition (Tithonian–Berriasian) in data regarding vertebrate diversity in lacustrine ecosystems 13 Journal of Iberian Geology during the Jurassic–Cretaceous transition of the Iberian Basin. The fossil diagenesis of the deposit is interesting Basin Rift System (IBRS). In this regard, the southern and particular, since the bones have suffered considerable basins of the IBRS (Maestrazgo and South Iberian basins) taphonomic modifications in which the original structure have yielded many of the important vertebrate Spanish has been replaced by quartz. The assemblage is formed localities (with multiple sites) from the Jurassic–Cretaceous by attrition in a lacustrine environment, and dominated by transition (e.g., Ruiz-Omeñaca et al., 2004; Royo-Torres fragmented bones, but with complete bones and associated et al., 2009; Suñer & Martín, 2009; Cobos et al., 2020), and articulated specimens (mainly fishes and partial turtle but these localities are mainly known for their dinosaur shells) also present. The identified vertebrate assemblage is remains, whereas other groups of vertebrates are compara- composed of fishes (Halecomorphi indet. and Neogingly- tively scarce or not studied in detail. It is also noteworthy modi indet.), crocodylomorphs (Goniopholididae indet.), that the Kimmeridgian–Berriasian deposits of the IBRS turtles (Testudinata indet.) and pterosaurs (Pterodactyloidea have yielded a low vertebrate diversity (see supplementary indet.). Counting with the turtle Pleurosternon moncay- material in Aurell et al., 2016; Campos-Soto et al., 2017, ensis, described in the same formation and a few hundred 2019) when compared with other Hauterivian–Barremian meters northwest of Ribota, there is a total of 6 minimum localities, especially those where microvertebrate remains taxa identified within the Matute Fm. The study of the verte- are abundant (Ruiz-Omeñaca et al., 2004; Canudo et al., brate record of the Matute Fm has been neglected for years, 2010). Accordingly, the Ribota assemblage in particular, considering the abundant vertebrate remains that have been and the Matute Formation in general, open a new window previously mentioned (Gómez-Fernández and Melendez, towards the understanding of the vertebrate fauna during the 1994). Therefore, the potential of the Ribota locality in par- latest Tithonian–earliest Berriasian of the Iberian Peninsula, ticular, and the geological unit in general (e.g., Bermudez- as it shows a high concentration of macrovertebrates and Rochas and Poyato-Ariza, 2014; Pérez-García et al., 2022), a moderate number of identified taxa (6 with Pleuroster- makes these deposits key towards the understanding of the non moncayensis, Pérez-García et al., 2022). Although the vertebrate ecosystems in the Jurassic–Cretaceous transi- number of taxa is not higher than other coeval sites on the tion of the Iberian Basin Rift System (especially because IBRS, the absence of more taxa could be produced by the this studied assemblage is palaeoenvironmentally different preliminary nature of this work, since only some of the best from the known coeval sites described within the IBRS). fossil remains recovered from the site were selected for the Nonetheless, this is just a preliminary study and a first step study. It is quite likely that additional field work, labora- towards the understanding of how the fossils have been tory preparation, and further detailed studies of the material preserved, how many bonebeds exist in the area, and how from Ribota site may increase the number of identified taxa. the vertebrate diversity was around the lake (for which the In addition, the Ribota site is a key site to providing new number of taxa will probably increase with further studies). data about the assemblages preserved in the fully continen- Therefore, the preparation of the material already collected tal realm (see palaeobiogeographical reconstructions of the and/or new recovery may provide a considerable increase in basin in Aurell et al., 2021) since the deposits of the Matute the information about these lacustrine assemblages. Formation in the area represent the deposition in a shallow carbonate lake with intense evaporation, and far from the Acknowledgements E.P-P was supported by a postdoctoral con- tract (María Zambrano) funded by the Ministry of Universities of marine influence (Gómez-Fernández and Melendez, 1994). the Government of Spain through the Next Generation EU funds This palaeoenvironment is compatible with the fauna iden- of the European Union. D.C has been supported by the Beatriu de tified so far, where the assemblage is dominated by obli- Pinós postdoctoral programme (BP2017-00195) of the Government gate aquatic taxa (Neoginglymodi indet. and Halecomorphi of Catalonia’s Secretariat for Universities and Research of the Min- istry of Economy and Knowledge. D.D.B-R was supported by the indet.), amphibian organisms (Testudinata indet. and Gonio- project PGC2018-094034-B-C21 of the Spanish Ministry of Science pholididae indet.) and flying pterosaurs (Pterodactyloidea and Innovation. M.M-A is supported by the MCIN/AEI/https://doi. indet.) This palaeoenvironment is also possibly the reason org/10.13039/501100011033 and co-financed by the NextGeneration why the assemblage is different to many of the mentioned EU/PRTR, Ramón y Cajal contract RYC2021-034473, Fundação para a Ciência e a Tecnologia (grant number PTDC/CTA-PAL/31656/2017 sites in the IBRS that are preserved in continental to transi- and research unit UIDB/04035/2020; GeoBioTec); Fieldwork has tional units and dominated by dinosaurs. been funded thanks to and ERG Grant of the European Association of Vertebrate Palaeontologists (EAVP). The study was subsidized in part by Project PID2021-122612OB- I00 of the Spanish Ministerio de Economía y Competitividad-ERDF, as well as by the Aragón Regional 7 Conclusions Government (Grupos de referencia Aragosaurus: Recursos Geológicos y Paleoambientales and E04_20R_FOCONTUR) and Unidad de Dino- The Ribota site represents a new locality that shows the saurios de Teruel financed by the Ministerio de Ciencia e Innovación highest bone concentration of macrovertebrate remains of (Gobierno de España). Authors would like to acknowledge the use of Servicio General de Apoyo a la Investigación-SAI, Universidad de the Tithonian–Berriasian sites within the eastern Cameros 13 Journal of Iberian Geology Zaragoza. Thanks to Marian Arlegui Sánchez, director of the Museo evolution of the Galve sub-basin (Spain) in the middle Tithonian– Numantino, provincial de Soria (Spain). 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Fernandes3,4 · A. Galobart5 · M. Moreno-Azanza1,6 · A. Pérez-García7 · D. Castanera8 2 E. Puértolas-Pascual Área de Didáctica de las Ciencias Experimentales, puertolas@unizar.es Departamento de Didácticas Específicas, Facultad de Formación de Profesorado y Educación, Universidad M. Aurell Autónoma de Madrid, c/ Francisco Tomás y Valiente 3, maurell@unizar.es 28049 Cantoblanco, Madrid, Spain D. D. Bermúdez-Rochas 3 Bayerische Staatssammlung für Paläontologie und Geologie, david.bermudez@uam.es SNSB, Richard-Wagner-Str. 10, 80333 Munich, Germany J. I. Canudo 4 Department of Earth and Environmental Sciences, Ludwig- jicanudo@unizar.es Maximilians-Universität, Richard-Wagner-Str. 10, A. E. Fernandes 80333 Munich, Germany fernandes@snsb.de 5 Institut Català de Paleontologia Miquel Crusafont, A. Galobart Universitat Autònoma de Barcelona, c/ Escola Industrial 23, angel.galobart@icp.cat 08201 Sabadell, Barcelona, Spain 6 M. Moreno-Azanza Department of Earth Sciences, GEOBIOTEC, NOVA School mmazanza@unizar.es of Science and Technology, Campus de Caparica, P- 2829 516 Caparica, Portugal A. Pérez-García 7 a.perez.garcia@ccia.uned.es Grupo de Biología Evolutiva, Facultad de Ciencias, UNED, Avda. Esparta s/n, 28232 Las Rozas, Madrid, Spain D. Castanera 8 castanera@fundaciondinopolis.org Fundación Conjunto Paleontológico de Teruel-Dinópolis/ Museo Aragonés de Paleontología, Avenida de Sagunto s/n, 1 Departamento de Ciencias de la Tierra, Facultad de Ciencias, 44002 Teruel, Spain Aragosaurus-IUCA Reconstrucciones Paleoambientales, Universidad de Zaragoza, Calle Pedro Cerbuna 12, 50009 Zaragoza, Spain 13