Paulo M Brito
email: pbritopaleo@yahoo.com.br

Romain Vullo
email: romain.vullo@univ-rennes.fr

A new species of Pleuropholis (Teleostei: Pleuropholidae) from the Upper Cretaceous Akrabou Formation of southeastern Morocco, with comments on the evolutionary history of the genus

Keywords: Actinopterygii, Neopterygii, Pleuropholid fish, New taxon, Cenomanian, Africa

A pleuropholid fish is described for the first time from the Upper Cretaceous (Cenomanian) Akrabou Formation, Agoult locality, southeastern Morocco. The new material, consisting of a single complete specimen, is attributed to a new species, Pleuropholis danielae, and is diagnosed by its slender body shape, the absence of a leptolepid notch in the dentary, a broad maxilla, covering almost the entirely lower jaw, and flank scales with an unserrated posterior margin. This species represents the last known occurrence of Pleuropholidae, a group so far unknown from the Late Cretaceous.

Introduction

Pleuropholidae is an extinct family erected by [START_REF] Saint-Seine | Les poissons des calcaires lithographiques de Cerin (Ain)[END_REF], with a chronostratigraphic range of Middle Jurassic to Lower Cretaceous [START_REF] Olive | Pleuropholis germinalis n. sp., a new Pleuropholidae (Neopterygii, Teleostei) from the Early Cretaceous of Bernissart, Belgium[END_REF]. Known from marine and continental strata in Europe (e.g. [START_REF] Egerton | British fossils -Pleuropholis. Figures and Descriptions Illustrative of British Organic Remains[END_REF][START_REF] Sauvage | Note sur le genre Pleuropholis[END_REF]Woodward, 1919;[START_REF] Biese | Uber einige Pholidophoriden aus den lithographischen Schiefern Bayerns[END_REF][START_REF] Bravi | Contributo allo studio del giacimento ad ittioliti di Pietraroja (Benevento). I. Pleuropholis decastroi n. sp. (Pisces, Actinopterygii, Pholidophoriformes)[END_REF][START_REF] Lambers | The actinopterygian fish fauna of the Late Kimmeridgian and Early Tithonian 'Plattenkalke' near Solnhofen (Bavaria, Germany): state of the art[END_REF][START_REF] Bonde | An Early Cretaceous (Ryazanian) fauna of "Purbeck-Wealden type" at Robbedale, Bornholm, Denmark[END_REF][START_REF] Bravi | Middle Jurassic Monte Fallano Plattenkalk (Campania, southern Italy): first report on terrestrial plants, decapod crustaceans and fishes[END_REF][START_REF] Schultze | Knochenfische im weiteren Sinne (Osteichthyes oder Osteognathostomata)[END_REF][START_REF] Poyato-Ariza | Osteichthyan fishes[END_REF][START_REF] Olive | Pleuropholis germinalis n. sp., a new Pleuropholidae (Neopterygii, Teleostei) from the Early Cretaceous of Bernissart, Belgium[END_REF], North America (Alvarado-Ortega and [START_REF] Alvarado-Ortega | A Jurassic pleuropholid fish (Teleostei, Pleuropholidae) in the Tethys Sea domain of North America[END_REF], South America [START_REF] Brito | A new pleuropholid, Gondwanapleuropholis longimaxillaris n. g., n. sp. (Actinopterygii: Teleostei) from the Jurassic of north east Brazil[END_REF][START_REF] Chiappe | Biotic association and palaeoenvironmental reconstruction of the "Loma del Pterodaustro" fossil site (Early Cretaceous, Argentina)[END_REF][START_REF] Giordiano | Zurupleuropholis gen. nov. (Teleostei, Albian, Argentina), first pleuropholids from the Cretaceous of South America[END_REF], Africa [START_REF] Saint-Seine | Poissons fossiles de l'étage de Stanleyville (Congo Belge) -Première partie : La faune des argilites et schistes bitumineux[END_REF] and the Middle East [START_REF] Janensch | Fische aus dem Dysodil des Wealden vom Libanon[END_REF][START_REF] Basse | Fossiles du Jurassique supérieur et des "Grès de Nubie" de la région de Sana (Yémen)[END_REF][START_REF] El Hajj | Dysodiles from the lower Barremian of Lebanon: Insights on the fossil assemblages and the depositional environment reconstruction[END_REF], pleuropholids are currently accepted as basal teleosts [START_REF] Patterson | Interrelationships of holosteans[END_REF][START_REF] Patterson | The contribution of paleontology to teleostean phylogeny[END_REF][START_REF] Pinna | Teleostean monophyly[END_REF] although their phylogenetic relationships within Teleostei are still unclear [START_REF] López-Arbarello | Jurassic fishes of Gondwana[END_REF]. Today, pleuropholids are represented by five genera (i.e. Pleuropholis [START_REF] Egerton | British fossils -Pleuropholis. Figures and Descriptions Illustrative of British Organic Remains[END_REF], Parapleuropholis Saint-Seine, 1955, Austropleuropholis Saint-Seine, 1955, Gondwanapleuropholis Brito and Gallo, 2002, and Zurupleuropholis Giordano et al., 2018) and about 24 nominal species comprising relatively small (usually < 10 cm in total length), slender fishes, readily recognized by their small and upturned mouth, the numerous supra and infraorbital bones, and the presence of a single row of deep (i.e. dorsoventrally elongated) flank scales covering almost the whole side of the body [START_REF] Brito | A new pleuropholid, Gondwanapleuropholis longimaxillaris n. g., n. sp. (Actinopterygii: Teleostei) from the Jurassic of north east Brazil[END_REF][START_REF] López-Arbarello | Jurassic fishes of Gondwana[END_REF][START_REF] Arratia | Morphology, taxonomy, and phylogeny of Triassic pholidophorid fishes (Actinopterygii, Teleostei)[END_REF].

In the present paper we describe a new species of Pleuropholis from the Akrabou Formation, which is clearly distinguishable from all the other known species. In addition to the inherent importance of this discovery, the presence of this taxon in the Upper Cretaceous of Morocco increases the distribution and diversity of pleuropholids and represents the youngest member of this family.

The specimen described here comes from the Plattenkalk Konservat-Lagerstätte of Agoult, located at the top of the Gara Sbâa hill, in the Kem Kem region of southeastern Morocco (Cavin et al., 2010: fig. 2;[START_REF] Martill | A new Plattenkalk Konservat Lagerstätte in the Upper Cretaceous of Gara Sbaa, southeastern Morocco[END_REF] fig. 1). These transgressive laminated limestones (Gara Sbâa Member) correspond to the base of the Akrabou Formation and assigned to the upper Cenomanian-?lower Turonian [START_REF] Cavin | Vertebrate assemblages from the early Late Cretaceous of southeastern Morocco: an overview[END_REF][START_REF] Martill | A new Plattenkalk Konservat Lagerstätte in the Upper Cretaceous of Gara Sbaa, southeastern Morocco[END_REF]. This site has mainly yielded a rich and diverse actinopterygian assemblage [START_REF] Cavin | A new Cenomanian ichthyofauna from southeastern Morocco and its relationships with other early Late Cretaceous Moroccan faunas[END_REF][START_REF] Murray | A new Late Cretaceous macrosemiid fish (Neopterygii, Halecostomi) from Morocco, with temporal and geographical range extensions for the family[END_REF], 2011[START_REF] Arratia | Morphology, taxonomy, and phylogeny of Triassic pholidophorid fishes (Actinopterygii, Teleostei)[END_REF], 2014;Murray et al., 2013;Taverne and Capasso, 2021); the assemblage also includes ichnofossils, plants, worms, arthropods, echinoderms, elasmobranchs, and an undescribed lizard specimen [START_REF] Garassino | New decapod assemblage from the Upper Cretaceous (Cenomanian-Turonian) of Gara Sbaa, southeastern Morocco[END_REF][START_REF] Guinot | Marocarcinidae, a new eubrachyuran family, and Marocarcinus pasinii n. gen., n. sp. from the Upper Cretaceous (Cenomanian-Turonian) of Gara Sbaa, southeastern Morocco (Crustacea, Decapoda, Brachyura)[END_REF][START_REF] Martill | A new Plattenkalk Konservat Lagerstätte in the Upper Cretaceous of Gara Sbaa, southeastern Morocco[END_REF][START_REF] Engel | A new Cenomanian-Turonian (Late Cretaceous) insect assemblage from southeastern Morocco[END_REF][START_REF] Krassilov | New Cenomanian florule and a leaf mine from southeastern Morocco: palaeoecological and climatological inferences[END_REF][START_REF] Vullo | The first articulated specimen of the Cretaceous mackerel shark Haimirichia amonensis gen. nov. (Haimirichiidae fam. nov.) reveals a novel ecomorphological adaptation within the Lamniformes (Elasmobranchii)[END_REF][START_REF] Lamsdell | A new limulid (Chelicerata, Xiphosurida) from the Late Cretaceous (Cenomanian-Turonian) of Gara Sbaa, southeast Morocco[END_REF]. The Agoult fossil assemblage, consisting of both marine and terrestrial organisms, indicates that these laminated carbonates were deposited in a shallow marine coastal environment, most likely a quiet muddy lagoon [START_REF] Garassino | New decapod assemblage from the Upper Cretaceous (Cenomanian-Turonian) of Gara Sbaa, southeastern Morocco[END_REF][START_REF] Cavin | Vertebrate assemblages from the early Late Cretaceous of southeastern Morocco: an overview[END_REF][START_REF] Martill | A new Plattenkalk Konservat Lagerstätte in the Upper Cretaceous of Gara Sbaa, southeastern Morocco[END_REF][START_REF] Ibrahim | Geology and paleontology of the Upper Cretaceous Kem Kem Group of eastern Morocco[END_REF].

Material and methods

The specimen is cataloged in the collections of the Muséum d'Histoire naturelle de Marrakech, Morocco (MHNM), but is currently housed at the Muséum national d'Histoire naturelle of Paris, France (MNHN), for study, within an agreement between both museums. Photographs were taken using ultraviolet light, following the technique proposed by [START_REF] Tischlinger | Ultraviolet light as a tool for investigating Mesozoic fishes, with a focus on the ichthyofauna of the Solnhofen archipelago[END_REF].

General anatomical terminology follows [START_REF] Brito | Révision des Aspidorhynchidae (Pisces, Actinopterygii) du Mésozoïque : ostéologie, relations phylogénétiques, données environnementales et biogéographiques[END_REF]. For the anatomy of the caudal region, we used the terminology proposed by [START_REF] Arratia | Actinopterygian postcranial skeleton with special reference to the diversity of fin ray elements, and the problem of identifying homologies[END_REF].

The publication and the newly described species are registered in Zoobank. The LSID (Life Science Identifier) for the publication is: urn:lsid:zoobank.org:pub:93E130B3-AFBE-480F-BA42-9EA726DDD0E8; the LSID for the new species is: urn:lsid:zoobank.org:act:21F4D5D2-06E9-4846-8504-F43B230DABC6.

Results

J

Figs. 1-5

Diagnosis. Large-sized pleuropholid differing from all other species of the genus by its slenderer body shape, the absence of a leptolepid notch in the dentary, a broad maxilla, covering almost the entirely lower jaw, and flank scales with an unserrated posterior margin.

Holotype. MHNM.KK.393, complete flattened specimen preserved in left lateral view.

Etymology. The specific epithet is dedicated to Daniela Vullo, the mother of one of us (R.V.), who passed away in May 2022, for her constant support.

Type-locality and horizon. The specimen comes from outcrops of the Gara Sbaa hill, near the locality of Agoult, in the Kem Kem region of southeastern Morocco. Gara Sbaa Member of the Akrabou Formation, Upper Cretaceous (Cenomanian).

Description

This is a large-sized Pleuropholis (115 mm in standard length and about 130 mm in total length) and by far one of the largest known pleuropholid fishes. The body is very elongated and slender (standard length/body height = 7.7) (Fig. 1). The deepest part of the body is constant The between the pectoral and the anal fin (body height = 15 mm). The head, including the opercular series, measures approximately 1/6 of the standard length. The dermal bones and scales are covered with ganoin.

The description of some skull elements is difficult due to poor preservation. Most of the skull bones are recognizable, but in some cases, some elements are damaged. This situation makes it difficult to interpret the shape of several bones, especially in the upper jaw.

Only a small fragment of the rostral is preserved and both its shape and suture with the posterior bones are not clearly discernible. The nasal is longer than wide, with its anterior half slightly thinner than the posterior one. The nasal overlaps the frontal posteriorly, and contacts the antorbital antero-laterally. The frontal lies just behind the nasal, and is the longest element of the dermal skull, having a length approximately three times that of the nasal. It extends posteriorly to the parietal and dermopterotic. The parietal is a well-developed bone, slightly longer than wide and having approximately half of the length of the frontal. The lateral border of the parietal is almost linear and contacts ventrally the dermopterotic. The dermopterotic is a subrectangular bone, placed lateral to the parietal and posterior to the frontal. This bone is directed backwards, narrowing at its most posterior part. The extrascapular is much smaller than the parietal. The occipital sensory canal passes through the mid-length of the extrascapulars. Anteriorly, the extrascapular partly overlaps the posterior margins of the parietal and the dermopterotic; posteriorly, it overlaps the anterior margin of the posttemporal, and laterally, it is in contact with the dorsal part of the opercle.

The orbital series is formed by one antorbital, the lacrimal, four infraorbitals, the dermosphenotic, and at least two supraorbitals (Figs. 2,3). The antorbital is a subtriangular bone; this bone is positioned above the lacrimal. The lacrimal is an elongated bone with the infraorbital canal extending into it. Both antorbital and lacrimal form the anterior margin of the orbit. The first two infraorbitals border the orbit ventrally, followed by two covering the postorbital region of the orbit. The infraorbital canal extends in the middle part of the bones. The posterior infraorbitals border the anterior edge of the vertical limb of the preopercle. Although not very well preserved, As a characteristic of the genus, the jaws are short, upturned, and positioned ahead of the orbit. The maxilla is poorly preserved with only few fragments present. However, these fragments point to an extremely broadened bone that covers almost entirely the lower jaws. It has a straight dorsal edge. The supramaxilla is not preserved.

The lower jaw seems to be somewhat triangular; the dentary and the angular bones are exposed laterally (Figs. 2,3). The dentary is a short stout subtriangular bone that presents a rounded symphysis. The mandibular sensory canal extends along the ventral border of the bone. Contrary to the pattern found in Pleuropholis decastroi Bravi, 1988, from ,4). The dorsal fin presents two short, unsegmented procurrent ray followed by seven principal rays. The first ray is segmented, longer and thicker than the preceding procurrent rays.The longest ray of the dorsal fin is the second principal ray, which is segmented and branched. Elongated fringing fulcra are placed in the anterior margin of the first and the second principal rays.

The anal fin is larger than the dorsal fin. Procurrent rays are not preserved in the specimen that has about 13 branched principal rays. The first and the second rays seem to have the same length. The main leading ray is preceded by numerous fringing fulcra. After the third ray, they decrease in size and width.

The vertebrae as well as the caudal endoskeleton are completely hidden by the scales and thus not observable in the studied specimen. The caudal fin is almost completely preserved, although some few elements are broken.

The caudal fin has a forked outline. As in other species of the genus (e.g. Egerton 1858: pl. 7, fig. 4;Woodward, 1919: text-fig. 36;Taverne and Capasso, 2019: fig. 8), the upper lobe is slightly longer than the lower lobe (Figs. 1,5). There are about fourteen epaxial basal fulcra, a series of elongated, small, epaxial fringing fulcra, 15 to 16 principal rays, a series of hypaxial fringing fulcra, segmented procurrent rays, and two hypaxial basal fulcra.

The epaxial basal fulcra are elongated, unsegmented leaf-like elements, partially covering the next fulcra. The epaxial fringing fulcra are associated with the first and second principal rays.

The first principal ray is shorter than the following segmented-and-branched principal ray. Both, together form the dorsal leading margin of the fin.

In the lower lobe, the hypaxial procurrent rays are segmented with their terminal segment resembling the fringing fulcra; fringing fulcra extend along the ventral margin of the last principal rays. As in the upper lobe, the last principal ray is shorter than the following segmented-andbranched principal ray (Fig. 5). segmented There are at least 39 lateral rows of thick, ganoid-type scales, from the supracleithrum to the base of the caudal fin (Fig. 5). These scales have different shapes according to the region of the body. The flank scales are deeper than long, progressively decreasing in size posteriorly and becoming diamond-shaped in the caudal region. They present no serrations in the posterior edge. Dorsal to the flank scales there are two rows of small scales. Ventral to the flank scales, there are three rows of small, rectangular scales.

Discussion

Pleuropholis danielae sp. nov. possesses synapomorphies of the family Pleuropholidae, including: the presence of a single row of deepened ganoid scales covering almost the whole side of the body, an edentulous mouth, and a complete orbital series formed by infraorbital and supraorbital bones (cf. [START_REF] Saint-Seine | Les poissons des calcaires lithographiques de Cerin (Ain)[END_REF][START_REF] Brito | A new pleuropholid, Gondwanapleuropholis longimaxillaris n. g., n. sp. (Actinopterygii: Teleostei) from the Jurassic of north east Brazil[END_REF][START_REF] Olive | Pleuropholis germinalis n. sp., a new Pleuropholidae (Neopterygii, Teleostei) from the Early Cretaceous of Bernissart, Belgium[END_REF].

The presence of an upturned maxilla and a jaw articulation lying anterior to the orbit (cf. Woodward, 1919;[START_REF] Saint-Seine | Les poissons des calcaires lithographiques de Cerin (Ain)[END_REF][START_REF] Patterson | Interrelationships of holosteans[END_REF][START_REF] Bravi | Contributo allo studio del giacimento ad ittioliti di Pietraroja (Benevento). I. Pleuropholis decastroi n. sp. (Pisces, Actinopterygii, Pholidophoriformes)[END_REF][START_REF] Alvarado-Ortega | A Jurassic pleuropholid fish (Teleostei, Pleuropholidae) in the Tethys Sea domain of North America[END_REF][START_REF] Taverne | New data on Pleuropholis decastroi (Teleostei, Pleuropholidae), a "pholidophoriform" fish from the Lower Cretaceous of the Eurafrican Mesogea[END_REF][START_REF] Olive | Pleuropholis germinalis n. sp., a new Pleuropholidae (Neopterygii, Teleostei) from the Early Cretaceous of Bernissart, Belgium[END_REF] places the new taxon in the genus

Pleuropholis.

One of the biggest problems concerning the genus Pleuropholis is the validity of most of its contained species, especially those from the Jurassic. Therefore, a comprehensive revision of all these taxa would be necessary, especially because many of them lack any autapomorphies to sustain their taxonomic status, or they are based on body proportions from imperfect or distorted specimens (e.g. [START_REF] Patterson | Interrelationships of holosteans[END_REF][START_REF] Alvarado-Ortega | A Jurassic pleuropholid fish (Teleostei, Pleuropholidae) in the Tethys Sea domain of North America[END_REF][START_REF] Taverne | New data on Pleuropholis decastroi (Teleostei, Pleuropholidae), a "pholidophoriform" fish from the Lower Cretaceous of the Eurafrican Mesogea[END_REF][START_REF] Olive | Pleuropholis germinalis n. sp., a new Pleuropholidae (Neopterygii, Teleostei) from the Early Cretaceous of Bernissart, Belgium[END_REF]. J o u r n a l P r e -p r o o f problems from the same locality, that needs taxonomic revision and does not appear to belong to the genus Pleuropholis), P. decastroi Bravi, 1988, andP. germinalis Olive, Taverne and[START_REF] Olive | Pleuropholis germinalis n. sp., a new Pleuropholidae (Neopterygii, Teleostei) from the Early Cretaceous of Bernissart, Belgium[END_REF] Although not well preserved, the maxilla of P. danielae sp. nov. seems to be broad, almost entirely covering the lower jaw, a condition also known in P. thiollieri Sauvage, 1883, P. decastroi and P. cisnerosorum Alvarado-Ortega and Brito, 2016. Finally, flank scales with an unserrated posterior margin are known in P. germinalis, P. longicauda [START_REF] Egerton | British fossils -Pleuropholis. Figures and Descriptions Illustrative of British Organic Remains[END_REF], P. formosa Woodward, 1919, P. attenuata [START_REF] Egerton | British fossils -Pleuropholis. Figures and Descriptions Illustrative of British Organic Remains[END_REF]P. crassicauda Egerton, 1858. From a palaeobiogeographical point of view, the evolution and distribution of Pleuropholis appear to have been centered in the Tethyan realm (Fig. 6). So far, the only unambiguous Middle Jurassic occurrence of Pleuropholis is from the Bajocian-Bathonian of southern Italy [START_REF] Bravi | Middle Jurassic Monte Fallano Plattenkalk (Campania, southern Italy): first report on terrestrial plants, decapod crustaceans and fishes[END_REF]; this single occurrence, representing the oldest known record of Pleuropholis, is indicative of a low taxic diversity at that time and probably reflects a restricted distribution at the beginning of the evolutionary history of the lineage (neoendemism; see [START_REF] Cowman | The biogeography of tropical reef fishes: endemism and provinciality through time[END_REF]. The genus Pleuropholis reached its apparent highest diversity during the Late Jurassic-earliest Cretaceous (an observation possibly partly due to a preservational bias; see [START_REF] Cavin | The Late Jurassic ray-finned fish peak of diversity: biological radiation or preservational bias?[END_REF], resulting in several European species described from Cerin and the Meuse department in France [START_REF] Sauvage | Note sur le genre Pleuropholis[END_REF][START_REF] Saint-Seine | Les poissons des calcaires lithographiques de Cerin (Ain)[END_REF], Bavaria in Germany [START_REF] Egerton | British fossils -Pleuropholis. Figures and Descriptions Illustrative of British Organic Remains[END_REF][START_REF] Biese | Uber einige Pholidophoriden aus den lithographischen Schiefern Bayerns[END_REF], and the English Purbeck and coeval beds in Denmark [START_REF] Egerton | British fossils -Pleuropholis. Figures and Descriptions Illustrative of British Organic Remains[END_REF]Woodward, 1919;[START_REF] Bonde | An Early Cretaceous (Ryazanian) fauna of "Purbeck-Wealden type" at Robbedale, Bornholm, Denmark[END_REF]. Two additional species, P. cisnerosorum from the Kimmeridgian of southern Mexico (Alvarado-Ortega and [START_REF] Alvarado-Ortega | A Jurassic pleuropholid fish (Teleostei, Pleuropholidae) in the Tethys Sea domain of North America[END_REF] and P. bassei Lehman in [START_REF] Basse | Fossiles du Jurassique supérieur et des "Grès de Nubie" de la région de Sana (Yémen)[END_REF] from the Tithonian-Berriasian of Yemen [START_REF] Basse | Fossiles du Jurassique supérieur et des "Grès de Nubie" de la région de Sana (Yémen)[END_REF], indicate westward dispersal to the Hispanic Corridor and eastward dispersal to the Arabian platform, respectively; furthermore, pending a revision of P. jamotti and P. lannoyi Saint-Seine, 1955, the genus also occurs in Upper Jurassic freshwater deposits of the Democratic Republic of Congo [START_REF] Saint-Seine | Poissons fossiles de l'étage de Stanleyville (Congo Belge) -Première partie : La faune des argilites et schistes bitumineux[END_REF]Caillaud et al., 2017). During the Early Cretaceous, the genus Pleuropholis appears to have been decreasing in diversity and less widely distributed; it is known from five localities in the Middle East and Western Europe: the Barremian of Lebanon (Janensh, J o u r n a l P r e -p r o o f Egerton, 1924;[START_REF] El Hajj | Dysodiles from the lower Barremian of Lebanon: Insights on the fossil assemblages and the depositional environment reconstruction[END_REF], the Barremian of El Montsec and Las Hoyas in eastern Spain [START_REF] Wenz | Note préliminaire sur la faune ichthyologique du Jurassique supérieur du Montsech[END_REF][START_REF] Sanz | An Early Cretaceous faunal and floral continental assemblage: Las Hoyas fossil site (Cuenca, Spain)[END_REF][START_REF] Poyato-Ariza | Osteichthyan fishes[END_REF], the Barremian-Aptian of Belgium [START_REF] Olive | Pleuropholis germinalis n. sp., a new Pleuropholidae (Neopterygii, Teleostei) from the Early Cretaceous of Bernissart, Belgium[END_REF], and the Albian of southern Italy [START_REF] Bravi | Contributo allo studio del giacimento ad ittioliti di Pietraroja (Benevento). I. Pleuropholis decastroi n. sp. (Pisces, Actinopterygii, Pholidophoriformes)[END_REF]; with the exception of the latter, all these occurrences are from continental localities, indicating that Pleuropholis was a euryhaline genus that colonized brackish and freshwater environments at many times between the Late Jurassic and the middle Early Cretaceous. The present study reports the first Upper Cretaceous occurrence of Pleuropholis; therefore, P. danielae sp. nov. from the Cenomanian of Agoult, southeastern Morocco, represents the youngest known member of the genus and the last known occurrence of pleuropholid fishes (Fig. 6). The narrowly distributed species P. decastroi (Albian of Italy) and P. danielae sp. nov. suggest a range contraction for this formerly widespread genus, with the Mediterranean Tethys as a center of survival (paleoendemism; see [START_REF] Cowman | The biogeography of tropical reef fishes: endemism and provinciality through time[END_REF] The discovery of P. danielae sp. nov. confirms that several relictual lineages of fishes that had been flourishing during the Late Jurassic persisted until the early Late Cretaceous (a period during which they were probably severely affected by competition with diversifying modern teleosts; [START_REF] Guinot | Fish' (Actinopterygii and Elasmobranchii) diversification patterns through deep time[END_REF], and apparently became extinct at the end of the Cenomanian, possibly in relation to the Cenomanian-Turonian Boundary Event (CTBE). These forms include some holosteans such as the ionoscopiforms Spathiurus (Ionoscopidae) and Petalopteryx (?Ophiopsiellidae) from Lebanon and Agoultichthys (?Ophiopseliidae) from Morocco [START_REF] Ebert | Cerinichthys koelblae, gen. et sp. nov., from the Upper Jurassic of Cerin, France, and its phylogenetic setting, leading to a reassessment of the phylogenetic relationships of Halecomorphi (Actinopterygii)[END_REF][START_REF] El Hossny | Spathiurus dorsalis Davis, 1887, from the Upper Cretaceous of Haqel, Lebanon, and the evolutionary history of Ionoscopiformes[END_REF], the probable ionoscopiform Aphanepygus (?Ophiopsiellidae) from Croatia [START_REF] Bartram | The Macrosemiidae, a Mesozoic family of holostean fishes[END_REF], and some basal teleosts such as the newly described Pleuropholis (Pleuropholidae) species from Morocco. It is likely that these declining groups, known from very few specimens in Lebanese, Moroccan, and Croatian Cenomanian Konservat-Lagerstätten, were relatively rare in early Late Cretaceous (pre-CTBE) fish faunas and palaeoendemic, i.e. geographically restricted to small areas of the Mediterranean Tethys (Levant Platform, Atlas Gulf of the Saharan Platform, Apulian terrane). This suggests that the apparent absence in Cenomanian strata of other groups of ray-finned fishes with last known occurrences being from Albian deposits (e.g. macrosemiids and callipurbeckiid semionotiforms) could be due to problems with the detection of rare species in the fossil record (rarity bias), especially in undersampled Plattenkalk localities and understudied areas (collecting bias).

Conclusions

The Cenomanian beds of the Akrabou Formation of southeastern Morocco yielded a single specimen of pleuropholid fish, which is recognized as belonging to a new species of Pleuropholis. Interestingly, P. danielae sp. nov. is the first Late Cretaceous Pleuropholidae and thus represents the youngest known member of this family of basal teleosts. This discovery contributes to our understanding of the mid-Cretaceous turnover of marine fish faunas, which is a significant biological event characterized by the apparent extinction of relictual lineages such as pleuropholids and the diversification of modern groups such as neoteleosts. Ao, antorbital; Cl, cleithrum; De, dentary; Dsp, dermosphenotic; Dpt, dermopterotic; Ex, extrascapular; Fr, frontal; Io, infraorbital; Iop, interopercle; La, lacrimal; Mx, maxilla; Na, nasal; Op, opercle; Pa, parietal; Pmx, premaxilla; Pop, preopercle; Pt, posttemporal; Ro, rostral; Scl, supracleithrum; So, supraorbital; Sop, subopercle. Scale bar equals 10 mm. Ao, antorbital; De, dentary; Dsp, dermosphenotic; Dpt, dermopterotic; Ex, extrascapular; Fr, frontal; Io, infraorbital; Iop, interopercle; La, lacrimal; Mx, maxilla; Na, nasal; Op, opercle; Pa, parietal; Pmx, premaxilla; Pop, preopercle; Pt, posttemporal; Ro, rostral; Smx, supramaxilla; So, supraorbital; Sop, subopercle. Scale bar equals 10 mm.

  J o u r n a l P r e -p r o o f al. J o u r n a l P r e -p r o o f al., J o u r n a l P r e -p r o o f , 1998 J o u r n a l P r e -p r o o f 1998 Janensch J o u r n a l P r e -p r o o f Janensch, J o u r n a l P r e -p r o o f , 1925 J o u r n a l P r e -p r o o f 1925; J o u r n a l P r e -p r o o f ; Basse J o u r n a l P r e -p r o o f Basse teleosts J o u r n a l P r e -p r o o f teleosts ( J o u r n a l P r e -p r o o f (Patterson, J o u r n a l P r e -p r o o f Patterson, within J o u r n a l P r e -p r o o f within Teleostei J o u r n a l P r e -p r o o f Teleostei are J o u r n a l P r e -p r o o f are represented J o u r n a l P r e -p r o o f represented by J o u r n a l P r e -p r o o f by five J o u r n a l P r e -p r o o f five genera J o u r n a l P r e -p r o o f genera Austropleuropholis J o u r n a l P r e -p r o o f Austropleuropholis Zurupleuropholis J o u r n a l P r e -p r o o f Zurupleuropholis Giordano J o u r n a l P r e -p r o o f Giordano (usually J o u r n a l P r e -p r o o f (usually < J o u r n a l P r e -p r o o f < 10 J o u r n a l P r e -p r o o f 10 cm J o u r n a l P r e -p r o o f cm upturned J o u r n a l P r e -p r o o f upturned mouth, J o u r n a l P r e -p r o o f mouth, the J o u r n a l P r e -p r o o f the numerous J o u r n a l P r e -p r o o f numerous (i.e. J o u r n a l P r e -p r o o f (i.e. dorsoventrally J o u r n a l P r e -p r o o f dorsoventrally 2002; J o u r n a l P r e -p r o o f 2002;

  o u r n a l P r e -p r o o f indicates J o u r n a l P r e -p r o o f indicates that J o u r n a l P r e -p r o o f that environment, J o u r n a l P r e -p r o o f environment, most J o u r n a l P r e -p r o o f most likely J o u r n a l P r e -p r o o f likely et J o u r n a l P r e -p r o o f et al. J o u r n a l P r e -p r o o f al., J o u r n a l P r e -p r o o f , 2011; J o u r n a l P r e -p r o o f 2011; Ibrahim J o u r n a l P r e -p r o o f Ibrahim in J o u r n a l P r e -p r o o f in the J o u r n a l P r e -p r o o f the collections J o u r n a l P r e -p r o o f collections (MHNM) J o u r n a l P r e -p r o o f (MHNM), J o u r n a l P r e -p r o o f , but J o u r n a l P r e -p r o o f but is J o u r n a l P r e -p r o o f is currently J o u r n a l P r e -p r o o f currently (MNHN), J o u r n a l P r e -p r o o f (MNHN), for J o u r n a l P r e -p r o o f for study, J o u r n a l P r e -p r o o f study, within J o u r n a l P r e -p r o o f within ltraviolet J o u r n a l P r e -p r o o f ltraviolet light J o u r n a l P r e -p r o o f light 3.1. Systematic palaeontology Infraclass Teleostei Müller, 1846 (sensu Arratia, 1999) Order incertae sedis Family Pleuropholidae Saint-Seine, 1949 Genus Pleuropholis Egerton, 1858 Type species: P. attenuata Egerton, 1858 Pleuropholis danielae sp. nov. LSID: urn:lsid:zoobank.org:act:21F4D5D2-06E9-4846-8504-F43B230DABC6.

  J o u r n a l P r e -p r o o f 8504 J o u r n a l P r e -p r o o f 8504-J o u r n a l P r e -p r o o f -F43B230DABC6. J o u r n a l P r e -p r o o f F43B230DABC6. from J o u r n a l P r e -p r o o f from all J o u r n a l P r e -p r o o f all other J o u r n a l P r e -p r o o f other species J o u r n a l P r e -p r o o f species notch J o u r n a l P r e -p r o o f notch in J o u r n a l P r e -p r o o f in the J o u r n a l P r e -p r o o f the dentary, J o u r n a l P r e -p r o o f dentary, with J o u r n a l P r e -p r o o f with an J o u r n a l P r e -p r o o f an unserrated J o u r n a l P r e -p r o o f unserrated omplete J o u r n a l P r e -p r o o f omplete fla J o u r n a l P r e -p r o o f flattened J o u r n a l P r e -p r o o f ttened epithet J o u r n a l P r e -p r o o f epithet is J o u r n a l P r e -p r o o f is dedicated J o u r n a l P r e -p r o o f dedicated May J o u r n a l P r e -p r o o f May 2022, J o u r n a l P r e -p r o o f 2022, for J o u r n a l P r e -p r o o f for her J o u r n a l P r e -p r o o f her horizon J o u r n a l P r e -p r o o f horizon The J o u r n a l P r e -p r o o f

  J o u r n a l P r e -p r o o f wide, J o u r n a l P r e -p r o o f wide, with J o u r n a l P r e -p r o o f with frontal J o u r n a l P r e -p r o o f frontal posteriorly, J o u r n a l P r e -p r o o f posteriorly, nasal, J o u r n a l P r e -p r o o f nasal, and J o u r n a l P r e -p r o o f and is J o u r n a l P r e -p r o o f is the J o u r n a l P r e -p r o o f the times J o u r n a l P r e -p r o o f times that J o u r n a l P r e -p r o o f that of J o u r n a l P r e -p r o o f of the J o u r n a l P r e -p r o o f the nasal J o u r n a l P r e -p r o o f nasal a J o u r n a l P r e -p r o o f a well J o u r n a l P r e -p r o o f well-J o u r n a l P r e -p r o o f -developed J o u r n a l P r e -p r o o f developed of J o u r n a l P r e -p r o o f of the J o u r n a l P r e -p r o o f the frontal. J o u r n a l P r e -p r o o f frontal. The J o u r n a l P r e -p r o o f The dermopterotic. J o u r n a l P r e -p r o o f dermopterotic. The J o u r n a l P r e -p r o o f The posterior J o u r n a l P r e -p r o o f posterior to J o u r n a l P r e -p r o o f to J o u r n a l P r e -p r o o f the J o u r n a l P r e -p r o o f the frontal. J o u r n a l P r e -p r o o f frontal. extrascapular J o u r n a l P r e -p r o o f extrascapular is J o u r n a l P r e -p r o o f is mid J o u r n a l P r e -p r o o f mid-J o u r n a l P r e -p r o o f -length J o u r n a l P r e -p r o o f length of J o u r n a l P r e -p r o o f of the J o u r n a l P r e -p r o o f the the dermosphenotic is a somewhat diamond-shaped bone that borders the posterior dorsal angle of the orbit. It is in contact with the anterior part of the dorsal edge of infraorbital 4. At least two supraorbitals are preserved, forming the dorsal margin of the orbit.

  the Albian of Italy (cf. Taverne and Capasso, 2019) and P. germinalis Olive, Taverne and Brito, 2022 from the Barremian-Aptian of Belgium the new species does not present a leptolepid notch in the median-dorsal part of the dentary. Posteriorly, the dentary sutures with the angular. The jaw looks toothless. The preopercle is an L-shaped bone in which the horizontal limb is slightly larger than the vertical limb. These limbs are placed perpendicular to each other (Figs. 2, 3). The preopercular canal is close and parallel to the ventral and dorsal borders of the bone.The opercular series consists of the large opercle, a subtriangular subopercle and an elongated interopercle. Only part a relatively narrow branchiostegal ray is preserved. The posttemporal bone is somewhat oval and located behind the skull; the posterior edge overlies the dorsal end of the supracleithrum. The supracleithrum laterally overlaps the dorsal end of the cleithrum. The cleithrum has an elongated lower arm, less developed than the dorsal one. The pectoral fin is well developed and nine rays are preserved. It is positioned close to the ventral margin of the body, on the lower border between the flank scales and the scales immediately below. The pelvic fin is located at the level of the 15th transverse row of scales. Only four branched rays are preserved. No bones of the pelvic girdle are visible. J o u r n a l P r e -p r o o f and J o u r n a l P r e -p r o o f and the J o u r n a l P r e -p r o o f the triangular J o u r n a l P r e -p r o o f triangular bone J o u r n a l P r e -p r o o f bone that J o u r n a l P r e -p r o o f that the J o u r n a l P r e -p r o o f the ventral J o u r n a l P r e -p r o o f ventral border J o u r n a l P r e -p r o o f border 1988 J o u r n a l P r e -p r o o f 1988, J o u r n a l P r e -p r o o f , from J o u r n a l P r e -p r o o f from the J o u r n a l P r e -p r o o f the Albian J o u r n a l P r e -p r o o f Albian Taverne J o u r n a l P r e -p r o o f Taverne and J o u r n a l P r e -p r o o f and Brito, J o u r n a l P r e -p r o o f Brito, 2022 J o u r n a l P r e -p r o o f 2022 present J o u r n a l P r e -p r o o f present a J o u r n a l P r e -p r o o f a leptolepid J o u r n a l P r e -p r o o f leptolepid notch J o u r n a l P r e -p r o o f notch sutures J o u r n a l P r e -p r o o f sutures with J o u r n a l P r e -p r o o f with the J o u r n a l P r e -p r o o f the angular. J o u r n a l P r e -p r o o f angular. -J o u r n a l P r e -p r o o f -shaped J o u r n a l P r e -p r o o f shaped bone J o u r n a l P r e -p r o o f bone in J o u r n a l P r e -p r o o f in limbs J o u r n a l P r e -p r o o f limbs are J o u r n a l P r e -p r o o f are placed J o u r n a l P r e -p r o o f placed perpendicular J o u r n a l P r e -p r o o f perpendicular parallel J o u r n a l P r e -p r o o f parallel to J o u r n a l P r e -p r o o f to the J o u r n a l P r e -p r o o f the ventral J o u r n a l P r e -p r o o f ventral sub J o u r n a l P r e -p r o o f subtriangular J o u r n a l P r e -p r o o f triangular Both the dorsal and the anal fins are triangular and located posteriorly in the body, at the level of the 29th and 26th transversal row of scales, respectively (Figs. 1

  fig. 36; J o u r n a l P r e -p r o o f 36; Taverne J o u r n a l P r e -p r o o f Taverne J o u r n a l P r e -p r o o f lower J o u r n a l P r e -p r o o f lower lobe J o u r n a l P r e -p r o o f lobe ( J o u r n a l P r e -p r o o f (Figs. J o u r n a l P r e -p r o o f Figs.

  Despite these problems, P. danielae sp. nov. clearly differs from the other species of Pleuropholis by its considerably more slender body shape. The absence of a leptolepid notch in the dentary distinguishes P. danielae n. sp from P. jamotti Saint-Seine, 1955 (a species, like P. lannoyi J o u r n a l P r e -p r o o f synapomorphies J o u r n a l P r e -p r o o f synapomorphies of J o u r n a l P r e -p r o o f of the J o u r n a l P r e -p r o o f the family J o u r n a l P r e -p r o o f family ganoid J o u r n a l P r e -p r o o f ganoid scales J o u r n a l P r e -p r o o f scales covering J o u r n a l P r e -p r o o f covering complete J o u r n a l P r e -p r o o f complete orbital J o u r n a l P r e -p r o o f orbital series J o u r n a l P r e -p r o o f series formed J o u r n a l P r e -p r o o f formed Gallo J o u r n a l P r e -p r o o f Gallo, J o u r n a l P r e -p r o o f , 2002 J o u r n a l P r e -p r o o f 2002; J o u r n a l P r e -p r o o f ; Olive J o u r n a l P r e -p r o o f Olive upturned J o u r n a l P r e -p r o o f upturned maxilla J o u r n a l P r e -p r o o f maxilla and J o u r n a l P r e -p r o o f and a J o u r n a l P r e -p r o o f a Seine J o u r n a l P r e -p r o o f Seine, J o u r n a l P r e -p r o o f , 1949; J o u r n a l P r e -p r o o f 1949; Patterson J o u r n a l P r e -p r o o f Patterson Capasso J o u r n a l P r e -p r o o f Capasso, J o u r n a l P r e -p r o o f , 2019 J o u r n a l P r e -p r o o f 2019; J o u r n a l P r e -p r o o f ; Olive J o u r n a l P r e -p r o o f Olive biggest J o u r n a l P r e -p r o o f biggest problems

  J o u r n a l P r e -p r o o f and J o u r n a l P r e -p r o o f and distribution J o u r n a l P r e -p r o o f distribution far, J o u r n a l P r e -p r o o f far, t J o u r n a l P r e -p r o o f the J o u r n a l P r e -p r o o f he only J o u r n a l P r e -p r o o f only unambig J o u r n a l P r e -p r o o f unambig Bathonian J o u r n a l P r e -p r o o f Bathonian of J o u r n a l P r e -p r o o f of southern J o u r n a l P r e -p r o o f southern known J o u r n a l P r e -p r o o f known record J o u r n a l P r e -p r o o f record reflects J o u r n a l P r e -p r o o f reflects a J o u r n a l P r e -p r o o f a restricted J o u r n a l P r e -p r o o f restricted neoendemism J o u r n a l P r e -p r o o f neoendemism; J o u r n a l P r e -p r o o f ; see J o u r n a l P r e -p r o o f see highest J o u r n a l P r e -p r o o f highest diversity J o u r n a l P r e -p r o o f diversity due J o u r n a l P r e -p r o o f due to J o u r n a l P r e -p r o o f to a J o u r n a l P r e -p r o o f a preservational J o u r n a l P r e -p r o o f preservational described J o u r n a l P r e -p r o o f described from J o u r n a l P r e -p r o o f from Cerin J o u r n a l P r e -p r o o f Cerin Bavaria J o u r n a l P r e -p r o o f Bavaria in J o u r n a l P r e -p r o o f in Germany J o u r n a l P r e -p r o o f Germany ( J o u r n a l P r e -p r o o f (Egerton,

  . The new discovery extends the stratigraphic range of this group by about 15 million years and makes Pleuropholis one of the most temporally long-lived actinopterygian genera of all time, with a fossil record spanning ~75 million years. Based on 1108 time ranges calculated from Cavin et al.'s (2021a) database, Cavin et al. (2021b: fig. 7) found that the average time duration of the ray-finned fish genera having their first occurrence in the Devonian-Palaeocene interval is 13.3 million years all environments combined, with only a dozen genera having a longer range than that of Pleuropholis.

  fig. 7 J o u r n a l P r e -p r o o f 7) J o u r n a l P r e -p r o o f )

  to J o u r n a l P r e -p r o o f to problems J o u r n a l P r e -p r o o f problems especially J o u r n a l P r e -p r o o f especially in J o u r n a l P r e -p r o o f in undersampled J o u r n a l P r e -p r o o f undersampled the J o u r n a l P r e -p r o o f the Akrabou J o u r n a l P r e -p r o o f Akrabou Formation J o u r n a l P r e -p r o o f Formation fish, J o u r n a l P r e -p r o o f fish, which J o u r n a l P r e -p r o o f which is J o u r n a l P r e -p r o o f is recognized J o u r n a l P r e -p r o o f recognized danielae J o u r n a l P r e -p r o o f danielae sp. J o u r n a l P r e -p r o o f sp. nov. J o u r n a l P r e -p r o o f nov. is J o u r n a l P r e -p r o o f is the J o u r n a l P r e -p r o o f the known J o u r n a l P r e -p r o o f known member J o u r n a l P r e -p r o o f member of J o u r n a l P r e -p r o o f of this J o u r n a l P r e -p r o o f this mid J o u r n a l P r e -p r o o f mid Cretaceous J o u r n a l P r e -p r o o f Cretaceous J o u r n a l P r e -p r o o f teleosts. J o u r n a l P r e -p r o o f teleosts. In: J o u r n a l P r e -p r o o f In: Record. J o u r n a l P r e -p r o o f Record. Verlag J o u r n a l P r e -p r o o f Verlag Dr. J o u r n a l P r e -p r o o f Dr. skeleton J o u r n a l P r e -p r o o f skeleton with J o u r n a l P r e -p r o o f with special J o u r n a l P r e -p r o o f special identifying J o u r n a l P r e -p r o o f identifying homologies. J o u r n a l P r e -p r o o f homologies. Fishes J o u r n a l P r e -p r o o f Fishes 4 J o u r n a l P r e -p r o o f 4 -J o u r n a l P r e -p r o o f -Homology J o u r n a l P r e -p r o o f Homology Morphology, J o u r n a l P r e -p r o o f Morphology, taxonomy, J o u r n a l P r e -p r o o f taxonomy, and J o u r n a l P r e -p r o o f and Teleostei) J o u r n a l P r e -p r o o f Teleostei). J o u r n a l P r e -p r o o f . Soc. J o u r n a l P r e -p r o o f Soc. Vert. J o u r n a l P r e -p r o o f Vert. The J o u r n a l P r e -p r o o f The J o u r n a l P r e -p r o o f Actinopterygii) J o u r n a l P r e -p r o o f Actinopterygii) du J o u r n a l P r e -p r o o f du Mé J o u r n a l P r e -p r o o f Mé environnementales J o u r n a l P r e -p r o o f environnementales et J o u r n a l P r e -p r o o f et pleuropholid, J o u r n a l P r e -p r o o f pleuropholid, Gondwanapleuropholis J o u r n a l P r e -p r o o f Gondwanapleuropholis the J o u r n a l P r e -p r o o f the Jurassic J o u r n a l P r e -p r o o f Jurassic of J o u r n a l P r e -p r o o f of north J o u r n a l P r e -p r o o f north new J o u r n a l P r e -p r o o f new Cenomanian J o u r n a l P r e -p r o o f Cenomanian other J o u r n a l P r e -p r o o f other early J o u r n a l P r e -p r o o f early Late J o u r n a l P r e -p r o o f Late Cretac J o u r n a l P r e -p r o o f Cretac Boudad, J o u r n a l P r e -p r o o f Boudad, L., J o u r n a l P r e -p r o o f L., Meister, J o u r n a l P r e -p r o o f Meister, Dyke, J o u r n a l P r e -p r o o f Dyke, G., J o u r n a l P r e -p r o o f G., Hua, J o u r n a l P r e -p r o o f Hua, southeastern J o u r n a l P r e -p r o o f southeastern J o u r n a l P r e -p r o o f Upper J o u r n a l P r e -p r o o f Upper Jurassic J o u r n a l P r e -p r o o f Jurassic the J o u r n a l P r e -p r o o f the phylogenetic J o u r n a l P r e -p r o o f phylogenetic relationships J o u r n a l P r e -p r o o f relationships 38, J o u r n a l P r e -p r o o f 38, e1420071. J o u r n a l P r e -p r o o f e1420071. , J o u r n a l P r e -p r o o f , Garcia J o u r n a l P r e -p r o o f Garcia, J o u r n a l P r e -p r o o f , G. J o u r n a l P r e -p r o o f G., J o u r n a l P r e -p r o o f , Horne J o u r n a l P r e -p r o o f Horne lower J o u r n a l P r e -p r o o f lower Barremian J o u r n a l P r e -p r o o f Barremian of J o u r n a l P r e -p r o o f of environ J o u r n a l P r e -p r o o f environment J o u r n a l P r e -p r o o f ment reconstruction. J o u r n a l P r e -p r o o f reconstruction. D., J o u r n a l P r e -p r o o f D., Abi J o u r n a l P r e -p r o o f Abi Saad, J o u r n a l P r e -p r o o f Saad, P., J o u r n a l P r e -p r o o f P., Cretaceous J o u r n a l P r e -p r o o f Cretaceous of J o u r n a l P r e -p r o o f of Haqel, J o u r n a l P r e -p r o o f Haqel, Cret. J o u r n a l P r e -p r o o f Cret. Res. J o u r n a l P r e -p r o o f Res. 120, J o u r n a l P r e -p r o o f 120, 104619 J o u r n a l P r e -p r o o f 104619 1858 J o u r n a l P r e -p r o o f 1858. J o u r n a l P r e -p r o o f . British J o u r n a l P r e -p r o o f British fossils J o u r n a l P r e -p r o o f fossils Remains J o u r n a l P r e -p r o o f Remains J o u r n a l P r e -p r o o f paleontology J o u r n a l P r e -p r o o f paleontology 928, J o u r n a l P r e -p r o o f 928, 1 J o u r n a l P r e -p r o o f 1-J o u r n a l P r e -p r o o f -216. J o u r n a l P r e -p r o o f 216. vom J o u r n a l P r e -p r o o f vom Libanon. J o u r n a l P r e -p r o o f Libanon. Cenomanian J o u r n a l P r e -p r o o f Cenomanian florule J o u r n a l P r e -p r o o f florule and J o u r n a l P r e -p r o o f and climatological J o u r n a l P r e -p r o o f climatological inferences. J o u r n a l P r e -p r o o f inferences. actinopterygian J o u r n a l P r e -p r o o f actinopterygian fish J o u r n a l P r e -p r o o f fish fauna J o u r n a l P r e -p r o o f fauna Solnhofen J o u r n a l P r e -p r o o f Solnhofen (Bavaria, J o u r n a l P r e -p r o o f r n a l P r e -p r o o f G., Pasini, J o u r n a l P r e -p r o o f Pasini, A., J o u r n a l P r e -p r o o f A., Garassino, J o u r n a l P r e -p r o o f Garassino, from J o u r n a l P r e -p r o o f from the J o u r n a l P r e -p r o o f the Late J o u r n a l P r e -p r o o f Late Cretaceous J o u r n a l P r e -p r o o f Cretaceous 106, J o u r n a l P r e -p r o o f 106, J o u r n a l P r e -p r o o f acanthomorph J o u r n a l P r e -p r o o f acanthomorph fi J o u r n a l P r e -p r o o f fishes J o u r n a l P r e -p r o o f shes from J o u r n a l P r e -p r o o f from , J o u r n a l P r e -p r o o f , B.D.E., J o u r n a l P r e -p r o o f B.D.E., 2013 J o u r n a l P r e -p r o o f 2013 actinopterygian J o u r n a l P r e -p r o o f actinopterygian fauna J o u r n a l P r e -p r o o f fauna on J o u r n a l P r e -p r o o f on the J o u r n a l P r e -p r o o f the palaeoenvironment. J o u r n a l P r e -p r o o f palaeoenvironment. Mesozoic J o u r n a l P r e -p r o o f Mesozoic Fishes J o u r n a l P r e -p r o o f Fishes 5 J o u r n a l P r e -p r o o f 5 -J o u r n a l P r e -p r o o f -Global J o u r n a l P r e -p r o o f Global nchen, J o u r n a l P r e -p r o o f nchen, pp. J o u r n a l P r e -p r o o f pp. 525 J o u r n a l P r e -p r o o f 525-J o u r n a l P r e -p r o o f -548. J o u r n a l P r e -p r o o f 548. B J o u r n a l P r e -p r o o f Brito J o u r n a l P r e -p r o o f rito, J o u r n a l P r e -p r o o f , P. J o u r n a l P r e -p r o o f P.M. J o u r n a l P r e -p r o o f M., J o u r n a l P r e -p r o o f , 2022 J o u r n a l P r e -p r o o f 2022 Teleostei) J o u r n a l P r e -p r o o f Teleostei) from J o u r n a l P r e -p r o o f from J o u r n a l P r e -p r o o f Jimenez J o u r n a l P r e -p r o o f Jimenez Early J o u r n a l P r e -p r o o f Early Cretaceous J o u r n a l P r e -p r o o f Cretaceous faunal J o u r n a l P r e -p r o o f faunal (Cuenca, J o u r n a l P r e -p r o o f (Cuenca, Spain). J o u r n a l P r e -p r o o f Spain). Geobios J o u r n a l P r e -p r o o f Geobios Bull J o u r n a l P r e -p r o o f Bull. J o u r n a l P r e -p r o o f . Soc. J o u r n a l P r e -p r o o f Soc. Géol. 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New J o u r n a l P r e -p r o o f New data J o u r n a l P r e -p r o o f data on J o u r n a l P r e -p r o o f on fi J o u r n a l P r e -p r o o f fish J o u r n a l P r e -p r o o f sh from J o u r n a l P r e -p r o o f from the J o u r n a l P r e -p r o o f the Lower J o u r n a l P r e -p r o o f Lower G. J o u r n a l P r e -p r o o f G.

Fig. 1 .

 1 Fig. 1. Pleuropholis danielae sp. nov., holotype MHNM.KK.393 photographed under natural light

Fig. 2 .

 2 Fig. 2. Pleuropholis danielae sp. nov., holotype MHNM.KK.393, detail of the head region

Fig. 3 .

 3 Fig. 3. Reconstruction of the skull of Pleuropholis danielae sp. nov. Abbreviations: Ang, angular;

Fig. 4 .

 4 Fig. 4. Pleuropholis danielae sp. nov., holotype MHNM.KK.393, detail of the dorsal and anal fins

Fig. 5 .Fig. 6 .

 56 Fig. 5. Pleuropholis danielae sp. nov., holotype MHNM.KK.393, detail of the caudal region

J o u r n a l P r e -p r o o f J o u r n a l P r e -p r o o f J o u r n a l P r e -p r o o f

Acknowledgements

We wish to express our gratitude to D. Martill and D. Dutheil for their comments on the manuscript and to L. Cazes and P. Loubry for the photographs. We are grateful to N.-E. Jalil for providing a MHNM collection number. We also thank E. Koutsoukos as well as L. Cavin and an J o u r n a l P r e -p r o o f (Espagne). Bull. Soc. Géol. Fr., Sér. 7, 10, 116-119. Woodward, A.S., 1919. The fossil fishes of the English Wealden and Purbeck formations. Part III. Monogr. Palaeontogr. Soc. 71, Issue 340, 105-148. J o u r n a l P r e -p r o o f

The specimen is assigned to a new species of the genus Pleuropholis, P. danielae. This is the first known occurrence of Pleuropholidae in the Upper Cretaceous.

Pleuropholid fishes may have become extinct at the Cenomanian-Turonian boundary.

This adds to our understanding of the mid-Cretaceous turnover of marine fish faunas.

J o u r n a l P r e -p r o o f

Cretaceous turnover of marine fish faunas J o u r n a l P r e -p r o o f Cretaceous turnover of marine fish faunas

Declaration of interests

☒ The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

☐The authors declare the following financial interests/personal relationships which may be considered as potential competing interests: