An unexpected New World mole (Scalopini, Mammalia) from the Pliocene of Europe sheds light on the phylogeny of talpids

An unexpected New World mole (Scalopini, Mammalia) from the Pliocene of Europe sheds light on the phylogeny of talpids | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Article An unexpected New World mole (Scalopini, Mammalia) from the Pliocene of Europe sheds light on the phylogeny of talpids Adriana Linares-Martín, Marc Furió, Bruno Gómez Soler, Jordi Agustí, and 6 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-6215385/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 10 Jul, 2025 Read the published version in Scientific Reports → Version 1 posted 10 You are reading this latest preprint version Abstract The Pliocene Konservat-Lagerstätten maar lake site of Camp dels Ninots (NE Iberian Peninsula) has recently delivered a partial skeleton of a mole (family Talpidae) with many elements in anatomical connection. At a first glance, molar and humerus size, geological time interval, and geographical location suggested that this specimen could correspond to Talpa minor . However, after some mechanical preparation of the clay block (matrix removal, consolidation, and cleaning) and a micro-CT scan, this excellently preserved specimen turned out to be an unknown species to science. The resulting 3D models of this new form, Vulcanoscaptor ninoti gen. et sp. nov., revealed some peculiar morphological traits in teeth, mandible, and postcranial elements, which according to the phylogenetic analysis carried out, would allocate this new species within the tribe Scalopini. This is surprising, because the representatives of this tribe are nowadays restricted to North America and Asia, and only some taxa had been previously reported in the Oligocene and Miocene fossil record from Europe. The postcranial construction of this specimen reveals a highly fossorial lifestyle supported by a complex forelimb structure. How such a specialized digging animal reached the maar lake sediments where it was finally preserved is still to be solved. Some hypotheses consider swimming abilities for this extinct species. Alternatively, this specimen could be the remaining portions of a floated or scavenged carcass whose remains fell into the lake and reached the anoxic bottom. Biological sciences/Evolution/Palaeontology Biological sciences/Evolution/Phylogenetics Biological sciences/Evolution/Taxonomy Camp dels Ninots Maar Konservat-Lagerstätten Spain Fossorial Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Figure 8 Introduction The Talpidae are probably one of the most intriguing families of mammals from the paleobiogeographic point of view. Only a few species within this group can be considered truly cursorial, because most of them show in some extent adaptations to fossorial activity. Indeed, many talpids are extremely adapted to subterranean and aquatic lifestyles, and their subaerial movements or out of water displacements are rather clumsy. Out of tunnels or rivers, most species of moles are highly vulnerable to potential predators. Thus, anyone would expect these animals to follow a rather predictable and constant (paleo-)biogeographic evolutionary history, with no significant migrations, considering all the possible physical and biological barriers that could constrain their distribution. Surprisingly, the fossil record is rather indicating a quite different story. The genus Eotalpa is the oldest fossorial representative of the family known hitherto 1 , 2 . According to the fossil record of the younger representatives, few groups have originated and stayed in the same region that they currently inhabit. For instance, Schwermann et al . 3 conducted a thorough study on the evolutionary history of the Scalopini, concluding that numerous transcontinental colonization events occurred between Asia, Europe, and North America, to the extent that no unambiguous location for the origin of the group could be identified. The recent discovery of Alpiscaptulus medogensis , a living species of Scalopini from the Himalayas 4 , has just added some more questions to this enigma. In a similar way, uropsilines are nowadays represented only by the genus Uropsilus (but see disagreements in He et al. 5 ) and restricted to Central and Southeastern Asia. The fossil record shows, however, that its Oligocene, Miocene and Pliocene relatives ( Desmanella , Mystipterus , Asthenoscapter , Mygatalpa , Theratiskos ) lived in European, Minor Asian and North American lands 6 – 11 . Another example is found in the tribe condylurini, nowadays being solely represented by the endemic North American star-nosed mole, Condylura cristata . However, unequivocal representatives of the genus have been found in the Pliocene of Poland 12 , 13 and the Middle Miocene of Kazakhstan 14 . Likewise, Neurotrichus gibbsii is the only living representative of the tribe Neurotrichini, according to Hutterer 15 . The hypothesis of a North American origin of Neurotrichus judging by its solely current occurrence in this continent would apparently be parsimonious. Nonetheless, the cladistic analysis performed by Schwermann and Thompson 16 suggested that this genus could find its closest living relative in the genus Scaptonyx , which mostly currently inhabits elevated areas from China. The extinct genus Rzebikia is a very similar form to Neurotrichus , but it is only found in Plio-Pleistocene sites from Europe 17 . Both genera, Neurotrichus and Rzebikia are putative descendants of an Asian form, Quyania 18 . Far from being simple, the most likely explanation to such distribution according to Sansalone et al. 17 is that the original Asian stock of moles finally derived into N. gibbsii after colonizing North America in the Early Pliocene, while another one or two migration waves towards Eastern Europe could have resulted in several European forms. The case of the swimming forms of the Desmanini is similarly puzzling. The Russian desman ( Desmana moschata ), an extant endemic species living in Russia, Ukraine and Kazakhstan, is the only survivor of a genus that apparently originated in the south of the Iberian Peninsula 19 . Ironically, the Iberian desman ( Galemys pyrenaicus ), an endemic form that currently inhabits some stream headwaters in this part of the European continent, belongs to a genus of unknown geographic origin. During the Late Neogene, the tribe had a much wider distribution, although restricted to Europe and parts of Asia Minor 20 . Some of these apparent inconsistencies are probably rooted in a very limited view of the talpid fossil record. On the one hand, moles are rather conservative in their morphology, likely due to the constant conditions where they thrive. It could be that once acquired an efficient capacity for digging (or swimming), there were few possible morphological modifications to adapt to different lifestyles 21 . On the other hand, the fossil finds of talpids usually correspond to disarticulated and/or incomplete elements, not always easy to identify to the species level. In many cases, it is difficult to discriminate whether the (little) variation in size and morphology observed in the fossil assemblages is due to interspecific or intraspecific variability. A good example is found on the uncertainty of how many species of Talpa occur in the European fossil record, a debate that has been extended for decades in specialized literature 9 , 22 , 23 . For a comprehensive list of approaches to link humeri and dentitions in fossil talpids, the reader is referred to van den Hoek Ostende and Fejfar 24 . Fortunately, the fossil record is not always limited to patchy finds and fragmentary elements. In the last years, some fossil-Lagerstätten 25 sites have provided several exceptional specimens with many skeletal elements that can be clearly ascribed to the same individual 16 , 26 . Such finds are two-fold precious. On the one hand, they allow checking useful morphological characters located in fragile parts, which are not frequently preserved in the fossil record. On the other hand, they link dental to postcranial elements, thus becoming very useful in the calculation of the relative sizes between both. Each one of these exceptional finds constitutes a reference milestone to which compare many other isolated fossil elements of talpids, mostly teeth and humeri. In the present work, we study a new partial fossil skeleton of a burrowing mole that has been recently discovered in the Pliocene locality of Camp dels Ninots (CN) in NE Spain. This exceptional fossil find preserves several postcranial elements associated to almost complete dentitions and the whole mandible, all of them clearly belonging to the same individual. This specimen was tentatively ascribed to Talpa minor at a first glance, considering the chronology and location of the fossil site in combination to its general size. Nevertheless, our colleague Dr. Lars van den Hoek Ostende (Naturalis Biodiversity Center, Leiden, The Netherlands) noticed in a previous stage of the present work some morphological details against its ascription to the genus Talpa . The rest of the talpid genera hitherto identified in Pliocene sites from Spain 27 , namely Archaeodesmana , Desmana , Galemys , and Desmanella could be easily discarded as well. Therefore, in the present study we aim to solve: 1) which is the species represented in this fossil site, 2) the phylogenetic position of this fossil among extant and extinct talpids, 3) how adapted was this species to excavate and live underground, and more specifically, 4) how did this specimen reach the sedimentary environment where it was finally preserved. Geological settings CN is a Pliocene site located in the town of Caldes de Malavella, in the province of Girona, in the NE of the Iberian Peninsula (Fig. 1 ). Some analyses using magneto and cyclostratigraphic techniques had provided an age of ca 3.25 Ma 28 . However, recent 40Ar/39Ar datation techniques are indicative of a much older age for the site, likely Early Pliocene (work in progress). The first fragmentary bone from this locality was found in 1985 and it was assigned to the genus Leptobos by Vicente 29 . Some later prospections resulted in new fossil inputs 30 – 34 , but it was in 2003 when the Institut Català de Paleoecologia Humana i Evolució Social (IPHES-CERCA) started the systematic research projects and excavations. Since then, this locality has delivered several articulated skeletons of vertebrates of bovids ( Alephis tigneresi) , tapirs ( Tapirus arvernensis) , rhinoceros ( Stephanorhinus cf. jeanvireti) , snakes ( Natrix maura ), turtles ( Mauremys leprosa and Chelydropsis cf. pontica ), amphibians (Pleurodelinae indet., Lissotriton aff. helveticus , and Pelophylax sp.), and freshwater fishes ( Leuciscus sp. and Luciobarbus sp.) 31 – 34 . To a lesser extent, also diatoms, invertebrates (insects), micro- and macroflora (such as Laurophyllum sp. ) have been found 28 , 35 , 36 . CN site was in origin a maar lake of a volcano found in the Catalan Volcanic Complex (CVC) at the southern border of La Selva basin. There, volcanic activity ranged from the Miocene till the Holocene. The CVC is part of the volcanic provinces of the European Rift System into the Catalan Coastal Ranges 37 (Fig. 1 ). La Selva basin was the result of an extensional tectonic episode during the Miocene-Pliocene, generated by the movement of two sets of faults oriented ENE-WSW and NW-SE of a previously fractured Paleozoic basement (for further details about the geotectonic setting see Roca et al. 38 and Tassone et al. 39 ). The tectonic activity favored the volcanism during the Miocene, with intense phases during the Pliocene 40 , 41 . Several volcanic episodes are related with the meteoric water infiltration according to fractures of the basement and porosity of Pliocene materials, which filled the basin. The near surface magma-water interactions generated a mixed volcanism with an alternating eruptive-effusive phase. Phreatomagmatic vulcanism of explosive phases resulted in the formation of a maar crater 30 , 42 . Post volcanic sediments were accumulated within the maar crater as lacustrine deposits. The sedimentary infill of CN is formed by the typical vertical stratigraphic succession in maars 43 , 44 . The one described below is taken as a reference for this fossil site and represents a thick section of 8 meters at the Can Argilera excavation sector, with four units from its base to the top (Figs. 2 , 3 ). Unit 1 encompasses greyish clays, sandstones and diatomites. Unit 2 is represented by greenish laminated clays with diatoms, and it is differentiated in other four subunits; 2.1, 2.2 and 2.4 are characterized by the presence of carbonates. The subunit 2.3 (see 'Detailed Location' in Fig. 2 ) has been recorded in the Can Argilera sector and it is divided in fine-grained reddish sands with silty admixture (Layer 10) and lacustrine greenish silts with clay laminations (Layer 11). The latter subunit is the most remarkable one in terms of articulated skeletons of mammals and plant remains. Lastly, unit 3 is formed by reddish laminated clays and silty slope wash deposits. For further details on the specific geology of the site the reader is referred to Gómez de Soler et al. 31 , Jiménez-Moreno et al. 28 , Oms et al. 42 , Rodríguez-Salgado et al. 45 . Results Systematic paleontology Order Eulipotyphla Waddell, Okada & Hasegawa, 1999 Family Talpidae Fischer, 1814 Tribe Scalopini Gill, 1875 Genus Vulcanoscaptor nov. Vulcanoscaptor ninoti sp. nov. (Figs 4, 5, 6 and 7). Authorship of genus and species: Linares-Martín, 2025 Etymology. Name of the genus derived from the Latin word of ‘Vulcan’, the Roman god of fire, in reference to the volcanic nature of the source area, and ‘-scaptor’, from the ancient Greek word ‘scaptein’, to dig. Name of the species invoking ‘ninot’, the local word to refer the opaline nodules ‘doll-shaped’ typically found in the type-locality of the species, Camp dels Ninots. Holotype. CN10-O17-NIV11-12, partial skeleton with cranial and postcranial elements. Storage. IPHES-CERCA facilities. Stratigraphic range: Hitherto restricted to its type locality, Lower Pliocene. Diagnosis (genus and species). Small sized mole with dental formula ???3/2143. Doubled mesostyle in M1 and M2. Presence of paraconule and absence of hypocone in M2. Double rooted P4. Presence of a parastyle in P4. Lower premolar row without gaps. Absence of i3 and enlarged i2. Absence of metastylid in m2. Robust and small postcranial remains. Pit for M. flexor digitorum profundus ligament present. Straight medial edge of humeral trochlea. Fusiform shape of the humeral capitulum. Well-developed and transverse olecranon crest. Anconal and coronoid processes present in the ulna. Presence of capitular process in the radius. Scaphoid and lunar not co-ossified. Differential diagnosis. See Supplementary Information 1. Supplementary Tables of measurements and comparisons with other selected species of Talpidae can be found in Supplementary Information 2 (Tables S1–S9). Description The partial skeleton of the mole lies partially embedded by the sediment exposing the lateral side of the remains. Some skeletal elements are found in anatomical connection such as the mandible, the dentition and the postcranial elements (right forelimb). The tibiofibula is isolated from the rest of elements (Fig. 4). Skull and upper dentition. The skull is partially preserved but strongly damaged. None of the cranial structures can be identified through digital reconstruction. Because the remains are not scattered, the outline can be delineated. The left upper tooth row is incomplete, but it preserves the molars, three premolars and one incisor (Fig. 5a). The incisor is displaced pointing mesially. The length of the premolars decreases towards the anterior part, P4>P3>P2. The premolars are all doubled-rooted. P2 (L = 0.80; W = 0.38) is clearly unicuspid. P3 (L = 0.85; W = 0.47) is mostly dominated by a high central cusp with a curved posterior ridge which bears a metastyle at its posterior end. The anterior ridge of both premolars is straight. The protocone in P4 (L = 1.36; W = 1.13) lies mesiolingual to the paracone. The postparacrista is straight and slightly curved at the posterobuccal end. The preparacrista is curved and wide at the end. The molars are double-rooted and become smaller distally, M1>M2>M3. There is no cingulum in any of them. In the M1 (L = 2.27; W = 1.78) the metacone is expanded distolingually being the paracone slightly larger than the metacone. The trigon basin is deep and wide. The protocone of the M1 is lower than that of M2 and there is no evidence of the presence of the hypocone. The mesostyle is divided into two cusps separated by a deep valley. The parastyle and the metastyle are also well developed. In the M2 (L = 1.82; W = 1.66), the trigon basin forms a deep valley as in M1. The paracone is the highest cusp and the protocone is the smallest. The paracrista is a bit shorter than the postparacrista and similar in length with the postmetacrista. No hypocone is discerned, whereas the metastyle and parastyle are well-developed. The mesostyle valley follows the same pattern as in M1. In the M3 (L = 1.52; W = 1.35), the trigon basin is wide but not as deep as in the other molars. The metacone and the paracone have a similar height in contrast to the protocone, which is smaller. All the cusps are wide and rounded. The parastyle is well developed. Mandible and lower dentition. Both hemimandibles and most of the lower dentition (Fig. 5b) are preserved. Only the right hemimandible and its whole dentition (Fig. 5c) is exposed but digital reconstructions demonstrates that the left hemimandible additionally preserves two molars, two premolars, the canine and one incisor. Anterior mental foramina is observed below p2 and p3, and a less clear posterior one is placed below m1 and m2. The corpus mandibulae is slender and elongated. The distal part becomes wider and convex towards the angular process. The anterior profile of the coronoid process is slightly curved and straightens towards the tip of the coronoid process. It has a convex shape with a faint notch. The posterior profile delineates a concave curvature that connects to the condylar process. The condylar process points backwards, and it is longer than the angular process. The angular process is short, and the tip ends anterior to the angular process and posterior to the coronoid process. Measurements (mm): Length of the corpus mandibulae = 14.63; maximum thickness of mandible (below m2) = 1.40; maximum height of mandible (below m2) = 1.45, minimum thickness of mandible (below i1) = 0.43; Minimum height of mandible (below i2) = 1.12; height of coronoid process = 4.94. Regarding the dentition, the incisors and the canine present an elongated and flat crown. Both incisors are single-rooted. The i2 is the largest and widest of the incisors. Its shape is spatulated. The canine is smaller than the incisors. The premolars are doubled-rooted unicuspids and there are no gaps between them. The lateral outlines of their crowns are almost triangular. All of them have divergent and rather stout roots with rounded tips. Towards the distal side, premolars are progressively larger, p1< p2 < p3 < p4. The cusps in all of them are rather high and sharp. The p1 (Right: L = 0.52; W = 0.35; Left: L = 0.50, W = 0.38) stands as a simple single-cusped tooth. The posterior ridge of p2 (Right: L = 0.65; W = 0.33; Left: L = 0.60; W = 0.32) and p3 (L = 0.71; W = 0.33) are curved and wide towards the posterior end. In p4 (L = 0.88; W = 0.59), the anterior ridge of the protoconid is straight. The posterior ridge is slightly convex towards the entoconid. The molars are doubled-rooted, and they are progressively smaller towards the distal side, m1 > m2 > m3. In the m1 (L = 2.04; W = 1.33), the protoconid is higher than all the lingual cusps, but similar in size to the hypoconid. The talonid basin is delimited anteriorly by the oblique cristid which ends anterolingually between the metaconid and entoconid without the development of a metastylid. The entoconid and metaconid cusps have the same height, being slightly larger than those shown in the paraconid. No talonid notch is observed. There is a well-developed entostylid at the distolingual side. In the m2 (Right: L = 1.92; W = 1.32; m2: L = 1.89; W = 1.29), the talonid basin is reduced compared with its corresponding in the m1. The paracristid in m2 is higher than in m1. The oblique cristid follows the same pattern as in m1. The metaconid is higher than the paraconid and slightly higher than the entoconid. Anteriorly, the praecingulid is narrow. The protoconid is higher than the hypoconid. The talonid notch is not observed but the entostylid is well developed as in the m1. The talonid basin in m3 (Right: L = 1.54, W = 1.07; Left: L = 1.53, W = 1.04) is smaller than that of m1 and m2. The metaconid is similar in height to the paraconid but slightly higher than the entoconid. The praecingulid is narrow and the protoconid is higher than the hypoconid as in the m2. The entostylid is not developed. Postcranial elements. The digital reconstructions show a mash of numerous rib fragments and vertebrae flattened by compaction. It has been impossible to restore any element of the postcranial axial skeleton in an objective way. Similarly, one clavicle and scapula are preserved and partially exposed. However, in digital reconstructions they are flattened and strongly fragmented, so it has been impossible to describe the original morphology of any element from the shoulder girdle. Humerus. Only the distal part of the right humerus is preserved (Fig. 6a). At the distal end of the epiphysis, the ectepicondylar (=Lateral epicondyle) and the entepicondylar (=Medial epicondyle) processes are rather well preserved but their tips are missing. In the ectepicondyle, the capitulum is laterally elongated with a fusiform shape. The surface of the entepicondyle is smaller than that of the ectepicondyle. The entepicondylar foramen is shown as a deep groove. Adjacent to this groove, a wide elliptical fossa for M. flexor digitorum profundus ligament is observed. At the posterior side of the distal end, the epicondyles are separated by a large trochlear area associated with the broadening of the humerus in which there is a small projection separating the trochlea from the fossa for M. flexor digitorum profundus ligament. This area becomes deep towards the shaft of the humerus, giving rise to the olecranon fossa. Measurements (mm): Length = 7.14; maximum distal width = 6.04; shaft thickness = 2.37; shaft width (minimum diaphysal widht) = 3.30. Radius. The right radius (Fig. 6b) is preserved in lateral connection with the distal part of the humerus. At its proximal end, the strong and stout capitular process is followed by the glenoid cavity. The glenoid cavity is deep and concave. In medial view, the ulnar articular facet of the capitular process is elongated and clearly defined. In lateral view, a conspicuous crest extends distally from the capitular process, showing a small groove for the attachment of abductor pollicis muscle. In medial view, next to this crest, a wide fossa is observed in which the radial head of abductor muscle is attached. In the distal part of the radius, the scars for tendon of extensor carpi radialis muscle forms a slight protuberance between them. The articular facets for the lunar and scaphoid are narrow and convex. Measurements (mm): Length (Glenoid cavity – articular facet) = 7.56; shaft length = 6.09; proximal width = 2.57; distal width = 2.77; maximum distal thickness = 1.56. Ulna. The right ulna (Fig. 6c) is preserved in connection with the radius and the distal end of the humerus. The proximal crest is elongated in anterior view with a widely extended area of insertion for the triceps. The decrease of the extension of this area towards its distal part forms the medial and lateral olecranon crest, which ends with the protuberance of the anconaeus process. Medial and lateral olecranon crests are elongated increasing the length of the ulna over the radius. In lateral view, a protruding proximal crest is enhanced by the depth of the abductor fossa. Towards its distal part, at the anconaeus-level, this fossa presents a small groove of the abductor scar. Below them, the coronoid process is observed with a small protuberance compared with the anconaeus. This difference in size conforms the semilunar notch. Next to the coronoid process, the radial articular facet overhangs the abductor fossa expanded towards the laterodistal side. In anterior view, the humeral articular facet is observed between the anconaeus and the coronoid process, which forms a slight depression. Below the coronoid process there is a small scar for the insertion of brachialis muscle. The area delimited by the anconoeus process and the coronoid process is the functional zone for the rotation of the humerus, namely the trochlear area. The distal part of the ulna is wide at the connection of the posterior crest and the abductor fossa with the shaft. In lateral view, the shaft ends with the styloid process, which is large with a stout and rounded terminal process. The ulnar articular facet is narrow and moderately deep ending in a stout cuneiform articular facet. Measurements (mm): Length = 13.00; olecranon length (proximal crest – anconaeus process = 4.11; mediolateral diameter = 1.30; width of proximal crest = 4.22. Carpal bones. All the carpalia are preserved (Fig. 7a). The triquetrum, hamate, centrale, trapezoid, trapezium and capitate are in connection below the third and fourth metacarpal. The lunate and the scaphoid are disconnected from the hand. In the scaphoid, the groove for the insertion of the flexor carpi radialis muscle is observed (Fig. 7b). Metacarpal bones. Four elements of the metacarpalia are preserved, the second (II) being the longest of them (Fig. 7b). The proximal prominence is rounded and stout. The outline of the phalanx articular facet is strong and it forms two protuberances. Generally, the metacarpals sustain the shape of the phalanx articular facet. Furthermore, the size of the metacarpals decreases from II to V. The proximal prominences of the third metacarpal are stouter than that of the second. The metacarpal articular facet is wide. The unciform articular facet is hardly visible. The fifth metacarpal is shorter and narrower than the rest. The unciform articular facet is strong and rounded. Measurements (mm): Mc II: L = 2.66, W = 1.45, Distal W = 1.63; Mc III: L = 2.31, W = 1.63, Distal W = 1.67; Mc IV: L = 1.57, W = 1.43, Distal W = 1.46; Mc V: L = 1.52, W = 1.37, Distal W = 1.40. Phalanges. Six phalanges are preserved (Fig. 7b). The proximal phalanges are shorter than the metacarpals. They are stout and the proximal articular facet is wide in their connection with the metacarpals. The distal articular facet is wide and rounded to the sides with a similar shape to that of the middle phalanges. The middle phalanges are shorter than the proximal phalanges, and they become narrower towards the distal end. The proximal articular facets are wide and rounded. The distal articular facets are slightly thinner than the proximal ones. Distal phalanges are long, and they flatten towards their distal ends. They have a characteristic shovel-like shape with a small notch in the extreme. The proximal articular facet is narrower than the distal articular facet of the middle phalanges. Measurements (mm): Pp III: L = 1.44, W= 1.67, Distal W = 1.54 ; Pp IV: L = 1.39 , W = 1.46, Distal W = 1.38; Mp III: L = 1.28, W= 1.54 , Distal W =1.01; Mp IV: L = 1.1, W = 1.38 , Distal W = 1.06; Dp III: L = 4.28, W= 1.01 , Distal W =0.47; Dp IV: L = 3.60, W= 1.06 , Distal W =0.30. Sesamoid bones. The proximal part of one sesamoid bone is partially preserved (Fig. 7b). The proximal articular facet of this sickle-shaped bone is wide. The distal part is stout. This element has been found below the other bones of the hand. In addition, a small accessory sesamoid bone is observed. Measurements (mm): L = 3.13; W = 1.28. Tibiofibula. The tibiofibula (Fig. 6d) is placed a few centimeters away from the rest of the connected skeleton of the animal. The tibia is thicker than the fibula and the two bones are separated by the interosseous space. Tibia and fibula are fused at their distal part, thus ending in a wide shaft. In the proximal part, the head of the fibula forms a rounded protuberance. In the tibia, the rounded medial condyle is separated from the lateral condyle by a notch that conforms the intercondyloid fossa. The lateral condyle is longer than the medial one. In the posterior part of this notch, there is a tuberosity which continues towards the distal part of the dorsomedial ridge. This ridge becomes slightly wider mediolaterally at its distal part ending with a slight depression before the medial malleolus protuberance. Next to medial malleolus, in lateral view, a tuberosity conforms the lateral malleolus. In the posterior surface of the distal part, the groove for the M. flexor digitorum tibialis is observed. A low narrow ridge of the tibiofibular shaft separates the previous groove from that for the tendon of the flexor digitorum fibularis muscle. In the distal extreme, a short and narrow crest separates the groove for the flexor digitorum tibialis from the that for the tibialis posterior tendon. The articular facet is widely extended laterally. Measurements (mm): Length = 13.97; width = 2.79; width of the distal articular facet = 2.42; interosseous space = 1.68. Additional hindlimb bones. Other than the tibiofibula, one femur, one metatarsal, and one distal phalanx are the only elements of the hindlimb preserved. However, the digital models reconstructed from the micro-CT scans are too flattened and fragmented to be considered a good approach of the original morphologies of these elements. Therefore, these bones do not provide sufficient resolution to be confidently described. Discussion Taxonomy and phylogenetic relationships The specimen from CN herein studied is, to our knowledge, the most complete Pliocene talpid skeleton ever reported in Europe. Other similar exceptionally preserved fossils of talpids are the Oligocene specimen of Geotrypus antiquus from the German locality of Enspel 26 , and the Miocene samples of Mygalea jaegeri , Proscapanus sansaniensis , and Geotrypus montisasini reported by Schwermann and Thompson 16 . Except for these privileged finds, the fossil specimens of the family Talpidae are usually restricted to isolated teeth and some characteristic postcranial elements, which make their correlation difficult. The discovery of this mole in the Pliocene site of CN turned into an exceptional landmark by the combination of unexpected characters in the only specimen recovered. The European fossil record of fossorial moles during the Pliocene was dominated by Talpa 5 . However, some other less frequent talpids have been documented in literature 11–13,17,46,47 . None of the unusual fossil forms described in these works completely fit the morphology or size of the specimen from CN. Indeed, all the Pliocene occurrences of fossorial moles in Spain have been traditionally limited to the species T. fossilis and T. minor 27 , directly ascribing all the wide and robust fossil humeri of talpids to one of these two species 48 . Actually, the specimen CN10-O17-NIV11-12 was tentatively identified as Talpa minor at first sight (see Introduction). Subsequent detailed scrutiny of the fossil resulted in the observation of an unusual configuration of the anterior lower toothrow, not typically found in the Talpini, which required a thorough phylogenetic analysis (see Material and methods). Together with our new species found, we took the chance to include some other taxa not considered in previous analyses, namely the genera Myxomygale (data taken from M. hutchisoni , M. antiqua ) and Hugueneya (data taken from H. primitiva , and Hugueneya sp. in Lopatin 49 , and the species Mongoloscapter zhegalloi , Skoczenia copernici , and Alpiscaptulus medogensis . The results of the cladistic analyses performed are shown in Fig. 8. Applying Goloboff's criterion (K=2) and equally weighted characters of different types (ordered and unordered) results in the most parsimonious tree possible (CI = 0.3910, CI excluding uninformative characters = 0.3902, HI = 0.7065, HI excluding uninformative characters = 0.6098, RI=0.6148, RC = 0.2404, G. fit = -97.02592, tree length = 862) (Fig. 8). These results are mostly in line with previous studies 2,3,14,16 , but there are some differences in the resulting evolutionary tree which deserve to be pinpointed. It is worth noting that in our analyses (Fig. 8): 1) Tegulariscaptor minor is closer to the Urotrichini than to the Uropsilinae (as proposed by Sansalone et al . 14 ), but it is phylogenetically distinct from Geotrypus (stem group to true moles); 2) Geotrypus montisasini stays separated from Geotrypus antiquus (explained by Schwermann and Thompson 16 ; 3) Eotalpa is placed in a more basal position than Uropsilus ; 4) Neurotrichus is not the sister group of Scaptonyx ; 5) Euroscaptor and Mogera are closer to Talpa than to the rest of the Talpini taxa; 6) Parascaptor is not the sister group of Scaptochirus [indeed the latter is a more derived form]; 7) Scalopus is a more derived form than Scapanus ; and 8) Mioscalops is more derived than Leptoscaptor . Similarly, the introduction of new taxa modifies the trends of previous phylogenetic trees in that: 1) Myxomygale is placed as a more basal taxon than the Desmaninae, clearly separated from the Urotrichini (in which it had been previously included by Ziegler 50 and Hugueney & Maridet 51 ); 2) Mongoloscapter , which is included in the Scaptonychini according to Lopatin 52 , shows a more derived form than those, being the sister group of Neurotrichus; 3) Skoczenia is placed in the Talpini tribe, as stated by Rzebik-Kowalska 13 , close to Scaptochirus ; and 4) Alpiscaptulus and Hugueneya are clustered with other members of the tribe Scalopini, as predicted by the works of Chen et al. 4 and Lopatin 49 . According to this analysis, Vulcanoscaptor ninoti gen. et sp. nov., must be also included in the tribe Scalopini, being closely related with Scalopus and Scapanus . The position of this new taxon in the resulting phylogenetic tree is supported by 11 characters that differentiate it from the rest of the species of the tribe Scalopini, namely: 1) the number of roots of P4 (c. 11); 2) the presence or absence of a paraconule in M2 (c.16); 3) the dimensions of the postmetacrista and preparacrista in M2 (c.20); 4) the presence or absence of a talonid notch in m1-m2 ( c.24); 5) the absence or presence of gaps in the lower premolar row (c.27); 6) the position of the protocone in P4 (c.32); 7) the length of M2 (c.34); 8) the length of M3 (c.35); 9) the length of m1 (c.44); 10) the length of m2 (c.45); and 11) the location of the posterior mental foramen in the lower mandible (c.69). These results strongly support the nature of Vulcanoscaptor ninoti gen. et sp. nov. as a new Scalopini. In this sense, the phylogenetic analysis carried out is reinforcing the results previously reached by Barrow & MacLeod 53 on the morphology of talpid mandibles or on the humeri 54,55 . Nevertheless, this Scalopini form is clearly different from any other genus in the tribe because many of them do not share the same dental formula and / or they display different configurations of their postcranial bones. More specifically, the humerus of our mole resembles that of Condylura in gross proportions, but the distal part acquires an intermediate shape between Parascalops and Scapanus , with a notch in the trochlear area less pronounced than in Parascalops (also than in Scalopoides and Scapanulus ) , but more evident than in Scapanus . Moreover, the lateral and medial epicondyles of Vulcanoscaptor gen. nov. are more robust than in Parascalops, but weaker than in Scapanus . Similarly, the proximal end of the ulna of Vulcanoscaptor ninoti gen. et sp. nov. is similar to Scalopoides in the broad area for the insertion of the triceps muscle. This morphology is also similar to that of Condylura (see Hutchinson 56 ), with the exception of the olecranon process and the articular facets, which show an intermediate morphology between Parascalops and Scapanus . Paleobiology and taphonomy Adaptation efforts in subterranean environments imply a high development of the humerus and a subsequent impact on other functional traits, so the anterior mobility of the forelimb is affected by maximizing the abduction movement 57 . According to Meier et al. 54 , such modifications are clearly reflected in an extremely short, broad and compact bone specialized for digging. The complexity of the humerus is the result from the high load that the moles must overcome with the forelimbs when digging. Supporting such intense mechanical strains need from a great development of the muscles involved, mainly those of the triceps muscular complex and their attachments sites 54,57,58 . The dimensions and the compaction of the humerus herein described, together with the strong development of the forearm, imply a huge development of the abductor muscles (i.e. teres major , pectoralis , subscapularis and latissimus dorsi ). According to Gambaryan et al. 59 , these traits are related with a major stabilization of the forearm at the elbow and wrist joints in order to maintain the humerus and the hand in the correct position for the lateral thrust. Meier et al. 54 detailed that in the fossorial clades (i.e., Talpini and Scalopini) the humeri showed a rather buckled outline and a slightly elliptic medullary cavity, reflecting the torsion of this element and the deep reaching distal end of the deltopectoral crest. Therefore, there is no doubt that Vulcanoscaptor ninoti gen. nov. et sp. was a highly specialized burrower (for further details, see extended discussion in Supplementary Information 3). The find in lacustrine sediments of a mole highly specialized in burrowing deserves some comments. It can be speculated that the presence of this specimen in the anoxic bottom of the lake could be related to a possible semi-aquatic practice other than strictly burrowing. It would be difficult to assess this from the scarce record found at CN and more evidence and analysis are needed to sustain this hypothesis. However, several authors have documented swimming abilities among talpids adapted to strictly fossorial lifestyles (e.g., Hickman 60 and references therein). The same adaptation of the humerus needed for digging could also be useful for swimming 61 . Moreover, other features described are comparable with the aquatic Condylura. The presence of a mole in the sediments of a lake suggest that this individual should have lived in the nearby area. It could have been dragged into the lake by a predator, like a bird 62 or by floods, or accidentally fell and drowned. The specimen here described is incomplete, and semi-articulated as it preserves a good anatomical connection among several skeletal elements but not within each portion ( e.g. cervical vertebras). Dispersal of the skeletal elements could have occurred after or before deposition of the remains at the anoxic bottom of the lake as suggested by the missing anatomical portions 26,62–65 . A rapid burial in the ground or in deep waters 66 , instead, would have preserved most of the anatomical connections by impeding refloat to happen after developing decomposition gasses. Moreover, the anoxic bottom of the lake also prevents major bioturbators to disturb the carcass. Complete articulated body are common in maar sites 67 , including CN 31,68 . However, this mole specimen represents an exception. The lateral position could suggest that it arrived in the final deposition site already incomplete, as similar situations suggest that moles will lay ventrally or dorsally, but not laterally due to the shoulder girdle 26,62 . This specimen could represent the remains of a scavenged carcass accidentally felt in the lake or a floated carcass that sank after decomposition gasses escaped. Lastly, post depositional processes such as faulting that affected CN could also be responsible for the differential preservation of some elements 69,70 . For further details, see extended discussion in Supplementary Information 3. Conclusions The discovery of the partial skeleton of a mole in CN enlarges the fossil record of small mammals from this locality. Comparisons of our specimen with the Pliocene and extant talpids result in its identification as a new genus and species included in the tribe Scalopini by an unusual combination of morphological characters of its dentition and postcranial elements. This is an unexpected find, considering that Scalopini moles were not frequent in Europe after the Miocene. The phylogenetic analysis performed to place this new find has resulted in a major consensus tree, which fits quite well the models previously obtained by other authors. In this sense, only a few new locations of some taxa within the phylogeny of talpids are discerned. The inclusion of the characters codified for Myxomygale , Hugueneya , Mongoloscapter , Skoczenia , and Alpiscaptulus , together with the new ones of Vulcanoscaptor gen. nov. has resulted in new positions for the genera Eotalpa , Tegulariscaptor , Geotrypus , and Neurotrichus . With respect to its adaptations, digital reconstructions display several traits of the forelimb which implies intense modifications of the forearm as a respond of the biomechanics of the humerus to achieve a great efficiency throughout digging. This is clearly evidencing that Vulcanoscaptor ninoti nov. gen et sp. was adapted to a fossorial lifestyle. The strange occurrence of this mole in the fossil site of CN is a difficult issue to tackle for which different scenarios are suggested. The disposition and the optimal preservation of the partial skeleton of the mole indicates that whatever the cause of death, the specimen probably deposited already partially disarticulated. Posteriorly, the soft tissues were decomposed promoting the disarticulation of other remains before their compaction and fragmentation. The capability of the mole for swimming whereby it would have reached the lake deliberately, or the absence of this quality, could determine whether the mole died on the shore during a regular soak or drowned after falling into the lake. The real causes of death and how the specimen arrived at lacustrine sediments remain uncertain and they will deserve further research. Material and methods The great extension of the excavation area (approximately 275,000 m 2 ) required a complex level of organization, divided into several sectors, pits, sections, layers, units and subunits. Up to date, five sectors have been excavated: Can Argilera, Can Pol, Can Cateura, Comercial and Butano sectors. To define the limits of the lake and the fossiliferous layers, several pits of about 4 or 5 meters of maximum depth were carried out with a backhoe. Their dimensions varied depending on the evidence of the potential fossil content. Subsequently, the fossil strata were manually excavated. Reference systems such as UTM (ETRS89) and Cartesian coordinates were used to position the pits and fossil occurrences. The placement of the fossil was recorded by means of a registration code (acronym of the site -CN-, year, sector, pit, excavation unit, square and number of register). Depending on the preservation of the remains and the materials in which they were included, different methods were applied to unearth them 31 . The specimen studied in the present work was found in the layer 11 of the subunit 2.3 (Fig. 3). In terms of registration, this subunit belongs to the pit 7/8 from the Can Argilera sector. The remains were preserved at the surface of green clayish sediments with fine-grained sandy admixture. The specimen was originally extracted in its matrix from the excavation with expanded polyurethane. After extraction, the remains were taken to the IPHES-CERCA laboratory for preparation. Once there, its excavation was completed using mechanical methods and the aid of a binocular lens. After delimiting the skeletal remains, the clay block and the specimen were consolidated with ethyl silicate (Estel 1000), which was drip-applied using a syringe onto the surface of the block and the skeleton. In this case, the specimen was consolidated in several sessions without saturating the sample to avoid an excess of siliceous crystallizations on the surface. Once hardened, the block was reduced and prepared for Computerized Microtomography dividing it in five fragments. Subsequently, the fragments were adhered with acrylic resin, Paraloid® B-72 dissolved in 20% acetone. Finally, a structural reintegration was carried out with a putty made from the sediment of the same block agglutinated with the same acrylic resin used for adhesion. For storage, a custom-made polyethylene-based support was made in different formats: Ethafoam® foam, Tivek® tissue and a bag. Because the remains are partially embedded, the morphological description of this mole for subsequent taxonomic identification is complicated. To avoid the extraction of the fossil, the virtual reconstruction of the remains was carried out using computerized tomography, a nondestructive technique. Five µ-CT scanners, with a 3D spatial resolution of up to 6 µm, were made with the V|Tome|X s 240 (GE Sensing & Inspections Technologies) at the CENIEH in Burgos (Spain). The set of images obtained after the scans were processed with the free and open access software ‘3D slicer’. Subsequently, and based on these models, the descriptions and measurements were made according to Hutchison 7,56 . Cladistic analysis - A total of 41 extant and extinct species have been scored based on morphological characters, both cranial and postcranial elements (Supplementary Information 4). The list of characters and taxa suggested for phylogenetic analyses was carried out based on that proposed by Schwermann et al . 3 and its preceding works 2,16,61 . When some specific information was missing, the data matrix was completed checking the characters of selected specimens of some subfamilies, tribes, genera and species (Supplementary Information 4). For details of the different species and the skeletal remains see Schwermann et al. 3 . A total of 175 characters have been considered to score the different taxa based on dentition, humerus, hand and tibiofibula. Of these discrete characters 114 are binary and 61 are multistates. Some taxa (e.g., Mioscalops isodens , Domninoides mimicus , Parascalops breweri ) show polymorphism for some characters which are named as letters codifying different states (Supplementary Information 4). In view of the absence of preserved skeletal elements of some species, characters have been scored as missing with the symbol "?". Finally, in the present work we have added some taxa to the list: 1) Tegulariscaptor minor from Sansalone 14 ; 2) Myxomygale hutchisoni sensu Klietmann et al . 71 and Myxomygale antiqua sensu Hugueney & Maridet 51 ; 3) Mongoloscapter zhegalloi according to Lopatin 52 ; 4) Skoczenia copernici sensu Rzebik-Kowalska 13 , Alpiscaptulus medogensis by Chen et al . 4 ; 5) Hugueneya sp. in Lopatin 49 and Hugueneya primitiva according to Van den Hoek Ostende 72 , and 6) the new taxon herein described, Vulcanoscaptor ninoti gen. et sp. nov. For cladistic analyses the software PAUP 4.0 73 has been used to obtain the most parsimonious tree possible. The parsimony criteria have been applied by means of a heuristic search. For further veracity of the obtained tree, Bremer values 74 were calculated in addition to the application of the scheme of implied weights from Goloboff 75 . For the latter criterion, a value of K=2 was applied to reduce the homoplasy. On the other hand, when setting character types, the criterion of equal weights and a status as unordered or non-additive has been applied to all characters. Exceptionally, 27 of that characters have been considered as ordered or additive based on Wagner's parsimony criterion due to the assumption of an ordered sequence according to their position in the symbol list 76 . Finally, in our analyses Erinaceus europaeus is considered as an outgroup following different works 2,16,61 . Anatomical abbreviations – abf: abductor fossa, abs: abductor scar, acs: accessory sesamoid bone agp: angular process, anc: ancanoeus process, brs: brachialis scar, c: oblique cristid, caf: cuneiform articular facet, cdp: condylar process, cpp: capitular process, cpt: capitulum , crp: coronoid process, dmr: dorso medial ridge, dp: distal phalanx, ef: entepicondylar foramen, end: entoconid, fdf: flexor digitorium fibularis tendon, fdt: flexor digitorium tibialis , ffd: fossa of m. flexor digitorium ligament, flc: falciform, ga: groove for abductor pollicis longus tendon, hf: head of the fibula, hff: humeral articular facet, hyd: hypoconid, hyl: hypoconulid, i: incisor, ins: interosseus space, le: lateral epicondyle, llc: lateral olecranon, lm: lateral malleolus, lrf: lunar articular facet, ltc: lateral condyle, m: molar, mc: metacarpal, md: medial phalanx, mdc: medial condyle, me: medial epicondyle, me: metacone, med: metaconid, mes: mesostyle, met: metastyle, mld: medial olecranon, mm: medial malleolus, of: olecranon fossa, p: premolar, pa: paracone, pad: paraconid, par: parastyle, pc: pectoral crest, pp: proximal phalanx, pr: protocone, prc: praecingulid, prd: protoconid, prt: protoconule, pxc: proximal crest, raf: fossa for radial head of m. abductor, rf: radial articular facet, scp: scaphoide, sf: scaphoid articular facet, sn: semilunar notch, sty: styloid process, ta: trochlear area, tmp: terminal process, tp: tibialis posterior tendon, ulf: ulnar articular facet. Institutional abbreviations – CENIEH (Centro Nacional de Investigación sobre la Evolución Humana, Burgos, Spain); IPHES (Institut de Paleoecologia Humana i Evolució Social, Tarragona, Spain). Declarations Data availability Holotype: The fossil elements of Vulcanoscaptor ninoti gen. et sp. nov. are hosted in the IPHES-CERCA with the reference: CN’10. Can Argilera sector. Pit 7/8. Layers 11. Square O17. Number 12. The 3D virtual models of Vulcanoscaptor ninoti gen. et sp. nov. are accessible for viewing in the open-source 3D repository Morphosource (https://doi.org/10.17602/M2/M614677; https://doi.org/10.17602/M2/M614683; https://doi.org/10.17602/M2/M609966). Acknowledgements The CN project is sponsored by the Caldes de Malavella town hall. Funding for this research has been provided by the Catalan Government (Generalitat de Catalunya) by means of the Departament de Cultura project CLT009/22/000043 and the research groups 2021 SGR 01238 and 2021 SGR 00127. The Spanish Government and the European Union have supported this study with the projects PID2021-122533NB-I00, PID2021-123092NB-C21 and PID2020-117289GB-I00 (Agencia Estatal de Investigación and European Regional Development Fund of the European Union, AEI/FEDER EU). The Institut Català de Paleoecologia Humana i Evolució Social (IPHES-CERCA) received financial support from the Spanish Ministry of Science and Innovation through the ‘María de Maeztu’ program for Units of Excellence (CEX2019-000945-M). The research of B.G.S., G.C., H.A.B, P.P. and M.F. is funded by the CERCA Programme/Generalitat de Catalunya. P.P. is supported by a ‘‘Ramón y Cajal” contract (grant RYC2023-044218-I) funded by MICIU/AEI/10.13039/501100011033 and “ESF+”. The CENIEH is acknowledged for providing the micro-CT scans of the remains and Josep Fortuny (ICP) and Ivan Rey-Rodríguez (UVigo-IPHES-URV) for their useful advices with the software to manage the digital reconstructions. This work is part of the Ph.D. Dissertation of the first author, in the framework of the Ph.D. Programme ‘Quaternari i Prehistòria’ of the Universitat Rovira i Virgili (Tarragona, Spain). Author contributions statement M.F. conceptualized and supervised the research; A.L.-M. and M.F. performed the descriptive part of the investigation and the writing of the original draft; A.L.-M. performed the microCT scans and data curation, the formal phylogenetic analysis, the software management, and the methodological creation of models; B.G.d.S., G.C., and J.A. worked on the resources, funding acquisition, and project administration; E.M.-R. prepared the fossil specimen and described this process; O.O. worked on the formal analysis and investigation of the geological context of the site; A.L.-M. and O.O. worked on the visualization and prepared the figures; F.G. worked on the formal analysis and investigation of the taphonomy; B.G.d.S., G.C., J.A., H.-A.B., and P.P. performed the analysis of the fossil faunal content; M.F., A.L.-M., H.-A.B. and P.P. improved later versions of the original manuscript; all the authors were involved in the discussion of the results, the review and editing of the main manuscript text. Competing interests The authors declare no competing interests. Additional Information Supplementary information is available for this paper at doi:10.34810/data1921. References Sigé, B., Crochet, J.-Y. & Insole, A. Les plus vieilles taupes. Geobios, Mem. Spec. 1 141–157. https://doi.org/10.1016/S0016-6995(77)80014-4 (1977). Hooker, J. J. Skeletal adaptations and phylogeny of the oldest mole Eotalpa (Talpidae, Lipotyphla, Mammalia) from the UK Eocene: The beginning of fossoriality in moles. Palaeontology 59 , 195–216. https://doi.org/10.1111/pala.12221 (2016). Schwermann, A. H., He, K., Peters, B. J., Plogschties, T. & Sansalone, G. Systematics and macroevolution of extant and fossil scalopine moles (Mammalia, Talpidae). Palaeontology 62 , 661–676. https://doi.org/10.1111/pala.12422 (2019). Chen, Z. Z. et al. Morphology and phylogeny of scalopine moles (Eulipotyphla: Talpidae: Scalopini) from the eastern Himalayas, with descriptions of a new genus and species. Zool J Linn Soc 193 , 432–444. https://doi.org/10.1093/zoolinnean/zlaa172 (2021). He, K., Shinohara, A., Jiang, X. L. & Campbell, K. L. Multilocus phylogeny of talpine moles (Talpini, Talpidae, Eulipotyphla) and its implications for systematics. Mol Phylogenet Evol 70 , 513–521 (2014). Hugueney, M. Les talpidés (Mammalia, Insectivora) de Coderet-Bransat (Allier) et l’évolution de cette famille au cours de l’Oligocène et du Miocène inférieur d’Europe. Travaux et Documents des Laboratoires de Géologie de Lyon 50 , 1–81. https://doi.org/10.1016/j.ympev.2013.10.002 (1972). Hutchison, J. H. Notes on type specimens of European Miocene Talpidae and a tentative classification of old world Tertiary Talpidae (Insectivora: Mammalia). Geobios 7 , 211–256. https://doi.org/10.1016/S0016-6995(74)80009-4 (1974). Ziegler, R. Talpiden (Mammalia, Insectivora) aus dem Orleanium and Astaracium Bayerns. Mitteilungen der Bayerische Staatssammlung für Paläontologie und historische Geologie 25 , 131–175 (1985). Van Cleef-Roders, J. T. & Van Den Hoek Ostende, L. W. Dental morphology of Talpa europaea and Talpa occidentalis (Mammalia: Insectivora) with a discussion of fossil Talpa in the Pleistocene of Europe. Zool. Med. Leiden 75 , 51–68 (2001). García-Alix, A., Furió, M., Minwer-Barakat, R., Martín Suárez, E. & Freudenthal, M. Environmental control on the biogeographical distribution of Desmanella (Soricomorpha, Mammalia) in the Miocene of the Iberian Peninsula. Palaeontology 54 , 753–762. https://doi.org/10.1111/j.1475-4983.2011.01062.x (2011). Cailleux, F., van den Hoek Ostende, L. W. & Joniak, P. The Late Miocene Talpidae (Eulipotyphla, Mammalia) from the Pannonian Region, Slovakia. J Paleontol 98 , 128–151. https://doi.org/10.1017/jpa.2023.95 (2024). Skoczen, S. Condylurini Dobson, 1883 (Insectivora, Mammalia) in the Pliocene of Poland. Acta Palaeontol Pol 21 , 291–313 (1976). Rzebik-Kowalska, B. Review of the Pliocene and Pleistocene Talpidae. Palaeontologia Electronica 17 , (2014). Sansalone, G., Kotsakis, T., Schwermann, A. H., Van den Hoek Ostende, L. W. & Piras, P. When moles became diggers: Tegulariscaptor gen. nov., from the early Oligocene of south Germany, and the evolution of talpid fossoriality. J Syst Palaeontol 16 , 645–657. https://doi.org/10.1080/14772019.2017.1329235 (2017). Hutterer, R. Order Soricomorpha. In:Wilson DE, Reeder DM (eds) Mammals Species of the World. A Taxonomic and Geographic Reference, 3rd edition. The Johns Hopkins University Press, Baltimore. (2005). Schwermann, A. H. & Thompson, R. S. Extraordinarily preserved talpids (Mammalia, Lipotyphla) and the evolution of fossoriality. J Vertebr Paleontol 35 . https://doi.org/10.1080/02724634.2014.934828 (2015). Sansalone, G., Kotsakis, T. & Piras, P. New Systematic Insights about Plio-Pleistocene Moles from Poland. Acta Palaeontol Pol 61 , 221–229. https://doi.org/10.4202/app.00116.2014 (2016). Storch, G. & Qiu, Z. The Neogene mammalian faunas of Ertemte and Harr Obo in Inner Mongolia (Nei Mongol), China. 2. Moles-Insectivora: Talpidae. Senckenbergiana Lethaea 64 , 89–127 (1983). Minwer-Barakat, R., García-Alix, A., Martín-Suárez, E. & Freudenthal, M. Early Pliocene Desmaninae (Mammalia, Talpidae) from Southern Spain and the Origin of the Genus Desmana. J Vertebr Paleontol 40 . https://doi.org/10.1080/02724634.2020.1835936 (2020). Rümke, C. G. A review of fossil and recent Desmaninae (Talpidae, Insectivora). Utrecht Micropaleontological Bulletins Special Publication 4 , 1–241 (1985). Sansalone, G. et al. Impact of transition to a subterranean lifestyle on morphological disparity and integration in talpid moles (Mammalia, Talpidae). BMC Evol Biol 19 , (2019). Doukas, C. S., L.W., V. den H. O., C.D., T. & J.W.F., R. The vertebrate locality Maramena (Macedonia, Greece) at the Turolian- Ruscinian Boundary (Neogene). Münchner Geowissenschaftliche Abhandlungen (A) 28 , 43–64 (1995). Sansalone, G., Kotsakis, T. & Piras, P. Talpa fossilis or Talpa europaea? Using geometric morphometrics and allometric trajectories of humeral moles remains from Hungary to answer a taxonomic debate. Palaeontologia Electronica 18 , 1–17. https://doi.org/10.26879/560 (2015). Hoek Ostende, L. W. Van Den & Fejfar Oldrich. Erinaceidae and Talpidae (Erinaceomorpha , Soricomor- pha , Mammalia) from the Lower Miocene of Merkur-Nord (Czech Republic , MN 3). Beiträge zur Paläontologie 30 , 175–203 (2006). Kimmig, J., Schiffbauer, J. D. A modern definition of Fossil-Lagerstätten. Trends Ecol. 39 , 621–624 (2024). Schwermann, A. H. & Martin, T. A partial skeleton of Geotrypus antiquus (Talpidae, Mammalia) from the Late Oligocene of the Enspel fossillagerstätte in Germany. Palaontol Z 86 , 409–439 (2012). Furió, M., van den Hoek Ostende, L. W., Agustí, J. & Minwer-Barakat, R. Evolución de las asociaciones de insectívoros (Eulipotyphla, Mammalia) en España y su relación con los cambios climáticos del Neógeno y el Cuaternario. Ecosistemas 27 , 38–51. https://doi.org/10.7818/ECOS.1454 (2018). Jiménez-Moreno, G. et al. Late Pliocene vegetation and orbital-scale climate changes from the western Mediterranean area. Glob Planet Change 108 , 15–28. https://doi.org/10.1016/j.gloplacha.2013.05.012 (2013). Vicente, J. Troballa d’un Leptobos a Caldes de Malavella (La Selva). Societat d’Història Natural, Butlletí del Centre d’Estudis de la Natura del Barcelonès Nord 1 , 86–88 (1985). Vehí, M., Pujadas, A., Roqué, C. & Buxó, L. P. Un edifici volcànic inèdit a Caldes de Malavella (la Selva, Girona): El volcà del Camp dels Ninots. Quaderns de la Selva 11 , 45–67 (1999). Gómez de Soler, B. et al. A new key locality for the Pliocene vertebrate record of Europe: The Camp dels Ninots maar (NE Spain). Geologica Acta (2012) doi:10.1344/105.000001702. Claude, J., De Soler, B. G., Campeny, G., Agusti, J. & Oms, O. Presence of a chelydrid turtle in the late Pliocene Camp dels Ninots locality (Spain). Bulletin de la Societe Geologique de France 185 , 253–256. https://doi.org/10.2113/gssgfbull.185.4.253 (2014). Přikryl, T. et al. Fish fauna of the Camp dels Ninots locality (Pliocene; Caldes de Malavella, province of Girona, Spain) – first results with notes on palaeoecology and taphonomy. Hist Biol 28 , 347–357. https://doi.org/10.1080/08912963.2014.934820 (2016). Blain, H. et al. Water frogs (Anura, Ranidae) from the Pliocene Camp dels Ninots Konservat-Lagerstätte (Caldes de Malavella, NE Spain). Abstracts volume 7th International Maar Conference - Olot, Catalonia, Spain. 162–163 (2018). Robles, S., Barrón, E. & Cebolla, C. Estudio paleobotánico preliminar del afloramiento plioceno de Camp dels Ninots (Caldes de Malavella, Girona, España). Macroflora del sector de Can Argilera. Boletín de la Real Sociedad Española de Historia Natural, Sección Geológica. 17 , 75–89 (2013). Oms, O. et al. Early lake sedimentation in the Pliocene Camp dels Ninots maar (Catalan Coastal Ranges, Spain). Abstracts volume 7th International Maar Conference - Olot, Catalonia, Spain 172-173. 172–173 (2018). Martí, J., Mitjavila, J., Roca, E. & Aparicio, A. Cenozoic magmatism of the valencia trough (western mediterranean): Relationship between structural evolution and volcanism. Tectonophysics 203 , 145–165. https://doi.org/10.1016/0040-1951(92)90221-Q (1992). Roca, E., Sans, M., Cabrera, L. & Marzo, M. Oligocene to Middle Miocene evolution of the central Catalan margin (northwestern Mediterranean). Tectonophysics 315 , 209–229 (1999). Tassone, A., Roca, E., Munoz, J. A., Cabrera, L. & Canals, M. Evolucion del sector septentrional del margen continental catalan durante el Cenozoico. Acta Geologica Hispanica 29 , 3–37 (1994). Guardia, P. Volcans tertiaires et quaternaires de la province de Gerona et paléomagnétisme de leurs coulées. (1964). Donville, B. Ages potassium-argon des roches volcaniques de la Depression de la Selva (nord-est de l’espagne). (1973). Oms, O. et al. Structure of the Pliocene Camp dels Ninots maar-diatreme (Catalan Volcanic Zone, NE Spain). Bull Volcanol 77 , (2015). Lindner, H., Gabriel, G., Götze, H.-J., Kaeppler, R. & Suhr, P. Geophysical and geological investigation of maar structures in the Upper Lusatia region (East Saxony). Zeitschrift der Deutschen Gesellschaft für Geowissenschaften 157 , 355–372 (2006). Pirrung, M. et al. Lithofacies succession of maar crater deposits in the Eifel area (Germany). Terra Nova 15 , 125–132. https://doi.org/10.1046/j.1365-3121.2003.00473.x (2003). Rodríguez-Salgado, P. et al. Mineralogical proxies of a Pliocene maar lake recording changes in precipitation at the Camp dels Ninots (Pliocene, NE Iberia). Sediment Geol 418 , 105910. https://doi.org/10.1016/j.sedgeo.2021.105910 (2021). Skoczen, S. Scaptonychini Van Valen, 1967, Urotrichini and Scalopini Dobson, 1883 (Insectivora, Mammalia) in the Pliocene and Pleistocene of Poland. Acta Zoologica Cracoviensia 24 , 411–448 (1980). Skoczen, S. New records of Parascalops, Neurotrichus and Condylura (Talpinae, Insectivora) from the Pliocene of Poland. Acta Theriol (Warsz) 38 , 125–137 (1993). Furió, M. & Angelone, C. Insectivores (Erinaceidae, Soricidae, Talpidae; Mammalia) from the Pliocene of Capo Mannu D1 (Mandriola, central-western Sardinia, Italy). Neues Jahrb Geol Palaontol Abh 258 , 229–242 (2010). Lopatin, A. V. Early Miocene Small Mammals from the North Aral Region (Kazakhstan) with Special Reference to their Biostratigraphic Significance. Paleontological journal 38 , 217–323 (2004). Ziegler, R. Moles (Talpidae) from the late Middle Miocene of South Germany. Acta Palaeontol Pol 48 , 617–648 (2003). Hugueney, M. & Maridet, O. Evolution of Oligo-Miocene talpids (Mammalia, Talpidae) in Europe: focus on the genera Myxomygale and Percymygale n. gen. Hist Biol 30 , 267–275. https://doi.org/10.1080/08912963.2017.1282477 (2018). Lopatin, A. V. An oligocene mole (Talpidae, Insectivora, Mammalia) from Mongolia. Paleontologicheskii Zhurnal 36 , 91–92 (2002). Barrow, E. & MacLeod, N. Shape variation in the mole dentary (Talpidae: Mammalia). Zool J Linn Soc 153 , 187–211. https://doi.org/10.1111/j.1096-3642.2008.00376.x (2008). Meier, P. S., Bickelmann, C., Scheyer, T. M., Koyabu, D. & Sánchez-Villagra, M. R. Evolution of bone compactness in extant and extinct moles (Talpidae): Exploring humeral microstructure in small fossorial mammals. BMC Evol Biol 13 , (2013). Sansalone, G. et al. Influence of Evolutionary Allometry on Rates of Morphological Evolution and Disparity in strictly Subterranean Moles (Talpinae, Talpidae, Lipotyphla, Mammalia). J Mamm Evol 25 , 1–14 (2018). Hutchison, J. H. Fossil Talpidae (lnsectivora, Mammalia) from the later Tertiary of Oregon. Bulletin of the Museum of Natural History 1–117 (1968). Sansalone, G. et al. Decoupling Functional and Morphological Convergence, the Study Case of Fossorial Mammalia. Front Earth Sci (Lausanne) 8 , 112 (2020). Furio, M. The shrew pleads ‘not guilty’ to the mole’s murder: comment on Bennàsar et al. (2015). Hist Biol 29 , 230-233. https://doi.org/10.1080/08912963.2016.1151016 (2017). Gambaryan, P. P., Gasc, J.-P. & Renous, S. Cinefluorographical study of the burrowing movements in the common mole, Talpa europaea (Lipotyphla, Talpidae). Russ J Theriol 1 , 91–109 (2002). Hickman, G. C. Swimming ability of talpid moles, with particular reference to the semi-aquatic Condylura cristata. Mammalia 48 , 505–514. https://doi.org/10.1515/mamm.1984.48.4.505 (1984). Sánchez-Villagra, M. R., Horovitz, I. & Motokawa, M. A comprehensive morphological analysis of talpid moles (Mammalia) phylogenetic relationships. Cladistics 22 , 59–88. https://doi.org/10.1111/j.1096-0031.2006.00087.x (2006). Mähler, B., Schwermann, A. H., Wuttke, M., Schultz, J. A. & Martin, T. Four-dimensional virtopsy and the taphonomy of a mole from the Oligocene of Lake Enspel (Germany). Paleobiodivers Paleoenviron 95 , 115–131 (2015). Behrensmeyer, A. K. Terrestrial vertebrate accumulations. In: Allison, P.A., Briggs, D.E.G. (Eds.), Taphonomy: Releasing the Data Locked in the Fossil Record. Taphonomy: Releasing the Data Locked in the Fossil Record. Plenum Press, New York 291–335 (1991). Sabol, M. et al. Early Late Pliocene site of Hajnáčka I (Southern Slovakia) - Geology, palaeovolcanic evolution, fossil assemblages and palaeoenvironment. CFS Courier Forschungsinstitut Senckenberg 261–274 (2006). Syme, C.E., Salisbury, S. W. Patterns of aquatic decay and disarticulation in juvenile Indo-Pacific crocodiles (Crocodylus porosus), and implications for the taphonomic interpretation of fossil crocodyliform material. Palaeogeogr Palaeoclimatol Palaeoecol 412 , 108–123 (2014). Moore, M., J., Mitchell, G. H., Rowles, T., K., Early, G. Dead Cetacean? Beach, Bloat, Float, Sink. Front. Mar. Sci. 7 . https://doi.org/10.3389/fmars.2020.00333 (2020). Uhl, D., Wuttke, M., Aiglstorfer, M., Gee, C.T., Grandi, F., Höltke, O., Kaiser, T.M., Kaulfuss, U., Lee, D., Lehmann, T., Oms, O., Poschmann, M.J., Rasser, M.J., Schindler, T., Smith, K.T., Suhr, P., Wappler, T., Wedmann, S. Deep‑time maar lakes and other volcanogenic lakes as Fossil‑Lagerstatten – An overview. Paleobiodivers Paleoenviron 104 , 763–848 (2024). Campeny Vall-Llosera, G. & Gómez de Soler, B. El Camp Dels Ninots: Rastres de l’Evolució . El Camp dels Ninots: Rastres de l’evolució (Cambridge University Press, Caldes de Malavella, 2010). Bolós, X., Oms, O., Rodríguez-Salgado, P., Martí, J., Gómez De Soler, B., Campeny, G. Eruptive evolution and 3D geological modeling of Camp dels Ninots maar-diatreme (Catalonia) through continuous intra-crater drill coring. J. Volcanol. Geotherm. Res. 419, 107369. 419 . https://doi.org/10.1016/j.jvolgeores.2021.107369 (2021). Grandi, F., Del Valle, H., Cáceres, I., Rodríguez-Salgado, P., Oms, O., Fernández-Jalvo, Y., García, F., Campeny, G & Gómez de Soler, B. Exceptional preservation of large fossil vertebrates in a volcanic setting (Camp dels Ninots, Spain). Hist. Biol . 35 , 1234-1249. https://doi.org/10.1080/08912963.2022.2085570 (2023). Klietmann, J., Nagel, D., Rummel, M. & van den Hoek Ostende, L. W. A gap in digging: the Talpidae of Petersbuch 28 (Germany, Early Miocene). Palaontol Z 89 , 563–592 (2015). Van den Hoek Ostende, L. W. The Talpidae (Insectivora, Mammalia) of Eggingen-Mittelhart (Baden-Wurttenberg, FRG) with special reference to the Paratalpa-Desmanodan lineage. Preprint at (1988). Swofford, D. L. PAUP*. Phylogenetic Analysis Using Parsimony (*And Other Methods), version 4. Sinauer Associates, Sunderland, Massachusetts. Version 4 , (2002). Bremer, K. R. Branch support and tree stability. Cladistics 10 , 295–304 (1994). Goloboff, P. A. Estimating character weights during tree search. Cladistics 9 , 83–91 (1993). Swofford, D. L., & Maddison, W. P. Reconstructing ancestral character states under Wagner parsimony. Math Biosci 87 , 199–229 (1987). Additional Declarations No competing interests reported. Supplementary Files LinaresetalSUPPLEMENTARYINFORMATION1.docx LinaresetalSUPPLEMENTARYINFORMATION2.docx LinaresetalSUPPLEMENTARYINFORMATION3.docx LinaresetalSUPPLEMENTARYINFORMATION4.docx Cite Share Download PDF Status: Published Journal Publication published 10 Jul, 2025 Read the published version in Scientific Reports → Version 1 posted Editorial decision: Revision requested 17 Apr, 2025 Reviews received at journal 14 Apr, 2025 Reviews received at journal 06 Apr, 2025 Reviewers agreed at journal 28 Mar, 2025 Reviewers agreed at journal 28 Mar, 2025 Reviewers invited by journal 28 Mar, 2025 Editor assigned by journal 28 Mar, 2025 Editor invited by journal 19 Mar, 2025 Submission checks completed at journal 19 Mar, 2025 First submitted to journal 12 Mar, 2025 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-6215385","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Article","associatedPublications":[],"authors":[{"id":436510987,"identity":"8f1d98e8-5dcb-4665-879a-7c9216f41d89","order_by":0,"name":"Adriana Linares-Martín","email":"","orcid":"","institution":"Institut Català de Paleoecologia Humana i Evolució Social (IPHES-CERCA)","correspondingAuthor":false,"prefix":"","firstName":"Adriana","middleName":"","lastName":"Linares-Martín","suffix":""},{"id":436510988,"identity":"4a6dfd68-980f-4782-b710-5a0efd6fa5a8","order_by":1,"name":"Marc Furió","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAAyklEQVRIiWNgGAWjYNACGwYGNgbmAwyMDURrSQNpYUuAaGEjVgsDA48BcVp0288+/MCQcDifj73nm8TPHQzy/PMJuM7sTLqxBFCLZRvP2W2SvWcYDGccI2CL2YE0NgbGH4cN2CRyt0nwtjEkMBDUcv4Z0O0JIC05zyT/ArXIE9RyIw2uhU0aZIsBYS3PmCUSEtIN2HiOGVvLnpEw3HgsgZDD0hg/fEiwNpBvb3548+0OG3m5wwcIWAMCSMZKEKF8FIyCUTAKRgFBAAB+eDveYuW6iQAAAABJRU5ErkJggg==","orcid":"","institution":"Universitat Autònoma de Barcelona (UAB)","correspondingAuthor":true,"prefix":"","firstName":"Marc","middleName":"","lastName":"Furió","suffix":""},{"id":436510989,"identity":"473cca36-cb07-44ae-9442-af6272d0d3fb","order_by":2,"name":"Bruno Gómez Soler","email":"","orcid":"","institution":"Institut Català de Paleoecologia Humana i Evolució Social (IPHES-CERCA)","correspondingAuthor":false,"prefix":"","firstName":"Bruno","middleName":"Gómez","lastName":"Soler","suffix":""},{"id":436510990,"identity":"3df1ce27-5fab-4eab-b434-cd1994e506d3","order_by":3,"name":"Jordi Agustí","email":"","orcid":"","institution":"Institut Català de Paleoecologia Humana i Evolució Social (IPHES-CERCA)","correspondingAuthor":false,"prefix":"","firstName":"Jordi","middleName":"","lastName":"Agustí","suffix":""},{"id":436510991,"identity":"b835e79d-b502-4b35-811f-b4c335942ad8","order_by":4,"name":"Oriol Oms","email":"","orcid":"","institution":"Universitat Autònoma de Barcelona (UAB)","correspondingAuthor":false,"prefix":"","firstName":"Oriol","middleName":"","lastName":"Oms","suffix":""},{"id":436510992,"identity":"1e7a223e-1978-4fc4-aa49-ea8be733d7a2","order_by":5,"name":"Federica Grandi","email":"","orcid":"","institution":"Institut Català de Paleoecologia Humana i Evolució Social (IPHES-CERCA)","correspondingAuthor":false,"prefix":"","firstName":"Federica","middleName":"","lastName":"Grandi","suffix":""},{"id":436510993,"identity":"41ae29ee-effd-4cb9-8ced-9cc6ec281fd7","order_by":6,"name":"Hugues-Alexandre Blain","email":"","orcid":"","institution":"Institut Català de Paleoecologia Humana i Evolució Social (IPHES-CERCA)","correspondingAuthor":false,"prefix":"","firstName":"Hugues-Alexandre","middleName":"","lastName":"Blain","suffix":""},{"id":436510994,"identity":"0752f7ac-e795-476a-af16-d824de5894b5","order_by":7,"name":"Elena Moreno-Ribas","email":"","orcid":"","institution":"Institut Català de Paleoecologia Humana i Evolució Social (IPHES-CERCA)","correspondingAuthor":false,"prefix":"","firstName":"Elena","middleName":"","lastName":"Moreno-Ribas","suffix":""},{"id":436510996,"identity":"5180d643-f0fc-4c0d-bb7c-b896e1fa5226","order_by":8,"name":"Pedro Piñero","email":"","orcid":"","institution":"Universitat de València","correspondingAuthor":false,"prefix":"","firstName":"Pedro","middleName":"","lastName":"Piñero","suffix":""},{"id":436510999,"identity":"ece00d85-eed7-4025-b189-38a38ff6ab42","order_by":9,"name":"Gerard Campeny","email":"","orcid":"","institution":"Institut Català de Paleoecologia Humana i Evolució Social (IPHES-CERCA)","correspondingAuthor":false,"prefix":"","firstName":"Gerard","middleName":"","lastName":"Campeny","suffix":""}],"badges":[],"createdAt":"2025-03-13 01:08:08","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6215385/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6215385/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1038/s41598-025-10396-1","type":"published","date":"2025-07-10T15:57:46+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":80045996,"identity":"50d72615-1162-479c-8dca-452723da4ecc","added_by":"auto","created_at":"2025-04-07 09:47:32","extension":"jpg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":3031698,"visible":true,"origin":"","legend":"\u003cp\u003eGeographical and geological context of the locality of CN and Can Argilera site. LSB: La Selva Basin (Adapted from Gómez De Soler \u003cem\u003eet al.\u003c/em\u003e 2012).\u003c/p\u003e","description":"","filename":"LinaresetalFIGURE01.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6215385/v1/115ee5e993aa81f15d411b9b.jpg"},{"id":80046001,"identity":"a367c846-f90b-4ff4-9fb3-253d5d9f8bf8","added_by":"auto","created_at":"2025-04-07 09:47:32","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":3502814,"visible":true,"origin":"","legend":"\u003cp\u003eStratigraphic section of Can Argilera (Adapted from Gómez de Soler \u003cem\u003eet al.\u003c/em\u003e2012).\u003c/p\u003e","description":"","filename":"LinaresetalFIGURE02.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6215385/v1/61199aeba6f25264fa6d2586.jpg"},{"id":80046027,"identity":"6db68c51-7b87-47d2-b7cc-3f5f4807f476","added_by":"auto","created_at":"2025-04-07 09:47:33","extension":"jpg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":20824384,"visible":true,"origin":"","legend":"\u003cp\u003eExcavation area of Can Argilera in CN (Pit 7/8). The exact place where the fossil specimen was found is indicated with a black spot.\u003c/p\u003e","description":"","filename":"LinaresetalFIGURE03.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6215385/v1/72915210cd320b70bdad7ebb.jpg"},{"id":80047502,"identity":"4684c71d-318c-4d2b-a89f-aa76e4913c3c","added_by":"auto","created_at":"2025-04-07 09:55:32","extension":"jpg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":3420656,"visible":true,"origin":"","legend":"\u003cp\u003ePartial skeleton of \u003cem\u003eVulcanoscaptor ninoti\u003c/em\u003e gen. et sp. nov. (CN10-O17-NIV11-12) from CN. Cranium (1), skull (2), mandibles (3), ulna (4), carpals (5,12,13), metacarpals (6,14), proximal phalanges (7), medial phalanges (8), distal phalanges (9,20), sesamoid bone (11,21), accessory sesamoid bone (10), radius (15), humerus (16), ribs (17,23,24), clavicle (18), scapula (19), vertebrae (22), tibiofibula (25), metatarsal (26), foot distal phalange (27), femur (28). Scale bar 10 cm.\u003c/p\u003e","description":"","filename":"LinaresetalFIGURE04.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6215385/v1/d07923133747564c2c7e23c8.jpg"},{"id":80047498,"identity":"923370b5-1168-40c5-af47-826e34997652","added_by":"auto","created_at":"2025-04-07 09:55:32","extension":"jpg","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":807269,"visible":true,"origin":"","legend":"\u003cp\u003eDigital 3D reconstruction of the dentognathic elements of \u003cem\u003eVulcanoscaptor ninoti\u003c/em\u003e gen. et sp. nov. (CN10-O17-NIV11-12). \u003cstrong\u003ea. \u003c/strong\u003eLeft upper tooth-row (I3 + P2-M3) in occlusal view. \u003cstrong\u003eb.\u003c/strong\u003e Right lower tooth-row (m3-i2) in occlusal view. \u003cstrong\u003ec\u003c/strong\u003e. Righthemimandible in lateral view. Scale bar equals 1 mm in \u003cstrong\u003ea\u003c/strong\u003e and \u003cstrong\u003eb\u003c/strong\u003e, and 5 mm in \u003cstrong\u003ec\u003c/strong\u003e.\u003c/p\u003e","description":"","filename":"LinaresetalFIGURE05.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6215385/v1/ff994e1fe73d44db9ac83be7.jpg"},{"id":80047504,"identity":"329d521d-a096-48f4-8b5e-f33afa3d5074","added_by":"auto","created_at":"2025-04-07 09:55:32","extension":"jpg","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":2660478,"visible":true,"origin":"","legend":"\u003cp\u003eDigital 3D reconstruction of some forelimb elements of \u003cem\u003eVulcanoscaptor ninoti\u003c/em\u003e gen. et sp. nov. (CN10-O17-NIV11-12). \u003cstrong\u003ea. \u003c/strong\u003eRight humerus; a1 posterior view; a2 anterior view. \u003cstrong\u003eb.\u003c/strong\u003e Right radius: b1 lateral view; b2 proximal-medial view; b3 distal view. \u003cstrong\u003ec.\u003c/strong\u003e Right ulna; c1 left anterior view; c2 lateral view. \u003cstrong\u003ed.\u003c/strong\u003eRight tibiofibula; d1 anterior view; d2 posterior view. Scale bars at the corners equal 1 mm in all cases.\u003c/p\u003e","description":"","filename":"LinaresetalFIGURE06.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6215385/v1/6f593cea838653a348f880c4.jpg"},{"id":80047500,"identity":"42f740d1-f659-4dbb-a8b4-db9ea45aa937","added_by":"auto","created_at":"2025-04-07 09:55:32","extension":"jpg","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":5036562,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003ea. \u003c/strong\u003eDigital reconstruction of the right forelimb of \u003cem\u003eVulcanoscaptor ninoti\u003c/em\u003e gen. et sp. nov. (CN10-O17-NIV11-12) in its original position. \u003cstrong\u003eb.\u003c/strong\u003e Detail of the right manus articulation in volar view. Carpals, metacarpals and phalanges are highlighted using different colors: triquetrum (dark blue), hamate (purple), capitate (pink), centrale (light blue), trapezoid (white), trapezium (red), metacarpals (orange), proximal phalanges (yellow), medial phalanges (light green), distal phalanges (dark green). Falciform, accessory sesamoid, lunate and scaphoid are disconnected from the hand. Scale bar equals 5 mm in \u003cstrong\u003ea\u003c/strong\u003e and1 mm in \u003cstrong\u003eb\u003c/strong\u003e.\u003c/p\u003e","description":"","filename":"LinaresetalFIGURE07.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6215385/v1/5401dedfa01d19355d15bd34.jpg"},{"id":80047499,"identity":"8c8a4619-962e-4b64-a095-47d56dbcca4c","added_by":"auto","created_at":"2025-04-07 09:55:32","extension":"jpg","order_by":8,"title":"Figure 8","display":"","copyAsset":false,"role":"figure","size":389607,"visible":true,"origin":"","legend":"\u003cp\u003eStrict consensus of the most parsimonious phylogenetic tree for the Talpidae (see Material and methods section) including \u003cem\u003eVulcanoscaptor\u003c/em\u003e gen. nov.\u003c/p\u003e","description":"","filename":"LinaresetalFIGURE08.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6215385/v1/9d2dc26b51b69bb1a99df4b8.jpg"},{"id":86699450,"identity":"7cce03d7-9356-481e-88cb-26e1cd7dde42","added_by":"auto","created_at":"2025-07-14 16:09:51","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":40828689,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6215385/v1/3d7bda31-9a50-4320-8fbe-04a25ef3f6e6.pdf"},{"id":80045990,"identity":"0909e51c-a6c6-48be-bce6-517e9d34837d","added_by":"auto","created_at":"2025-04-07 09:47:32","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":34903,"visible":true,"origin":"","legend":"","description":"","filename":"LinaresetalSUPPLEMENTARYINFORMATION1.docx","url":"https://assets-eu.researchsquare.com/files/rs-6215385/v1/cd7ebe5c965090147b4f351c.docx"},{"id":80045993,"identity":"dcd8134e-bb34-41dd-8f89-242da6ed65a2","added_by":"auto","created_at":"2025-04-07 09:47:32","extension":"docx","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":70730,"visible":true,"origin":"","legend":"","description":"","filename":"LinaresetalSUPPLEMENTARYINFORMATION2.docx","url":"https://assets-eu.researchsquare.com/files/rs-6215385/v1/20d077ac7f07c7c7c854a91b.docx"},{"id":80045991,"identity":"a68296a1-9519-4832-b611-04e70fe7cb2b","added_by":"auto","created_at":"2025-04-07 09:47:32","extension":"docx","order_by":3,"title":"","display":"","copyAsset":false,"role":"supplement","size":33562,"visible":true,"origin":"","legend":"","description":"","filename":"LinaresetalSUPPLEMENTARYINFORMATION3.docx","url":"https://assets-eu.researchsquare.com/files/rs-6215385/v1/b9eb56a11165902dae24f3ef.docx"},{"id":80045995,"identity":"c1364cbd-0307-4d94-abc3-c7f2e69810f8","added_by":"auto","created_at":"2025-04-07 09:47:32","extension":"docx","order_by":4,"title":"","display":"","copyAsset":false,"role":"supplement","size":52561,"visible":true,"origin":"","legend":"","description":"","filename":"LinaresetalSUPPLEMENTARYINFORMATION4.docx","url":"https://assets-eu.researchsquare.com/files/rs-6215385/v1/1fe55994a08106f86c965ae2.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"An unexpected New World mole (Scalopini, Mammalia) from the Pliocene of Europe sheds light on the phylogeny of talpids","fulltext":[{"header":"Introduction","content":"\u003cp\u003eThe Talpidae are probably one of the most intriguing families of mammals from the paleobiogeographic point of view. Only a few species within this group can be considered truly cursorial, because most of them show in some extent adaptations to fossorial activity. Indeed, many talpids are extremely adapted to subterranean and aquatic lifestyles, and their subaerial movements or out of water displacements are rather clumsy. Out of tunnels or rivers, most species of moles are highly vulnerable to potential predators. Thus, anyone would expect these animals to follow a rather predictable and constant (paleo-)biogeographic evolutionary history, with no significant migrations, considering all the possible physical and biological barriers that could constrain their distribution. Surprisingly, the fossil record is rather indicating a quite different story.\u003c/p\u003e \u003cp\u003eThe genus \u003cem\u003eEotalpa\u003c/em\u003e is the oldest fossorial representative of the family known hitherto\u003csup\u003e\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e,\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e\u003c/sup\u003e. According to the fossil record of the younger representatives, few groups have originated and stayed in the same region that they currently inhabit. For instance, Schwermann \u003cem\u003eet al\u003c/em\u003e.\u003csup\u003e\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e3\u003c/span\u003e\u003c/sup\u003e conducted a thorough study on the evolutionary history of the Scalopini, concluding that numerous transcontinental colonization events occurred between Asia, Europe, and North America, to the extent that no unambiguous location for the origin of the group could be identified. The recent discovery of \u003cem\u003eAlpiscaptulus medogensis\u003c/em\u003e, a living species of Scalopini from the Himalayas\u003csup\u003e\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e4\u003c/span\u003e\u003c/sup\u003e, has just added some more questions to this enigma.\u003c/p\u003e \u003cp\u003eIn a similar way, uropsilines are nowadays represented only by the genus \u003cem\u003eUropsilus\u003c/em\u003e (but see disagreements in He \u003cem\u003eet al.\u003c/em\u003e\u003csup\u003e\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e5\u003c/span\u003e\u003c/sup\u003e) and restricted to Central and Southeastern Asia. The fossil record shows, however, that its Oligocene, Miocene and Pliocene relatives (\u003cem\u003eDesmanella\u003c/em\u003e, \u003cem\u003eMystipterus\u003c/em\u003e, \u003cem\u003eAsthenoscapter\u003c/em\u003e, \u003cem\u003eMygatalpa\u003c/em\u003e, \u003cem\u003eTheratiskos\u003c/em\u003e) lived in European, Minor Asian and North American lands\u003csup\u003e\u003cspan additionalcitationids=\"CR7 CR8 CR9 CR10\" citationid=\"CR6\" class=\"CitationRef\"\u003e6\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e11\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eAnother example is found in the tribe condylurini, nowadays being solely represented by the endemic North American star-nosed mole, \u003cem\u003eCondylura cristata\u003c/em\u003e. However, unequivocal representatives of the genus have been found in the Pliocene of Poland\u003csup\u003e\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e,\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e13\u003c/span\u003e\u003c/sup\u003e and the Middle Miocene of Kazakhstan\u003csup\u003e\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e\u003c/sup\u003e. Likewise, \u003cem\u003eNeurotrichus gibbsii\u003c/em\u003e is the only living representative of the tribe Neurotrichini, according to Hutterer\u003csup\u003e\u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e\u003c/sup\u003e. The hypothesis of a North American origin of \u003cem\u003eNeurotrichus\u003c/em\u003e judging by its solely current occurrence in this continent would apparently be parsimonious. Nonetheless, the cladistic analysis performed by Schwermann and Thompson\u003csup\u003e\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e\u003c/sup\u003e suggested that this genus could find its closest living relative in the genus \u003cem\u003eScaptonyx\u003c/em\u003e, which mostly currently inhabits elevated areas from China. The extinct genus \u003cem\u003eRzebikia\u003c/em\u003e is a very similar form to \u003cem\u003eNeurotrichus\u003c/em\u003e, but it is only found in Plio-Pleistocene sites from Europe\u003csup\u003e\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e17\u003c/span\u003e\u003c/sup\u003e. Both genera, \u003cem\u003eNeurotrichus\u003c/em\u003e and \u003cem\u003eRzebikia\u003c/em\u003e are putative descendants of an Asian form, \u003cem\u003eQuyania\u003c/em\u003e\u003csup\u003e\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e\u003c/sup\u003e. Far from being simple, the most likely explanation to such distribution according to Sansalone \u003cem\u003eet al.\u003c/em\u003e\u003csup\u003e\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e17\u003c/span\u003e\u003c/sup\u003e is that the original Asian stock of moles finally derived into \u003cem\u003eN. gibbsii\u003c/em\u003e after colonizing North America in the Early Pliocene, while another one or two migration waves towards Eastern Europe could have resulted in several European forms.\u003c/p\u003e \u003cp\u003eThe case of the swimming forms of the Desmanini is similarly puzzling. The Russian desman (\u003cem\u003eDesmana moschata\u003c/em\u003e), an extant endemic species living in Russia, Ukraine and Kazakhstan, is the only survivor of a genus that apparently originated in the south of the Iberian Peninsula\u003csup\u003e\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e\u003c/sup\u003e. Ironically, the Iberian desman (\u003cem\u003eGalemys pyrenaicus\u003c/em\u003e), an endemic form that currently inhabits some stream headwaters in this part of the European continent, belongs to a genus of unknown geographic origin. During the Late Neogene, the tribe had a much wider distribution, although restricted to Europe and parts of Asia Minor\u003csup\u003e\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eSome of these apparent inconsistencies are probably rooted in a very limited view of the talpid fossil record. On the one hand, moles are rather conservative in their morphology, likely due to the constant conditions where they thrive. It could be that once acquired an efficient capacity for digging (or swimming), there were few possible morphological modifications to adapt to different lifestyles\u003csup\u003e\u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e21\u003c/span\u003e\u003c/sup\u003e. On the other hand, the fossil finds of talpids usually correspond to disarticulated and/or incomplete elements, not always easy to identify to the species level. In many cases, it is difficult to discriminate whether the (little) variation in size and morphology observed in the fossil assemblages is due to interspecific or intraspecific variability. A good example is found on the uncertainty of how many species of \u003cem\u003eTalpa\u003c/em\u003e occur in the European fossil record, a debate that has been extended for decades in specialized literature\u003csup\u003e\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e,\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e,\u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e23\u003c/span\u003e\u003c/sup\u003e. For a comprehensive list of approaches to link humeri and dentitions in fossil talpids, the reader is referred to van den Hoek Ostende and Fejfar\u003csup\u003e\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e24\u003c/span\u003e\u003c/sup\u003e.\u003cdiv class=\"BlockQuote\"\u003e\u003cp\u003eFortunately, the fossil record is not always limited to patchy finds and fragmentary elements. In the last years, some fossil-Lagerst\u0026auml;tten\u003csup\u003e\u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e25\u003c/span\u003e\u003c/sup\u003e sites have provided several exceptional specimens with many skeletal elements that can be clearly ascribed to the same individual\u003csup\u003e\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e,\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e26\u003c/span\u003e\u003c/sup\u003e. Such finds are two-fold precious. On the one hand, they allow checking useful morphological characters located in fragile parts, which are not frequently preserved in the fossil record. On the other hand, they link dental to postcranial elements, thus becoming very useful in the calculation of the relative sizes between both. Each one of these exceptional finds constitutes a reference milestone to which compare many other isolated fossil elements of talpids, mostly teeth and humeri.\u003c/p\u003e\u003c/div\u003e\u003c/p\u003e \u003cp\u003eIn the present work, we study a new partial fossil skeleton of a burrowing mole that has been recently discovered in the Pliocene locality of Camp dels Ninots (CN) in NE Spain. This exceptional fossil find preserves several postcranial elements associated to almost complete dentitions and the whole mandible, all of them clearly belonging to the same individual. This specimen was tentatively ascribed to \u003cem\u003eTalpa minor\u003c/em\u003e at a first glance, considering the chronology and location of the fossil site in combination to its general size. Nevertheless, our colleague Dr. Lars van den Hoek Ostende (Naturalis Biodiversity Center, Leiden, The Netherlands) noticed in a previous stage of the present work some morphological details against its ascription to the genus \u003cem\u003eTalpa\u003c/em\u003e. The rest of the talpid genera hitherto identified in Pliocene sites from Spain\u003csup\u003e\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e27\u003c/span\u003e\u003c/sup\u003e, namely \u003cem\u003eArchaeodesmana\u003c/em\u003e, \u003cem\u003eDesmana\u003c/em\u003e, \u003cem\u003eGalemys\u003c/em\u003e, and \u003cem\u003eDesmanella\u003c/em\u003e could be easily discarded as well. Therefore, in the present study we aim to solve: 1) which is the species represented in this fossil site, 2) the phylogenetic position of this fossil among extant and extinct talpids, 3) how adapted was this species to excavate and live underground, and more specifically, 4) how did this specimen reach the sedimentary environment where it was finally preserved.\u003c/p\u003e"},{"header":"Geological settings","content":"\u003cp\u003eCN is a Pliocene site located in the town of Caldes de Malavella, in the province of Girona, in the NE of the Iberian Peninsula (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Some analyses using magneto and cyclostratigraphic techniques had provided an age of ca 3.25 Ma\u003csup\u003e\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e28\u003c/span\u003e\u003c/sup\u003e. However, recent 40Ar/39Ar datation techniques are indicative of a much older age for the site, likely Early Pliocene (work in progress). The first fragmentary bone from this locality was found in 1985 and it was assigned to the genus \u003cem\u003eLeptobos\u003c/em\u003e by Vicente\u003csup\u003e\u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e\u003c/sup\u003e. Some later prospections resulted in new fossil inputs\u003csup\u003e\u003cspan additionalcitationids=\"CR31 CR32 CR33\" citationid=\"CR30\" class=\"CitationRef\"\u003e30\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e34\u003c/span\u003e\u003c/sup\u003e, but it was in 2003 when the Institut Catal\u0026agrave; de Paleoecologia Humana i Evoluci\u0026oacute; Social (IPHES-CERCA) started the systematic research projects and excavations. Since then, this locality has delivered several articulated skeletons of vertebrates of bovids (\u003cem\u003eAlephis tigneresi)\u003c/em\u003e, tapirs (\u003cem\u003eTapirus arvernensis)\u003c/em\u003e, rhinoceros (\u003cem\u003eStephanorhinus\u003c/em\u003e cf. \u003cem\u003ejeanvireti)\u003c/em\u003e, snakes (\u003cem\u003eNatrix maura\u003c/em\u003e), turtles (\u003cem\u003eMauremys leprosa\u003c/em\u003e and \u003cem\u003eChelydropsis\u003c/em\u003e cf. \u003cem\u003epontica\u003c/em\u003e), amphibians (Pleurodelinae indet., \u003cem\u003eLissotriton\u003c/em\u003e aff. \u003cem\u003ehelveticus\u003c/em\u003e, and \u003cem\u003ePelophylax\u003c/em\u003e sp.), and freshwater fishes (\u003cem\u003eLeuciscus\u003c/em\u003e sp. and \u003cem\u003eLuciobarbus\u003c/em\u003e sp.)\u003csup\u003e\u003cspan additionalcitationids=\"CR32 CR33\" citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e34\u003c/span\u003e\u003c/sup\u003e. To a lesser extent, also diatoms, invertebrates (insects), micro- and macroflora (such as \u003cem\u003eLaurophyllum sp.\u003c/em\u003e) have been found\u003csup\u003e\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e28\u003c/span\u003e,\u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e35\u003c/span\u003e,\u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e36\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eCN site was in origin a maar lake of a volcano found in the Catalan Volcanic Complex (CVC) at the southern border of La Selva basin. There, volcanic activity ranged from the Miocene till the Holocene. The CVC is part of the volcanic provinces of the European Rift System into the Catalan Coastal Ranges\u003csup\u003e\u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e37\u003c/span\u003e\u003c/sup\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). La Selva basin was the result of an extensional tectonic episode during the Miocene-Pliocene, generated by the movement of two sets of faults oriented ENE-WSW and NW-SE of a previously fractured Paleozoic basement (for further details about the geotectonic setting see Roca \u003cem\u003eet al.\u003c/em\u003e\u003csup\u003e\u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e38\u003c/span\u003e\u003c/sup\u003e and Tassone \u003cem\u003eet al.\u003c/em\u003e\u003csup\u003e\u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e39\u003c/span\u003e\u003c/sup\u003e). The tectonic activity favored the volcanism during the Miocene, with intense phases during the Pliocene\u003csup\u003e\u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e40\u003c/span\u003e,\u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e41\u003c/span\u003e\u003c/sup\u003e. Several volcanic episodes are related with the meteoric water infiltration according to fractures of the basement and porosity of Pliocene materials, which filled the basin. The near surface magma-water interactions generated a mixed volcanism with an alternating eruptive-effusive phase. Phreatomagmatic vulcanism of explosive phases resulted in the formation of a maar crater\u003csup\u003e\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e30\u003c/span\u003e,\u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e42\u003c/span\u003e\u003c/sup\u003e. Post volcanic sediments were accumulated within the maar crater as lacustrine deposits.\u003c/p\u003e \u003cp\u003eThe sedimentary infill of CN is formed by the typical vertical stratigraphic succession in maars\u003csup\u003e\u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e43\u003c/span\u003e,\u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e44\u003c/span\u003e\u003c/sup\u003e. The one described below is taken as a reference for this fossil site and represents a thick section of 8 meters at the Can Argilera excavation sector, with four units from its base to the top (Figs.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e, \u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). Unit 1 encompasses greyish clays, sandstones and diatomites. Unit 2 is represented by greenish laminated clays with diatoms, and it is differentiated in other four subunits; 2.1, 2.2 and 2.4 are characterized by the presence of carbonates. The subunit 2.3 (see 'Detailed Location' in Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e) has been recorded in the Can Argilera sector and it is divided in fine-grained reddish sands with silty admixture (Layer 10) and lacustrine greenish silts with clay laminations (Layer 11). The latter subunit is the most remarkable one in terms of articulated skeletons of mammals and plant remains. Lastly, unit 3 is formed by reddish laminated clays and silty slope wash deposits. For further details on the specific geology of the site the reader is referred to G\u0026oacute;mez de Soler \u003cem\u003eet al.\u003c/em\u003e\u003csup\u003e\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e\u003c/sup\u003e, Jim\u0026eacute;nez-Moreno \u003cem\u003eet al.\u003c/em\u003e\u003csup\u003e\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e28\u003c/span\u003e\u003c/sup\u003e, Oms \u003cem\u003eet al.\u003c/em\u003e\u003csup\u003e\u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e42\u003c/span\u003e\u003c/sup\u003e, Rodr\u0026iacute;guez-Salgado \u003cem\u003eet al.\u003c/em\u003e\u003csup\u003e\u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e45\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003e\u003cstrong\u003eSystematic paleontology\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eOrder Eulipotyphla Waddell, Okada \u0026amp; Hasegawa, 1999\u003c/p\u003e\n\u003cp\u003eFamily Talpidae Fischer, 1814\u003c/p\u003e\n\u003cp\u003eTribe Scalopini Gill, 1875\u003c/p\u003e\n\u003cp\u003eGenus \u003cem\u003eVulcanoscaptor\u0026nbsp;\u003c/em\u003enov.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eVulcanoscaptor ninoti\u003c/em\u003e sp. nov. (Figs 4, 5, 6 and 7).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eAuthorship of genus and species:\u003c/em\u003e\u003c/strong\u003e Linares-Mart\u0026iacute;n, 2025\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eEtymology.\u003c/em\u003e\u003c/strong\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003eName of the genus derived from the Latin word of \u0026lsquo;Vulcan\u0026rsquo;, the Roman god of fire, in reference to the volcanic nature of the source area, and \u0026lsquo;-scaptor\u0026rsquo;, from the ancient Greek word \u0026lsquo;scaptein\u0026rsquo;, to dig. Name of the species\u0026nbsp;invoking \u0026lsquo;ninot\u0026rsquo;, the local word to refer the opaline nodules \u0026lsquo;doll-shaped\u0026rsquo; typically found in the type-locality of the species, Camp dels Ninots.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eHolotype.\u003c/em\u003e\u003c/strong\u003e CN10-O17-NIV11-12, partial skeleton with cranial and postcranial elements.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eStorage.\u003c/em\u003e\u003c/strong\u003e IPHES-CERCA facilities.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eStratigraphic range:\u003c/em\u003e\u003c/strong\u003e Hitherto restricted to its type locality, Lower Pliocene.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eDiagnosis (genus and species).\u003c/em\u003e\u003c/strong\u003e Small sized mole with dental formula ???3/2143. Doubled mesostyle in M1 and M2. Presence of paraconule and absence of hypocone in M2. Double rooted P4. Presence of a parastyle in P4. Lower premolar row without gaps. Absence of i3 and enlarged i2. Absence of metastylid in m2. Robust and small postcranial remains. Pit for M. flexor digitorum profundus ligament present. Straight medial edge of humeral trochlea. Fusiform shape of the humeral capitulum. Well-developed and transverse olecranon crest. Anconal and coronoid processes present in the ulna. Presence of capitular process in the radius. Scaphoid and lunar not co-ossified.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eDifferential diagnosis.\u003c/em\u003e\u003c/strong\u003e See Supplementary Information 1. Supplementary Tables of measurements and comparisons with other selected species of Talpidae can be found in Supplementary Information 2 (Tables S1\u0026ndash;S9).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eDescription\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe partial skeleton of the mole lies partially embedded by the sediment exposing the lateral side of the remains. Some skeletal elements are found in anatomical connection such as the mandible, the dentition and the postcranial elements (right forelimb). The tibiofibula is isolated from the rest of elements (Fig. 4).\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eSkull and upper dentition.\u0026nbsp;\u003c/em\u003eThe skull is partially preserved but strongly damaged. None of the cranial structures can be identified through digital reconstruction. Because the remains are not scattered, the outline can be delineated. The left upper tooth row is incomplete, but it preserves the molars, three premolars and one incisor (Fig. 5a). The incisor is displaced pointing mesially. The length of the premolars decreases towards the anterior part, P4\u0026gt;P3\u0026gt;P2. The premolars are all doubled-rooted. P2 (L = 0.80; W = 0.38) is clearly unicuspid. P3 (L = 0.85; W = 0.47) is mostly dominated by a high central cusp with a curved posterior ridge which bears a metastyle at its posterior end. The anterior ridge of both premolars is straight. The protocone in P4 (L = 1.36; W = 1.13) lies mesiolingual to the paracone. The postparacrista is straight and slightly curved at the posterobuccal end. The preparacrista is curved and wide at the end.\u003c/p\u003e\n\u003cp\u003eThe molars are double-rooted and become smaller distally, M1\u0026gt;M2\u0026gt;M3. There is no cingulum in any of them. In the M1 (L = 2.27; W = 1.78) the metacone is expanded distolingually being the paracone slightly larger than the metacone. The trigon basin is deep and wide. The protocone of the M1 is lower than that of M2 and there is no evidence of the presence of the hypocone. The mesostyle is divided into two cusps separated by a deep valley. The parastyle and the metastyle are also well developed. In the M2 (L = 1.82; W = 1.66), the trigon basin forms a deep valley as in M1. The paracone is the highest cusp and the protocone is the smallest. The paracrista is a bit shorter than the postparacrista and similar in length with the postmetacrista. No hypocone is discerned, whereas the metastyle and parastyle are well-developed. The mesostyle valley follows the same pattern as in M1. In the M3 (L = 1.52; W = 1.35), the trigon basin is wide but not as deep as in the other molars. The metacone and the paracone have a similar height in contrast to the protocone, which is smaller. All the cusps are wide and rounded. The parastyle is well developed.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eMandible and lower dentition.\u0026nbsp;\u003c/em\u003eBoth hemimandibles and most of the lower dentition (Fig. 5b) are preserved. Only the right hemimandible and its whole dentition (Fig. 5c) is exposed but digital reconstructions demonstrates that the left hemimandible additionally preserves two molars, two premolars, the canine and one incisor. Anterior mental foramina is observed below p2 and p3, and a less clear posterior one is placed below m1 and m2.\u003c/p\u003e\n\u003cp\u003eThe corpus mandibulae is slender and elongated. The distal part becomes wider and convex towards the angular process. The anterior profile of the coronoid process is slightly curved and straightens towards the tip of the coronoid process. It has a convex shape with a faint notch. The posterior profile delineates a concave curvature that connects to the condylar process. The condylar process points backwards, and it is longer than the angular process. The angular process is short, and the tip ends anterior to the angular process and posterior to the coronoid process.\u003c/p\u003e\n\u003cp\u003eMeasurements (mm): Length of the corpus mandibulae = 14.63; maximum thickness of mandible (below m2) = 1.40; maximum height of mandible (below m2) = 1.45, minimum thickness of mandible (below i1) = 0.43; Minimum height of mandible (below i2)\u0026nbsp;= 1.12; height of coronoid process = 4.94.\u003c/p\u003e\n\u003cp\u003eRegarding the dentition, the incisors and the canine present an elongated and flat crown. Both incisors are single-rooted. The i2 is the largest and widest of the incisors. Its shape is spatulated. The canine is smaller than the incisors. The premolars are doubled-rooted unicuspids and there are no gaps between them. The lateral outlines of their crowns are almost triangular. All of them have divergent and rather stout roots with rounded tips. Towards the distal side, premolars are progressively larger, p1\u0026lt; p2 \u0026lt; p3 \u0026lt; p4. The cusps in all of them are rather high and sharp. The p1 (Right: L = 0.52; W = 0.35; Left:\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003eL = 0.50, W = 0.38) stands as a simple single-cusped tooth. The posterior ridge of p2 (Right: L = 0.65; W = 0.33; Left: L = 0.60; W = 0.32) and p3 (L = 0.71; W = 0.33) are curved and wide towards the posterior end. In p4 (L = 0.88; W = 0.59), the anterior ridge of the protoconid is straight. The posterior ridge is slightly convex towards the entoconid.\u003c/p\u003e\n\u003cp\u003eThe molars are doubled-rooted, and they are progressively smaller towards the distal side, m1 \u0026gt; m2 \u0026gt; m3. In the m1 (L = 2.04; W = 1.33), the protoconid is higher than all the lingual cusps, but similar in size to the hypoconid. The talonid basin is delimited anteriorly by the oblique cristid which ends anterolingually between the metaconid and entoconid without the development of a metastylid. The entoconid and metaconid cusps have the same height, being slightly larger than those shown in the paraconid. No talonid notch is observed. There is a well-developed entostylid at the distolingual side. In the m2 (Right: L = 1.92; W = 1.32; m2: L = 1.89; W = 1.29), the talonid basin is reduced compared with its corresponding in the m1. The paracristid in m2 is higher than in m1. The oblique cristid follows the same pattern as in m1. The metaconid is higher than the paraconid and slightly higher than the entoconid. Anteriorly, the praecingulid is narrow. The protoconid is higher than the hypoconid. The talonid notch is not observed but the entostylid is well developed as in the m1. The talonid basin in m3 (Right: L = 1.54, W = 1.07; Left:\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003eL = 1.53, W = 1.04) is smaller than that of m1 and m2. The metaconid is similar in height to the paraconid but slightly higher than the entoconid. The praecingulid is narrow and the protoconid is higher than the hypoconid as in the m2. The entostylid is not developed.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003ePostcranial elements.\u0026nbsp;\u003c/em\u003eThe digital reconstructions show a mash of numerous rib fragments and vertebrae flattened by compaction. It has been impossible to restore any element of the postcranial axial skeleton in an objective way. Similarly, one clavicle and scapula are preserved and partially exposed. However, in digital reconstructions they are flattened and strongly fragmented, so it has been impossible to describe the original morphology of any element from the shoulder girdle.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eHumerus.\u0026nbsp;\u003c/em\u003eOnly the distal part of the right humerus is preserved (Fig. 6a). At the distal end of the epiphysis, the ectepicondylar (=Lateral epicondyle) and the entepicondylar (=Medial epicondyle) processes are rather well preserved but their tips are missing. In the ectepicondyle, the capitulum is laterally elongated with a fusiform shape. The surface of the entepicondyle is smaller than that of the ectepicondyle. The entepicondylar foramen is shown as a deep groove. Adjacent to this groove, a wide elliptical fossa for \u003cem\u003eM. flexor digitorum profundus\u003c/em\u003e ligament is observed. At the posterior side of the distal end, the epicondyles are separated by a large trochlear area associated with the broadening of the humerus in which there is a small projection separating the trochlea from the fossa for \u003cem\u003eM. flexor digitorum profundus\u003c/em\u003e ligament. This area becomes deep towards the shaft of the humerus, giving rise to the olecranon fossa.\u003c/p\u003e\n\u003cp\u003eMeasurements (mm): Length = 7.14; maximum distal width = 6.04; shaft thickness = 2.37; shaft width (minimum diaphysal widht) = 3.30.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eRadius.\u0026nbsp;\u003c/em\u003eThe right radius (Fig. 6b) is preserved in lateral connection with the distal part of the humerus. At its proximal end, the strong and stout capitular process is followed by the glenoid cavity. The glenoid cavity is deep and concave. In medial view, the ulnar articular facet of the capitular process is elongated and clearly defined. In lateral view, a conspicuous crest extends distally from the capitular process, showing a small groove for the attachment of \u003cem\u003eabductor pollicis\u003c/em\u003e muscle. In medial view, next to this crest, a wide fossa is observed in which the radial head of abductor muscle is attached. In the distal part of the radius, the scars for tendon of \u003cem\u003eextensor carpi radialis\u003c/em\u003e muscle forms a slight protuberance between them. The articular facets for the lunar and scaphoid are narrow and convex.\u003c/p\u003e\n\u003cp\u003eMeasurements (mm): Length (Glenoid cavity \u0026ndash; articular facet) = 7.56; shaft length = 6.09; proximal width = 2.57; distal width = 2.77; maximum distal thickness = 1.56.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eUlna.\u0026nbsp;\u003c/em\u003eThe right ulna (Fig. 6c)\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003eis preserved in connection with the radius and the distal end of the humerus. The proximal crest is elongated in anterior view with a widely extended area of insertion for the triceps. The decrease of the extension of this area towards its distal part forms the medial and lateral olecranon crest, which ends with the protuberance of the anconaeus process. Medial and lateral olecranon crests are elongated increasing the length of the ulna over the radius. In lateral view, a protruding proximal crest is enhanced by the depth of the abductor fossa. Towards its distal part, at the anconaeus-level, this fossa presents a small groove of the abductor scar. Below them, the coronoid process is observed with a small protuberance compared with the anconaeus. This difference in size conforms the semilunar notch. Next to the coronoid process, the radial articular facet overhangs the abductor fossa expanded towards the laterodistal side. In anterior view, the humeral articular facet is observed between the anconaeus and the coronoid process, which forms a slight depression. Below the coronoid process there is a small scar for the insertion of brachialis muscle. The area delimited by the anconoeus process and the coronoid process is the functional zone for the rotation of the humerus, namely the trochlear area. The distal part of the ulna is wide at the connection of the posterior crest and the abductor fossa with the shaft. In lateral view, the shaft ends with the styloid process, which is large with a stout and rounded terminal process. The ulnar articular facet is narrow and moderately deep ending in a stout cuneiform articular facet.\u003c/p\u003e\n\u003cp\u003eMeasurements (mm): Length = 13.00; olecranon length (proximal crest \u0026ndash; anconaeus process = 4.11; mediolateral diameter = 1.30; width of proximal crest = 4.22.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eCarpal bones.\u0026nbsp;\u003c/em\u003eAll the carpalia are preserved (Fig. 7a). The triquetrum, hamate, centrale, trapezoid, trapezium and capitate are in connection below the third and fourth metacarpal. The lunate and the scaphoid are disconnected from the hand. In the scaphoid, the groove for the insertion of the \u003cem\u003eflexor carpi radialis\u003c/em\u003e muscle is observed (Fig. 7b).\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eMetacarpal bones.\u0026nbsp;\u003c/em\u003eFour elements of the metacarpalia are preserved, the second (II) being the longest of them (Fig. 7b). The proximal prominence is rounded and stout. The outline of the phalanx articular facet is strong and it forms two protuberances. Generally, the metacarpals sustain the shape of the phalanx articular facet. Furthermore, the size of the metacarpals decreases from II to V. The proximal prominences of the third metacarpal are stouter than that of the second. The metacarpal articular facet is wide. The unciform articular facet is hardly visible. The fifth metacarpal is shorter and narrower than the rest. The unciform articular facet is strong and rounded.\u003c/p\u003e\n\u003cp\u003eMeasurements (mm): Mc II: L = 2.66, W = 1.45, Distal W = 1.63; Mc III: L = 2.31, W = 1.63, Distal W = 1.67; Mc IV: L = 1.57, W = 1.43, Distal W = 1.46; Mc V: L = 1.52, W = 1.37, Distal W = 1.40.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003ePhalanges.\u0026nbsp;\u003c/em\u003eSix phalanges are preserved (Fig. 7b). The proximal phalanges are shorter than the metacarpals. They are stout and the proximal articular facet is wide in their connection with the metacarpals. The distal articular facet is wide and rounded to the sides with a similar shape to that of the middle phalanges. The middle phalanges are shorter than the proximal phalanges, and they become narrower towards the distal end. The proximal articular facets are wide and rounded. The distal articular facets are slightly thinner than the proximal ones. Distal phalanges are long, and they flatten towards their distal ends. They have a characteristic shovel-like shape with a small notch in the extreme. The proximal articular facet is narrower than the distal articular facet of the middle phalanges.\u003c/p\u003e\n\u003cp\u003eMeasurements (mm): Pp III:\u0026nbsp;L = 1.44, W= 1.67, Distal W = 1.54 ; Pp IV: L = 1.39 , W = 1.46, \u0026nbsp; Distal W = 1.38; Mp III: L = 1.28, W= 1.54 , Distal W =1.01; Mp IV: L = 1.1, W = 1.38 , Distal W = 1.06; Dp III:\u0026nbsp;L = 4.28, W= 1.01 , Distal W =0.47; Dp IV:\u0026nbsp;L = 3.60, W= 1.06 , Distal W =0.30.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eSesamoid bones.\u0026nbsp;\u003c/em\u003eThe proximal part of one sesamoid bone is partially preserved (Fig. 7b). The proximal articular facet of this sickle-shaped bone is wide. The distal part is stout. This element has been found below the other bones of the hand. In addition, a small accessory sesamoid bone is observed.\u003c/p\u003e\n\u003cp\u003eMeasurements (mm): L = 3.13; W = 1.28.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eTibiofibula.\u0026nbsp;\u003c/em\u003eThe tibiofibula (Fig. 6d) is placed a few centimeters away from the rest of the connected skeleton of the animal. The tibia is thicker than the fibula and the two bones are separated by the interosseous space. Tibia and fibula are fused at their distal part, thus ending in a wide shaft. In the proximal part, the head of the fibula forms a rounded protuberance. In the tibia, the rounded medial condyle is separated from the lateral condyle by a notch that conforms the intercondyloid fossa. The lateral condyle is longer than the medial one. In the posterior part of this notch, there is a tuberosity which continues towards the distal part of the dorsomedial ridge. This ridge becomes slightly wider mediolaterally at its distal part ending with a slight depression before the medial malleolus protuberance. Next to medial malleolus, in lateral view, a tuberosity conforms the lateral malleolus. In the posterior surface of the distal part, the groove for the \u003cem\u003eM. flexor digitorum tibialis\u003c/em\u003e is observed. A low narrow ridge of the tibiofibular shaft separates the previous groove from that for the tendon of the \u003cem\u003eflexor digitorum fibularis\u003c/em\u003e muscle. In the distal extreme, a short and narrow crest separates the groove for the \u003cem\u003eflexor digitorum tibialis\u003c/em\u003e from the that for the tibialis posterior tendon. The articular facet is widely extended laterally.\u003c/p\u003e\n\u003cp\u003eMeasurements (mm): Length = 13.97; width = 2.79; width of the distal articular facet = 2.42; interosseous space = 1.68.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eAdditional hindlimb bones.\u0026nbsp;\u003c/em\u003eOther than the tibiofibula, one femur, one metatarsal, and one distal phalanx are the only elements of the hindlimb preserved. However, the digital models reconstructed from the micro-CT scans are too flattened and fragmented to be considered a good approach of the original morphologies of these elements. Therefore, these bones do not provide sufficient resolution to be confidently described.\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003e\u003cstrong\u003eTaxonomy and phylogenetic relationships\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe specimen from CN herein studied is, to our knowledge, the most complete Pliocene talpid skeleton ever reported in Europe. Other similar exceptionally preserved fossils of talpids are the Oligocene specimen of \u003cem\u003eGeotrypus antiquus\u0026nbsp;\u003c/em\u003efrom the German locality of Enspel\u003csup\u003e26\u003c/sup\u003e, and the Miocene samples of \u003cem\u003eMygalea jaegeri\u003c/em\u003e, \u003cem\u003eProscapanus sansaniensis\u003c/em\u003e, and \u003cem\u003eGeotrypus montisasini\u003c/em\u003e reported by\u0026nbsp;Schwermann and Thompson\u003csup\u003e16\u003c/sup\u003e. Except for these privileged finds,\u0026nbsp;the fossil specimens of the family Talpidae are usually restricted to isolated teeth and some characteristic postcranial elements, which make their correlation difficult. The discovery of this mole in the Pliocene site of CN turned into an exceptional landmark by the combination of unexpected characters in the only specimen recovered.\u003c/p\u003e\n\u003cp\u003eThe European fossil record of fossorial moles during the Pliocene was dominated by \u003cem\u003eTalpa\u003c/em\u003e\u003csup\u003e5\u003c/sup\u003e. However, some other less frequent talpids have been documented in literature\u003csup\u003e11\u0026ndash;13,17,46,47\u003c/sup\u003e. None of the unusual fossil forms described in these works completely fit the morphology or size of the specimen from CN. Indeed, all the Pliocene occurrences of fossorial moles in Spain have been traditionally limited to the species \u003cem\u003eT. fossilis\u003c/em\u003e and \u003cem\u003eT. minor\u003csup\u003e27\u003c/sup\u003e,\u003c/em\u003e directly ascribing all the wide and robust fossil humeri of talpids to one of these two species\u003csup\u003e48\u003c/sup\u003e. Actually, the specimen CN10-O17-NIV11-12 was tentatively identified as \u003cem\u003eTalpa minor\u0026nbsp;\u003c/em\u003eat first sight (see Introduction). Subsequent detailed scrutiny of the fossil resulted in the observation of an unusual configuration of the anterior lower toothrow, not typically found in the Talpini, which required a thorough phylogenetic analysis (see Material and methods). Together with our new species found, we took the chance to include some other taxa not considered in previous analyses, namely the genera \u003cem\u003eMyxomygale\u0026nbsp;\u003c/em\u003e(data taken from \u003cem\u003eM. hutchisoni\u003c/em\u003e, \u003cem\u003eM. antiqua\u003c/em\u003e) and \u003cem\u003eHugueneya\u003c/em\u003e (data taken from \u003cem\u003eH. primitiva\u003c/em\u003e, and \u003cem\u003eHugueneya\u003c/em\u003e sp. in Lopatin\u003csup\u003e49\u003c/sup\u003e, and the species \u003cem\u003eMongoloscapter zhegalloi\u003c/em\u003e, \u003cem\u003eSkoczenia copernici\u003c/em\u003e, and \u003cem\u003eAlpiscaptulus medogensis\u003c/em\u003e.\u003c/p\u003e\n\u003cp\u003eThe results of the cladistic analyses performed are shown in Fig. 8. Applying Goloboff\u0026apos;s criterion (K=2) and equally weighted characters of different types (ordered and unordered) results in the most parsimonious tree possible (CI = 0.3910, CI excluding uninformative characters = 0.3902, HI = 0.7065, HI excluding uninformative characters = 0.6098, RI=0.6148, RC = 0.2404, G. fit = -97.02592, tree length = 862) (Fig. 8). These results are mostly in line with previous studies\u003csup\u003e2,3,14,16\u003c/sup\u003e, but there are some differences in the resulting evolutionary tree which deserve to be pinpointed.\u003c/p\u003e\n\u003cp\u003eIt is worth noting that in our analyses (Fig. 8): 1) \u003cem\u003eTegulariscaptor minor\u003c/em\u003e is closer to the Urotrichini than to the Uropsilinae (as proposed by Sansalone \u003cem\u003eet al\u003c/em\u003e.\u003csup\u003e14\u003c/sup\u003e), but it is phylogenetically distinct from \u003cem\u003eGeotrypus\u003c/em\u003e (stem group to true moles); 2) \u003cem\u003eGeotrypus montisasini\u003c/em\u003e stays separated from \u003cem\u003eGeotrypus antiquus\u003c/em\u003e (explained by Schwermann and Thompson\u003csup\u003e16\u003c/sup\u003e; 3) \u003cem\u003eEotalpa\u0026nbsp;\u003c/em\u003eis placed in a more basal position than \u003cem\u003eUropsilus\u003c/em\u003e; 4) \u003cem\u003eNeurotrichus\u0026nbsp;\u003c/em\u003eis not the sister group of \u003cem\u003eScaptonyx\u003c/em\u003e; 5) \u003cem\u003eEuroscaptor\u003c/em\u003e and \u003cem\u003eMogera\u003c/em\u003e are closer to \u003cem\u003eTalpa\u003c/em\u003e than to the rest of the Talpini taxa; 6) \u003cem\u003eParascaptor\u0026nbsp;\u003c/em\u003eis not the sister group of \u003cem\u003eScaptochirus\u003c/em\u003e [indeed the latter is a more derived form]; 7) \u003cem\u003eScalopus\u003c/em\u003e is a more derived form than \u003cem\u003eScapanus\u003c/em\u003e; and 8) \u003cem\u003eMioscalops\u0026nbsp;\u003c/em\u003eis more derived\u003cem\u003e\u0026nbsp;\u003c/em\u003ethan \u003cem\u003eLeptoscaptor\u003c/em\u003e.\u003c/p\u003e\n\u003cp\u003eSimilarly, the introduction of new taxa modifies the trends of previous phylogenetic trees in that: 1) \u003cem\u003eMyxomygale\u003c/em\u003e is placed as a more basal taxon than the Desmaninae, clearly separated from the Urotrichini (in which it had been previously included by Ziegler\u003csup\u003e50\u003c/sup\u003e and Hugueney \u0026amp; Maridet\u003csup\u003e51\u003c/sup\u003e); 2) \u003cem\u003eMongoloscapter\u003c/em\u003e, which is included in the Scaptonychini according to Lopatin\u003csup\u003e52\u003c/sup\u003e, shows a more derived form than those, being the sister group of \u003cem\u003eNeurotrichus;\u003c/em\u003e 3) \u003cem\u003eSkoczenia\u003c/em\u003e is placed in the Talpini tribe, as stated by Rzebik-Kowalska\u003csup\u003e13\u003c/sup\u003e, close to \u003cem\u003eScaptochirus\u003c/em\u003e; and 4) \u003cem\u003eAlpiscaptulus\u003c/em\u003e and \u003cem\u003eHugueneya\u003c/em\u003e are clustered with other members of the tribe Scalopini, as predicted by the works of Chen \u003cem\u003eet al.\u003c/em\u003e\u003csup\u003e4\u003c/sup\u003e\u003csup\u003e\u0026nbsp;\u003c/sup\u003eand Lopatin\u003csup\u003e49\u003c/sup\u003e.\u003c/p\u003e\n\u003cp\u003eAccording to this analysis, \u003cem\u003eVulcanoscaptor ninoti\u003c/em\u003e gen. et sp. nov., must be also included in the tribe Scalopini, being closely related with \u003cem\u003eScalopus\u003c/em\u003e and \u003cem\u003eScapanus\u003c/em\u003e. The position of this new taxon in the resulting phylogenetic tree is supported by 11 characters that differentiate it from the rest of the species of the tribe Scalopini, namely: 1) the number of roots of P4 (c. 11); 2) the presence or absence of a paraconule in M2 (c.16); 3) the dimensions of the postmetacrista and preparacrista in M2 (c.20); 4) the presence or absence of a talonid notch in m1-m2 ( c.24); 5) the absence or presence of gaps in the lower premolar row (c.27); 6) the position of the protocone in P4 (c.32); 7) the length of M2 (c.34); 8) the length of M3\u003csup\u003e\u0026nbsp;\u003c/sup\u003e(c.35); 9) the length of m1 (c.44); 10) the length of m2 (c.45); and 11) the location of the posterior mental foramen in the lower mandible (c.69).\u003c/p\u003e\n\u003cp\u003eThese results strongly support the nature of \u003cem\u003eVulcanoscaptor ninoti\u003c/em\u003e gen. et sp. nov. as a new Scalopini. In this sense, the phylogenetic analysis carried out is reinforcing the results previously reached by Barrow \u0026amp; MacLeod\u003csup\u003e53\u003c/sup\u003e on the morphology of talpid mandibles or on the humeri\u003csup\u003e54,55\u003c/sup\u003e. Nevertheless, this Scalopini form is clearly different from any other genus in the tribe because many of them do not share the same dental formula and / or they display different configurations of their postcranial bones. More specifically, the humerus of our mole resembles that of \u003cem\u003eCondylura\u0026nbsp;\u003c/em\u003ein gross proportions, but the distal part acquires an intermediate shape between \u003cem\u003eParascalops\u0026nbsp;\u003c/em\u003eand \u003cem\u003eScapanus\u003c/em\u003e, with a notch in the trochlear area less pronounced than in \u003cem\u003eParascalops\u0026nbsp;\u003c/em\u003e(also than in \u003cem\u003eScalopoides\u0026nbsp;\u003c/em\u003eand \u003cem\u003eScapanulus\u003c/em\u003e)\u003cem\u003e,\u0026nbsp;\u003c/em\u003ebut more evident than in \u003cem\u003eScapanus\u003c/em\u003e. Moreover, the lateral and medial epicondyles of \u003cem\u003eVulcanoscaptor\u003c/em\u003e gen. nov. are more robust than in \u003cem\u003eParascalops,\u0026nbsp;\u003c/em\u003ebut weaker than in \u003cem\u003eScapanus\u003c/em\u003e. Similarly, the proximal end of the ulna of \u003cem\u003eVulcanoscaptor ninoti\u003c/em\u003e gen. et sp. nov. is similar to \u003cem\u003eScalopoides\u0026nbsp;\u003c/em\u003ein the broad area for the insertion of the triceps muscle. This morphology is also similar to that of \u003cem\u003eCondylura\u003c/em\u003e (see Hutchinson\u003csup\u003e56\u003c/sup\u003e), with the exception of the olecranon process and the articular facets, which show an intermediate morphology between \u003cem\u003eParascalops\u0026nbsp;\u003c/em\u003eand \u003cem\u003eScapanus\u003c/em\u003e.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003ePaleobiology and taphonomy\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAdaptation efforts in subterranean environments imply a high development of the humerus and a subsequent impact on other functional traits, so the anterior mobility of the forelimb is affected by maximizing the abduction movement\u003csup\u003e57\u003c/sup\u003e. According to Meier \u003cem\u003eet al.\u003c/em\u003e\u003cem\u003e\u003csup\u003e54\u003c/sup\u003e\u003c/em\u003e, such modifications are clearly reflected in an extremely short, broad and compact bone specialized for digging. The complexity of the humerus is the result from the high load that the moles must overcome with the forelimbs when digging. Supporting such intense mechanical strains need from a great development of the muscles involved, mainly those of the triceps muscular complex and their attachments sites\u003csup\u003e54,57,58\u003c/sup\u003e. The dimensions and the compaction of the humerus herein described, together with the strong development of the forearm, imply a huge development of the abductor muscles (i.e. \u003cem\u003eteres major\u003c/em\u003e, \u003cem\u003epectoralis\u003c/em\u003e, \u003cem\u003esubscapularis\u003c/em\u003e and \u003cem\u003elatissimus dorsi\u003c/em\u003e). According to Gambaryan \u003cem\u003eet al.\u003c/em\u003e\u003csup\u003e59\u003c/sup\u003e, these traits are related with a major stabilization of the forearm at the elbow and wrist joints in order to maintain the humerus and the hand in the correct position for the lateral thrust. Meier \u003cem\u003eet al.\u003c/em\u003e\u003cem\u003e\u003csup\u003e54\u003c/sup\u003e\u003c/em\u003e\u003cem\u003e\u0026nbsp;\u003c/em\u003edetailed that in the fossorial clades (i.e., Talpini and Scalopini) the humeri showed a rather buckled outline and a slightly elliptic medullary cavity, reflecting the torsion of this element and the deep reaching distal end of the deltopectoral crest. Therefore, there is no doubt that \u003cem\u003eVulcanoscaptor ninoti\u003c/em\u003e gen. nov. et sp. was a highly specialized burrower (for further details, see extended discussion in Supplementary Information 3).\u003c/p\u003e\n\u003cp\u003eThe find in lacustrine sediments of a mole highly specialized in burrowing deserves some comments. It can be speculated that the presence of this specimen in the anoxic bottom of the lake could be related to a possible semi-aquatic practice other than strictly burrowing. It would be difficult to assess this from the scarce record found at CN and more evidence and analysis are needed to sustain this hypothesis. However, several authors have documented swimming abilities among talpids adapted to strictly fossorial lifestyles (e.g., Hickman\u003csup\u003e60\u003c/sup\u003e and references therein). The same adaptation of the humerus needed for digging could also be useful for swimming\u003csup\u003e61\u003c/sup\u003e. Moreover, other features described are comparable with the aquatic \u003cem\u003eCondylura.\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe presence of a mole in the sediments of a lake suggest that this individual should have lived in the nearby area. It could have been dragged into the lake by a predator, like a bird\u003csup\u003e62\u003c/sup\u003e or by floods, or accidentally fell and drowned.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThe specimen here described is incomplete, and semi-articulated as it preserves a good anatomical connection among several skeletal elements but not within each portion (\u003cem\u003ee.g.\u003c/em\u003e cervical vertebras). Dispersal of the skeletal elements could have occurred after or before deposition of the remains at the anoxic bottom of the lake as suggested by the missing anatomical portions\u003csup\u003e26,62\u0026ndash;65\u003c/sup\u003e. A rapid burial in the ground or in deep waters\u003csup\u003e66\u003c/sup\u003e, instead, would have preserved most of the anatomical connections by impeding refloat to happen after developing decomposition gasses. Moreover, the anoxic bottom of the lake also prevents major bioturbators to disturb the carcass. Complete articulated body are common in maar sites\u003csup\u003e67\u003c/sup\u003e, including CN\u003csup\u003e31,68\u003c/sup\u003e. However, this mole specimen represents an exception. The lateral position could suggest that it arrived in the final deposition site already incomplete, as similar situations suggest that moles will lay ventrally or dorsally, but not laterally due to the shoulder girdle\u003csup\u003e26,62\u003c/sup\u003e. This specimen could represent the remains of a scavenged carcass accidentally felt in the lake or a floated carcass that sank after decomposition gasses escaped. Lastly, post depositional processes such as faulting that affected CN could also be responsible for the differential preservation of some elements\u003csup\u003e69,70\u003c/sup\u003e. For further details, see extended discussion in Supplementary Information 3.\u003c/p\u003e"},{"header":"Conclusions","content":"\u003cp\u003eThe discovery of the partial skeleton of a mole in CN enlarges the fossil record of small mammals from this locality. Comparisons of our specimen with the Pliocene and extant talpids result in its identification as a new genus and species included in the tribe Scalopini by an unusual combination of morphological characters of its dentition and postcranial elements. This is an unexpected find, considering that Scalopini moles were not frequent in Europe after the Miocene.\u003c/p\u003e \u003cp\u003eThe phylogenetic analysis performed to place this new find has resulted in a major consensus tree, which fits quite well the models previously obtained by other authors. In this sense, only a few new locations of some taxa within the phylogeny of talpids are discerned. The inclusion of the characters codified for \u003cem\u003eMyxomygale\u003c/em\u003e, \u003cem\u003eHugueneya\u003c/em\u003e, \u003cem\u003eMongoloscapter\u003c/em\u003e, \u003cem\u003eSkoczenia\u003c/em\u003e, and \u003cem\u003eAlpiscaptulus\u003c/em\u003e, together with the new ones of \u003cem\u003eVulcanoscaptor\u003c/em\u003e gen. nov. has resulted in new positions for the genera \u003cem\u003eEotalpa\u003c/em\u003e, \u003cem\u003eTegulariscaptor\u003c/em\u003e, \u003cem\u003eGeotrypus\u003c/em\u003e, and \u003cem\u003eNeurotrichus\u003c/em\u003e.\u003c/p\u003e \u003cp\u003eWith respect to its adaptations, digital reconstructions display several traits of the forelimb which implies intense modifications of the forearm as a respond of the biomechanics of the humerus to achieve a great efficiency throughout digging. This is clearly evidencing that \u003cem\u003eVulcanoscaptor ninoti\u003c/em\u003e nov. gen et sp. was adapted to a fossorial lifestyle.\u003c/p\u003e \u003cp\u003eThe strange occurrence of this mole in the fossil site of CN is a difficult issue to tackle for which different scenarios are suggested. The disposition and the optimal preservation of the partial skeleton of the mole indicates that whatever the cause of death, the specimen probably deposited already partially disarticulated. Posteriorly, the soft tissues were decomposed promoting the disarticulation of other remains before their compaction and fragmentation.\u003c/p\u003e \u003cp\u003eThe capability of the mole for swimming whereby it would have reached the lake deliberately, or the absence of this quality, could determine whether the mole died on the shore during a regular soak or drowned after falling into the lake. The real causes of death and how the specimen arrived at lacustrine sediments remain uncertain and they will deserve further research.\u003c/p\u003e"},{"header":"Material and methods","content":"\u003cp\u003eThe great extension of the excavation area (approximately 275,000 m\u003csup\u003e2\u003c/sup\u003e) required a complex level of organization, divided into several sectors, pits, sections, layers, units and subunits. Up to date, five sectors have been excavated: Can Argilera, Can Pol, Can Cateura, Comercial and Butano sectors. To define the limits of the lake and the fossiliferous layers, several pits of about 4 or 5 meters of maximum depth were carried out with a backhoe. Their dimensions varied depending on the evidence of the potential fossil content. Subsequently, the fossil strata were manually excavated. Reference systems such as UTM (ETRS89) and Cartesian coordinates were used to position the pits and fossil occurrences. The placement of the fossil was recorded by means of a registration code (acronym of the site -CN-, year, sector, pit, excavation unit, square and number of register). Depending on the preservation of the remains and the materials in which they were included, different methods were applied to unearth them\u003csup\u003e31\u003c/sup\u003e.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThe specimen studied in the present work was found in the layer 11 of the subunit 2.3 (Fig. 3). In terms of registration, this subunit belongs to the pit 7/8 from the Can Argilera sector. The remains were preserved at the surface of green clayish sediments with fine-grained sandy admixture. The specimen was originally extracted in its matrix from the excavation with expanded polyurethane. After extraction, the remains were taken to the IPHES-CERCA laboratory for preparation. Once there, its excavation was completed using mechanical methods and the aid of a binocular lens.\u003c/p\u003e\n\u003cp\u003eAfter delimiting the skeletal remains, the clay block and the specimen were consolidated with ethyl silicate (Estel 1000), which was drip-applied using a syringe onto the surface of the block and the skeleton. In this case, the specimen was consolidated in several sessions without saturating the sample to avoid an excess of siliceous crystallizations on the surface. Once hardened, the block was reduced and prepared for Computerized Microtomography dividing it in five fragments. Subsequently, the fragments were adhered with acrylic resin, Paraloid\u0026reg; B-72 dissolved in 20% acetone. Finally, a structural reintegration was carried out with a putty made from the sediment of the same block agglutinated with the same acrylic resin used for adhesion. For storage, a custom-made polyethylene-based support was made in different formats: Ethafoam\u0026reg; foam, Tivek\u0026reg; tissue and a bag.\u003c/p\u003e\n\u003cp\u003eBecause the remains are partially embedded, the morphological description of this mole for subsequent taxonomic identification is complicated. To avoid the extraction of the fossil, the virtual reconstruction of the remains was carried out using computerized tomography, a nondestructive technique. Five \u0026micro;-CT scanners, with a 3D spatial resolution of up to 6 \u0026micro;m, were made with the V|Tome|X s 240 (GE Sensing \u0026amp; Inspections Technologies) at the CENIEH in Burgos (Spain). The set of images obtained after the scans were processed with the free and open access software \u0026lsquo;3D slicer\u0026rsquo;. Subsequently, and based on these models, the descriptions and measurements were made according to Hutchison\u003csup\u003e7,56\u003c/sup\u003e.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eCladistic analysis -\u0026nbsp;\u003c/em\u003eA total of 41 extant and extinct species have been scored based on morphological characters, both cranial and postcranial elements (Supplementary Information 4). The list of characters and taxa suggested for phylogenetic analyses was carried out based on that proposed by Schwermann \u003cem\u003eet al\u003c/em\u003e.\u003csup\u003e3\u003c/sup\u003e and its preceding works\u003csup\u003e2,16,61\u003c/sup\u003e. When some specific information was missing, the data matrix was completed checking the characters of selected specimens of some subfamilies, tribes, genera and species (Supplementary Information 4). For details of the different species and the skeletal remains see Schwermann \u003cem\u003eet al.\u003csup\u003e3\u003c/sup\u003e.\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eA total of 175 characters have been considered to score the different taxa based on dentition, humerus, hand and tibiofibula. Of these discrete characters 114 are binary and 61 are multistates. Some taxa (e.g., \u003cem\u003eMioscalops isodens\u003c/em\u003e, \u003cem\u003eDomninoides mimicus\u003c/em\u003e, \u003cem\u003eParascalops breweri\u003c/em\u003e) show polymorphism for some characters which are named as letters codifying different states (Supplementary Information 4). In view of the absence of preserved skeletal elements of some species, characters have been scored as missing with the symbol \u0026quot;?\u0026quot;. Finally, in the present work we have added some taxa to the list: 1) \u003cem\u003eTegulariscaptor minor\u003c/em\u003e\u003csup\u003e\u0026nbsp;\u003c/sup\u003efrom Sansalone\u003csup\u003e14\u003c/sup\u003e; 2) \u003cem\u003eMyxomygale\u003c/em\u003e \u003cem\u003ehutchisoni\u0026nbsp;\u003c/em\u003esensu Klietmann \u003cem\u003eet al\u003c/em\u003e.\u003csup\u003e\u003cspan lang=\"EN-US\"\u003e71\u003c/span\u003e\u003c/sup\u003e and \u003cem\u003eMyxomygale antiqua\u0026nbsp;\u003c/em\u003esensu Hugueney \u0026amp; Maridet\u003csup\u003e\u003cspan lang=\"EN-US\"\u003e51\u003c/span\u003e\u003c/sup\u003e; 3) \u003cem\u003eMongoloscapter zhegalloi\u0026nbsp;\u003c/em\u003eaccording to Lopatin\u003csup\u003e\u003cspan lang=\"EN-US\"\u003e52\u003c/span\u003e\u003c/sup\u003e; 4) \u003cem\u003eSkoczenia copernici\u0026nbsp;\u003c/em\u003esensu Rzebik-Kowalska\u003csup\u003e\u003cspan lang=\"EN-US\"\u003e13\u003c/span\u003e\u003c/sup\u003e, \u003cem\u003eAlpiscaptulus medogensis\u0026nbsp;\u003c/em\u003eby Chen \u003cem\u003eet al\u003c/em\u003e.\u003csup\u003e\u003cspan lang=\"EN-US\"\u003e4\u003c/span\u003e\u003c/sup\u003e; 5) \u003cem\u003eHugueneya\u0026nbsp;\u003c/em\u003esp. in Lopatin\u003csup\u003e49\u003c/sup\u003e\u003cem\u003e\u0026nbsp;\u003c/em\u003eand \u003cem\u003eHugueneya\u003c/em\u003e \u003cem\u003eprimitiva\u003c/em\u003e according to Van den Hoek Ostende\u003csup\u003e72\u003c/sup\u003e, and 6) the new taxon herein described, \u003cem\u003eVulcanoscaptor ninoti\u003c/em\u003e gen. et sp. nov.\u003c/p\u003e\n\u003cp\u003eFor cladistic analyses the software PAUP 4.0 \u003csup\u003e73\u003c/sup\u003e has been used to obtain the most parsimonious tree possible. The parsimony criteria have been applied by means of a heuristic search. For further veracity of the obtained tree, Bremer values \u003csup\u003e74\u003c/sup\u003e were calculated in addition to the application of the scheme of implied weights from Goloboff\u003csup\u003e75\u003c/sup\u003e. For the latter criterion, a value of K=2 was applied to reduce the homoplasy. On the other hand, when setting character types, the criterion of equal weights and a status as unordered or non-additive has been applied to all characters. Exceptionally, 27 of that characters have been considered as ordered or additive based on Wagner\u0026apos;s parsimony criterion due to the assumption of an ordered sequence according to their position in the symbol list\u003csup\u003e76\u003c/sup\u003e. Finally, in our analyses \u003cem\u003eErinaceus europaeus\u003c/em\u003e is considered as an outgroup following different works\u003csup\u003e2,16,61\u003c/sup\u003e.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eAnatomical abbreviations\u0026nbsp;\u003c/em\u003e\u0026ndash; \u003cstrong\u003eabf:\u0026nbsp;\u003c/strong\u003eabductor fossa, \u003cstrong\u003eabs:\u0026nbsp;\u003c/strong\u003eabductor scar, \u003cstrong\u003eacs:\u003c/strong\u003e accessory sesamoid bone \u003cstrong\u003eagp:\u003c/strong\u003e angular process, \u003cstrong\u003eanc:\u0026nbsp;\u003c/strong\u003eancanoeus process, \u003cstrong\u003ebrs:\u0026nbsp;\u003c/strong\u003ebrachialis scar, \u003cstrong\u003ec:\u003c/strong\u003e oblique cristid, \u003cstrong\u003ecaf:\u0026nbsp;\u003c/strong\u003ecuneiform articular facet, \u003cstrong\u003ecdp:\u003c/strong\u003e condylar process, \u003cstrong\u003ecpp:\u0026nbsp;\u003c/strong\u003ecapitular process, \u003cstrong\u003ecpt:\u0026nbsp;\u003c/strong\u003e\u003cem\u003ecapitulum\u003c/em\u003e, \u003cstrong\u003ecrp:\u003c/strong\u003e coronoid process, \u003cstrong\u003edmr:\u003c/strong\u003e dorso medial ridge, \u003cstrong\u003edp:\u003c/strong\u003e distal phalanx, \u003cstrong\u003eef:\u003c/strong\u003e entepicondylar foramen, \u003cstrong\u003eend:\u003c/strong\u003e entoconid, \u003cstrong\u003efdf:\u0026nbsp;\u003c/strong\u003e\u003cem\u003eflexor digitorium fibularis\u003c/em\u003e tendon, \u003cstrong\u003efdt:\u003c/strong\u003e \u003cem\u003eflexor digitorium tibialis\u003c/em\u003e, \u003cstrong\u003effd:\u0026nbsp;\u003c/strong\u003efossa of \u003cem\u003em. flexor digitorium\u003c/em\u003e ligament, \u003cstrong\u003eflc:\u003c/strong\u003e falciform, \u003cstrong\u003ega:\u0026nbsp;\u003c/strong\u003egroove for \u003cem\u003eabductor pollicis longus\u003c/em\u003e tendon, \u003cstrong\u003ehf:\u003c/strong\u003e head of the fibula, \u003cstrong\u003ehff:\u0026nbsp;\u003c/strong\u003ehumeral articular facet, \u003cstrong\u003ehyd:\u003c/strong\u003e hypoconid, \u003cstrong\u003ehyl:\u003c/strong\u003e hypoconulid, \u003cstrong\u003ei:\u0026nbsp;\u003c/strong\u003eincisor, \u003cstrong\u003eins:\u003c/strong\u003e interosseus space, \u003cstrong\u003ele:\u003c/strong\u003e lateral epicondyle, \u003cstrong\u003ellc:\u0026nbsp;\u003c/strong\u003elateral olecranon, \u003cstrong\u003elm:\u003c/strong\u003e lateral malleolus, \u003cstrong\u003elrf:\u0026nbsp;\u003c/strong\u003elunar articular facet, \u003cstrong\u003eltc:\u003c/strong\u003e lateral condyle, \u003cstrong\u003em:\u0026nbsp;\u003c/strong\u003emolar, \u003cstrong\u003emc:\u003c/strong\u003e metacarpal, \u003cstrong\u003emd:\u003c/strong\u003e medial phalanx, \u003cstrong\u003emdc:\u003c/strong\u003e medial condyle, \u003cstrong\u003eme:\u003c/strong\u003e medial epicondyle, \u003cstrong\u003eme:\u003c/strong\u003e metacone, \u003cstrong\u003emed:\u003c/strong\u003e metaconid, \u003cstrong\u003emes:\u003c/strong\u003e mesostyle, \u003cstrong\u003emet:\u003c/strong\u003e metastyle, \u003cstrong\u003emld:\u0026nbsp;\u003c/strong\u003emedial olecranon, \u003cstrong\u003emm:\u003c/strong\u003e medial malleolus, \u003cstrong\u003eof:\u003c/strong\u003e olecranon fossa, \u003cstrong\u003ep:\u0026nbsp;\u003c/strong\u003epremolar, \u003cstrong\u003epa:\u003c/strong\u003e paracone, \u003cstrong\u003epad:\u003c/strong\u003e paraconid, \u003cstrong\u003epar:\u0026nbsp;\u003c/strong\u003eparastyle, \u003cstrong\u003epc:\u0026nbsp;\u003c/strong\u003epectoral crest, \u003cstrong\u003epp:\u003c/strong\u003e proximal phalanx, \u003cstrong\u003epr:\u003c/strong\u003e protocone, \u003cstrong\u003eprc:\u003c/strong\u003e praecingulid, \u003cstrong\u003eprd:\u003c/strong\u003e protoconid, \u003cstrong\u003eprt:\u003c/strong\u003e protoconule, \u003cstrong\u003epxc:\u0026nbsp;\u003c/strong\u003eproximal crest, \u003cstrong\u003eraf:\u0026nbsp;\u003c/strong\u003efossa for radial head of m. abductor, \u003cstrong\u003erf:\u0026nbsp;\u003c/strong\u003eradial articular facet, \u003cstrong\u003escp:\u003c/strong\u003e scaphoide, \u003cstrong\u003esf:\u0026nbsp;\u003c/strong\u003escaphoid articular facet, \u003cstrong\u003esn:\u0026nbsp;\u003c/strong\u003esemilunar notch, \u003cstrong\u003esty:\u0026nbsp;\u003c/strong\u003estyloid process, \u003cstrong\u003eta:\u003c/strong\u003e trochlear area, \u003cstrong\u003etmp:\u0026nbsp;\u003c/strong\u003eterminal process, \u003cstrong\u003etp:\u003c/strong\u003e tibialis posterior tendon, \u003cstrong\u003eulf:\u0026nbsp;\u003c/strong\u003eulnar articular facet.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eInstitutional abbreviations\u0026nbsp;\u003c/em\u003e\u0026ndash; CENIEH (Centro Nacional de Investigaci\u0026oacute;n sobre la Evoluci\u0026oacute;n Humana, Burgos, Spain); IPHES (Institut de Paleoecologia Humana i Evoluci\u0026oacute; Social, Tarragona, Spain).\u003c/p\u003e"},{"header":"Declarations","content":"\u003ch2\u003eData availability\u003c/h2\u003e\n\u003cp\u003eHolotype: The fossil elements of Vulcanoscaptor ninoti gen. et sp. nov. are hosted in the IPHES-CERCA with the reference: CN\u0026rsquo;10. Can Argilera sector. Pit 7/8. Layers 11. Square O17. Number 12. The 3D virtual models of Vulcanoscaptor ninoti gen. et sp. nov. are accessible for viewing in the open-source 3D repository Morphosource (https://doi.org/10.17602/M2/M614677; https://doi.org/10.17602/M2/M614683; https://doi.org/10.17602/M2/M609966).\u003c/p\u003e\n\u003ch2\u003eAcknowledgements\u003c/h2\u003e\n\u003cp\u003eThe CN project is sponsored by the Caldes de Malavella town hall. Funding for this research has been provided by the Catalan Government (Generalitat de Catalunya) by means of the Departament de Cultura project CLT009/22/000043 and the research groups 2021 SGR 01238 and 2021 SGR 00127. The Spanish Government and the European Union have supported this study with the projects PID2021-122533NB-I00, PID2021-123092NB-C21 and PID2020-117289GB-I00 (Agencia Estatal de Investigaci\u0026oacute;n and European Regional Development Fund of the European Union, AEI/FEDER EU). The Institut Catal\u0026agrave; de Paleoecologia Humana i Evoluci\u0026oacute; Social (IPHES-CERCA) received financial support from the Spanish Ministry of Science and Innovation through the \u0026lsquo;Mar\u0026iacute;a de Maeztu\u0026rsquo; program for Units of Excellence (CEX2019-000945-M). The research of B.G.S., G.C., H.A.B, P.P. and M.F. is funded by the CERCA Programme/Generalitat de Catalunya. P.P. is supported by a \u0026lsquo;\u0026lsquo;Ram\u0026oacute;n y Cajal\u0026rdquo; contract (grant RYC2023-044218-I) funded by MICIU/AEI/10.13039/501100011033 and \u0026ldquo;ESF+\u0026rdquo;. The CENIEH is acknowledged for providing the micro-CT scans of the remains and Josep Fortuny (ICP) and Ivan Rey-Rodr\u0026iacute;guez (UVigo-IPHES-URV) for their useful advices with the software to manage the digital reconstructions. This work is part of the Ph.D. Dissertation of the first author, in the framework of the Ph.D. Programme \u0026lsquo;Quaternari i Prehist\u0026ograve;ria\u0026rsquo; of the Universitat Rovira i Virgili (Tarragona, Spain).\u003c/p\u003e\n\u003ch2\u003eAuthor contributions statement\u003c/h2\u003e\n\u003cp\u003eM.F. conceptualized and supervised the research; A.L.-M. and M.F. performed the descriptive part of the investigation and the writing of the original draft; A.L.-M. performed the microCT scans and data curation, the formal phylogenetic analysis, the software management, and the methodological creation of models; B.G.d.S., G.C., and J.A. worked on the resources, funding acquisition, and project administration; E.M.-R. prepared the fossil specimen and described this process; O.O. worked on the formal analysis and investigation of the geological context of the site; A.L.-M. and O.O. worked on the visualization and prepared the figures; F.G. worked on the formal analysis and investigation of the taphonomy; B.G.d.S., G.C., J.A., H.-A.B., and P.P. performed the analysis of the fossil faunal content; M.F., A.L.-M., H.-A.B. and P.P. improved later versions of the original manuscript; all the authors were involved in the discussion of the results, the review and editing of the main manuscript text.\u003c/p\u003e\n\u003ch2\u003e\u003cstrong\u003eCompeting interests\u003c/strong\u003e\u003c/h2\u003e\n\u003cp\u003eThe authors declare no competing interests.\u003c/p\u003e\n\u003ch2\u003e\u003cstrong\u003eAdditional Information\u003c/strong\u003e\u003c/h2\u003e\n\u003cp\u003eSupplementary information is available for this paper at doi:10.34810/data1921.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eSig\u0026eacute;, B., Crochet, J.-Y. \u0026amp; Insole, A. Les plus vieilles taupes. \u003cem\u003eGeobios, Mem. Spec. 1\u003c/em\u003e 141\u0026ndash;157. https://doi.org/10.1016/S0016-6995(77)80014-4 (1977).\u003c/li\u003e\n\u003cli\u003eHooker, J. J. Skeletal adaptations and phylogeny of the oldest mole Eotalpa (Talpidae, Lipotyphla, Mammalia) from the UK Eocene: The beginning of fossoriality in moles. \u003cem\u003ePalaeontology\u003c/em\u003e \u003cstrong\u003e59\u003c/strong\u003e, 195\u0026ndash;216. https://doi.org/10.1111/pala.12221 (2016).\u003c/li\u003e\n\u003cli\u003eSchwermann, A. H., He, K., Peters, B. J., Plogschties, T. \u0026amp; Sansalone, G. Systematics and macroevolution of extant and fossil scalopine moles (Mammalia, Talpidae). \u003cem\u003ePalaeontology\u003c/em\u003e \u003cstrong\u003e62\u003c/strong\u003e, 661\u0026ndash;676. https://doi.org/10.1111/pala.12422 (2019).\u003c/li\u003e\n\u003cli\u003eChen, Z. Z. \u003cem\u003eet al.\u003c/em\u003e Morphology and phylogeny of scalopine moles (Eulipotyphla: Talpidae: Scalopini) from the eastern Himalayas, with descriptions of a new genus and species. \u003cem\u003eZool J Linn Soc\u003c/em\u003e \u003cstrong\u003e193\u003c/strong\u003e, 432\u0026ndash;444. https://doi.org/10.1093/zoolinnean/zlaa172 (2021).\u003c/li\u003e\n\u003cli\u003eHe, K., Shinohara, A., Jiang, X. L. \u0026amp; Campbell, K. L. Multilocus phylogeny of talpine moles (Talpini, Talpidae, Eulipotyphla) and its implications for systematics. \u003cem\u003eMol Phylogenet Evol\u003c/em\u003e \u003cstrong\u003e70\u003c/strong\u003e, 513\u0026ndash;521 (2014).\u003c/li\u003e\n\u003cli\u003eHugueney, M. Les talpid\u0026eacute;s (Mammalia, Insectivora) de Coderet-Bransat (Allier) et l\u0026rsquo;\u0026eacute;volution de cette famille au cours de l\u0026rsquo;Oligoc\u0026egrave;ne et du Mioc\u0026egrave;ne inf\u0026eacute;rieur d\u0026rsquo;Europe. \u003cem\u003eTravaux et Documents des Laboratoires de G\u0026eacute;ologie de Lyon\u003c/em\u003e \u003cstrong\u003e50\u003c/strong\u003e, 1\u0026ndash;81. https://doi.org/10.1016/j.ympev.2013.10.002 (1972).\u003c/li\u003e\n\u003cli\u003eHutchison, J. H. Notes on type specimens of European Miocene Talpidae and a tentative classification of old world Tertiary Talpidae (Insectivora: Mammalia). \u003cem\u003eGeobios\u003c/em\u003e \u003cstrong\u003e7\u003c/strong\u003e, 211\u0026ndash;256. https://doi.org/10.1016/S0016-6995(74)80009-4 (1974).\u003c/li\u003e\n\u003cli\u003eZiegler, R. Talpiden (Mammalia, Insectivora) aus dem Orleanium and Astaracium Bayerns. \u003cem\u003eMitteilungen der Bayerische Staatssammlung f\u0026uuml;r Pal\u0026auml;ontologie und historische Geologie\u003c/em\u003e \u003cstrong\u003e25\u003c/strong\u003e, 131\u0026ndash;175 (1985).\u003c/li\u003e\n\u003cli\u003eVan Cleef-Roders, J. T. \u0026amp; Van Den Hoek Ostende, L. W. Dental morphology of Talpa europaea and Talpa occidentalis (Mammalia: Insectivora) with a discussion of fossil Talpa in the Pleistocene of Europe. \u003cem\u003eZool. Med. Leiden\u003c/em\u003e \u003cstrong\u003e75\u003c/strong\u003e, 51\u0026ndash;68 (2001).\u003c/li\u003e\n\u003cli\u003eGarc\u0026iacute;a-Alix, A., Furi\u0026oacute;, M., Minwer-Barakat, R., Mart\u0026iacute;n Su\u0026aacute;rez, E. \u0026amp; Freudenthal, M. Environmental control on the biogeographical distribution of Desmanella (Soricomorpha, Mammalia) in the Miocene of the Iberian Peninsula. \u003cem\u003ePalaeontology\u003c/em\u003e \u003cstrong\u003e54\u003c/strong\u003e, 753\u0026ndash;762. https://doi.org/10.1111/j.1475-4983.2011.01062.x (2011).\u003c/li\u003e\n\u003cli\u003eCailleux, F., van den Hoek Ostende, L. W. \u0026amp; Joniak, P. The Late Miocene Talpidae (Eulipotyphla, Mammalia) from the Pannonian Region, Slovakia. \u003cem\u003eJ Paleontol\u003c/em\u003e \u003cstrong\u003e98\u003c/strong\u003e, 128\u0026ndash;151. https://doi.org/10.1017/jpa.2023.95 (2024).\u003c/li\u003e\n\u003cli\u003eSkoczen, S. Condylurini Dobson, 1883 (Insectivora, Mammalia) in the Pliocene of Poland. \u003cem\u003eActa Palaeontol Pol\u003c/em\u003e \u003cstrong\u003e21\u003c/strong\u003e, 291\u0026ndash;313 (1976).\u003c/li\u003e\n\u003cli\u003eRzebik-Kowalska, B. Review of the Pliocene and Pleistocene Talpidae. \u003cem\u003ePalaeontologia Electronica\u003c/em\u003e \u003cstrong\u003e17\u003c/strong\u003e, (2014).\u003c/li\u003e\n\u003cli\u003eSansalone, G., Kotsakis, T., Schwermann, A. H., Van den Hoek Ostende, L. W. \u0026amp; Piras, P. When moles became diggers: Tegulariscaptor gen. nov., from the early Oligocene of south Germany, and the evolution of talpid fossoriality. \u003cem\u003eJ Syst Palaeontol\u003c/em\u003e \u003cstrong\u003e16\u003c/strong\u003e, 645\u0026ndash;657. https://doi.org/10.1080/14772019.2017.1329235 (2017).\u003c/li\u003e\n\u003cli\u003eHutterer, R. Order Soricomorpha. In:Wilson DE, Reeder DM (eds) Mammals Species of the World. A Taxonomic and Geographic Reference, 3rd edition. The Johns Hopkins University Press, Baltimore. (2005).\u003c/li\u003e\n\u003cli\u003eSchwermann, A. H. \u0026amp; Thompson, R. S. Extraordinarily preserved talpids (Mammalia, Lipotyphla) and the evolution of fossoriality. \u003cem\u003eJ Vertebr Paleontol\u003c/em\u003e \u003cstrong\u003e35\u003c/strong\u003e. https://doi.org/10.1080/02724634.2014.934828 (2015).\u003c/li\u003e\n\u003cli\u003eSansalone, G., Kotsakis, T. \u0026amp; Piras, P. New Systematic Insights about Plio-Pleistocene Moles from Poland. \u003cem\u003eActa Palaeontol Pol\u003c/em\u003e \u003cstrong\u003e61\u003c/strong\u003e, 221\u0026ndash;229. https://doi.org/10.4202/app.00116.2014 (2016).\u003c/li\u003e\n\u003cli\u003eStorch, G. \u0026amp; Qiu, Z. The Neogene mammalian faunas of Ertemte and Harr Obo in Inner Mongolia (Nei Mongol), China. 2. Moles-Insectivora: Talpidae. \u003cem\u003eSenckenbergiana Lethaea\u003c/em\u003e \u003cstrong\u003e64\u003c/strong\u003e, 89\u0026ndash;127 (1983).\u003c/li\u003e\n\u003cli\u003eMinwer-Barakat, R., Garc\u0026iacute;a-Alix, A., Mart\u0026iacute;n-Su\u0026aacute;rez, E. \u0026amp; Freudenthal, M. Early Pliocene Desmaninae (Mammalia, Talpidae) from Southern Spain and the Origin of the Genus Desmana. \u003cem\u003eJ Vertebr Paleontol\u003c/em\u003e \u003cstrong\u003e40\u003c/strong\u003e. https://doi.org/10.1080/02724634.2020.1835936 (2020).\u003c/li\u003e\n\u003cli\u003eR\u0026uuml;mke, C. G. A review of fossil and recent Desmaninae (Talpidae, Insectivora). \u003cem\u003eUtrecht Micropaleontological Bulletins Special Publication\u003c/em\u003e \u003cstrong\u003e4\u003c/strong\u003e, 1\u0026ndash;241 (1985).\u003c/li\u003e\n\u003cli\u003eSansalone, G. \u003cem\u003eet al.\u003c/em\u003e Impact of transition to a subterranean lifestyle on morphological disparity and integration in talpid moles (Mammalia, Talpidae). \u003cem\u003eBMC Evol Biol\u003c/em\u003e \u003cstrong\u003e19\u003c/strong\u003e, (2019).\u003c/li\u003e\n\u003cli\u003eDoukas, C. S., L.W., V. den H. O., C.D., T. \u0026amp; J.W.F., R. The vertebrate locality Maramena (Macedonia, Greece) at the Turolian- Ruscinian Boundary (Neogene). \u003cem\u003eM\u0026uuml;nchner Geowissenschaftliche Abhandlungen (A)\u003c/em\u003e \u003cstrong\u003e28\u003c/strong\u003e, 43\u0026ndash;64 (1995).\u003c/li\u003e\n\u003cli\u003eSansalone, G., Kotsakis, T. \u0026amp; Piras, P. Talpa fossilis or Talpa europaea? Using geometric morphometrics and allometric trajectories of humeral moles remains from Hungary to answer a taxonomic debate. \u003cem\u003ePalaeontologia Electronica\u003c/em\u003e \u003cstrong\u003e18\u003c/strong\u003e, 1\u0026ndash;17. https://doi.org/10.26879/560 (2015).\u003c/li\u003e\n\u003cli\u003eHoek Ostende, L. W. Van Den \u0026amp; Fejfar Oldrich. Erinaceidae and Talpidae (Erinaceomorpha , Soricomor- pha , Mammalia) from the Lower Miocene of Merkur-Nord (Czech Republic , MN 3). \u003cem\u003eBeitr\u0026auml;ge zur Pal\u0026auml;ontologie\u003c/em\u003e \u003cstrong\u003e30\u003c/strong\u003e, 175\u0026ndash;203 (2006).\u003c/li\u003e\n\u003cli\u003eKimmig, J., Schiffbauer, J. D. A modern definition of Fossil-Lagerst\u0026auml;tten. \u003cem\u003eTrends Ecol.\u003c/em\u003e \u003cstrong\u003e39\u003c/strong\u003e, 621\u0026ndash;624 (2024).\u003c/li\u003e\n\u003cli\u003eSchwermann, A. H. \u0026amp; Martin, T. A partial skeleton of Geotrypus antiquus (Talpidae, Mammalia) from the Late Oligocene of the Enspel fossillagerst\u0026auml;tte in Germany. \u003cem\u003ePalaontol Z\u003c/em\u003e \u003cstrong\u003e86\u003c/strong\u003e, 409\u0026ndash;439 (2012).\u003c/li\u003e\n\u003cli\u003eFuri\u0026oacute;, M., van den Hoek Ostende, L. W., Agust\u0026iacute;, J. \u0026amp; Minwer-Barakat, R. Evoluci\u0026oacute;n de las asociaciones de insect\u0026iacute;voros (Eulipotyphla, Mammalia) en Espa\u0026ntilde;a y su relaci\u0026oacute;n con los cambios clim\u0026aacute;ticos del Ne\u0026oacute;geno y el Cuaternario. \u003cem\u003eEcosistemas\u003c/em\u003e \u003cstrong\u003e27\u003c/strong\u003e, 38\u0026ndash;51. https://doi.org/10.7818/ECOS.1454 (2018).\u003c/li\u003e\n\u003cli\u003eJim\u0026eacute;nez-Moreno, G. \u003cem\u003eet al.\u003c/em\u003e Late Pliocene vegetation and orbital-scale climate changes from the western Mediterranean area. \u003cem\u003eGlob Planet Change\u003c/em\u003e \u003cstrong\u003e108\u003c/strong\u003e, 15\u0026ndash;28. https://doi.org/10.1016/j.gloplacha.2013.05.012 (2013).\u003c/li\u003e\n\u003cli\u003eVicente, J. Troballa d\u0026rsquo;un Leptobos a Caldes de Malavella (La Selva). \u003cem\u003eSocietat d\u0026rsquo;Hist\u0026ograve;ria Natural, Butllet\u0026iacute; del Centre d\u0026rsquo;Estudis de la Natura del Barcelon\u0026egrave;s Nord\u003c/em\u003e \u003cstrong\u003e1\u003c/strong\u003e, 86\u0026ndash;88 (1985).\u003c/li\u003e\n\u003cli\u003eVeh\u0026iacute;, M., Pujadas, A., Roqu\u0026eacute;, C. \u0026amp; Bux\u0026oacute;, L. P. Un edifici volc\u0026agrave;nic in\u0026egrave;dit a Caldes de Malavella (la Selva, Girona): El volc\u0026agrave; del Camp dels Ninots. \u003cem\u003eQuaderns de la Selva\u003c/em\u003e \u003cstrong\u003e11\u003c/strong\u003e, 45\u0026ndash;67 (1999).\u003c/li\u003e\n\u003cli\u003eG\u0026oacute;mez de Soler, B. \u003cem\u003eet al.\u003c/em\u003e A new key locality for the Pliocene vertebrate record of Europe: The Camp dels Ninots maar (NE Spain). \u003cem\u003eGeologica Acta\u003c/em\u003e (2012) doi:10.1344/105.000001702.\u003c/li\u003e\n\u003cli\u003eClaude, J., De Soler, B. G., Campeny, G., Agusti, J. \u0026amp; Oms, O. Presence of a chelydrid turtle in the late Pliocene Camp dels Ninots locality (Spain). \u003cem\u003eBulletin de la Societe Geologique de France\u003c/em\u003e \u003cstrong\u003e185\u003c/strong\u003e, 253\u0026ndash;256. https://doi.org/10.2113/gssgfbull.185.4.253 (2014).\u003c/li\u003e\n\u003cli\u003ePřikryl, T. \u003cem\u003eet al.\u003c/em\u003e Fish fauna of the Camp dels Ninots locality (Pliocene; Caldes de Malavella, province of Girona, Spain) \u0026ndash; first results with notes on palaeoecology and taphonomy. \u003cem\u003eHist Biol\u003c/em\u003e \u003cstrong\u003e28\u003c/strong\u003e, 347\u0026ndash;357. https://doi.org/10.1080/08912963.2014.934820 (2016).\u003c/li\u003e\n\u003cli\u003eBlain, H. \u003cem\u003eet al.\u003c/em\u003e Water frogs (Anura, Ranidae) from the Pliocene Camp dels Ninots Konservat-Lagerst\u0026auml;tte (Caldes de Malavella, NE Spain). Abstracts volume 7th International Maar Conference - Olot, Catalonia, Spain. 162\u0026ndash;163 (2018).\u003c/li\u003e\n\u003cli\u003eRobles, S., Barr\u0026oacute;n, E. \u0026amp; Cebolla, C. Estudio paleobot\u0026aacute;nico preliminar del afloramiento plioceno de Camp dels Ninots (Caldes de Malavella, Girona, Espa\u0026ntilde;a). Macroflora del sector de Can Argilera. \u003cem\u003eBolet\u0026iacute;n de la Real Sociedad Espa\u0026ntilde;ola de Historia Natural, Secci\u0026oacute;n Geol\u0026oacute;gica.\u003c/em\u003e \u003cstrong\u003e17\u003c/strong\u003e, 75\u0026ndash;89 (2013).\u003c/li\u003e\n\u003cli\u003eOms, O. \u003cem\u003eet al.\u003c/em\u003e Early lake sedimentation in the Pliocene Camp dels Ninots maar (Catalan Coastal Ranges, Spain). Abstracts volume 7th International Maar Conference - Olot, Catalonia, Spain 172-173. 172\u0026ndash;173 (2018).\u003c/li\u003e\n\u003cli\u003eMart\u0026iacute;, J., Mitjavila, J., Roca, E. \u0026amp; Aparicio, A. Cenozoic magmatism of the valencia trough (western mediterranean): Relationship between structural evolution and volcanism. \u003cem\u003eTectonophysics\u003c/em\u003e \u003cstrong\u003e203\u003c/strong\u003e, 145\u0026ndash;165. https://doi.org/10.1016/0040-1951(92)90221-Q (1992).\u003c/li\u003e\n\u003cli\u003eRoca, E., Sans, M., Cabrera, L. \u0026amp; Marzo, M. Oligocene to Middle Miocene evolution of the central Catalan margin (northwestern Mediterranean). \u003cem\u003eTectonophysics\u003c/em\u003e \u003cstrong\u003e315\u003c/strong\u003e, 209\u0026ndash;229 (1999).\u003c/li\u003e\n\u003cli\u003eTassone, A., Roca, E., Munoz, J. A., Cabrera, L. \u0026amp; Canals, M. Evolucion del sector septentrional del margen continental catalan durante el Cenozoico. \u003cem\u003eActa Geologica Hispanica\u003c/em\u003e \u003cstrong\u003e29\u003c/strong\u003e, 3\u0026ndash;37 (1994).\u003c/li\u003e\n\u003cli\u003eGuardia, P. Volcans tertiaires et quaternaires de la province de Gerona et pal\u0026eacute;omagn\u0026eacute;tisme de leurs coul\u0026eacute;es. (1964).\u003c/li\u003e\n\u003cli\u003eDonville, B. Ages potassium-argon des roches volcaniques de la Depression de la Selva (nord-est de l\u0026rsquo;espagne). (1973).\u003c/li\u003e\n\u003cli\u003eOms, O. \u003cem\u003eet al.\u003c/em\u003e Structure of the Pliocene Camp dels Ninots maar-diatreme (Catalan Volcanic Zone, NE Spain). \u003cem\u003eBull Volcanol\u003c/em\u003e \u003cstrong\u003e77\u003c/strong\u003e, (2015).\u003c/li\u003e\n\u003cli\u003eLindner, H., Gabriel, G., G\u0026ouml;tze, H.-J., Kaeppler, R. \u0026amp; Suhr, P. Geophysical and geological investigation of maar structures in the Upper Lusatia region (East Saxony). \u003cem\u003eZeitschrift der Deutschen Gesellschaft f\u0026uuml;r Geowissenschaften\u003c/em\u003e \u003cstrong\u003e157\u003c/strong\u003e, 355\u0026ndash;372 (2006).\u003c/li\u003e\n\u003cli\u003ePirrung, M. \u003cem\u003eet al.\u003c/em\u003e Lithofacies succession of maar crater deposits in the Eifel area (Germany). \u003cem\u003eTerra Nova\u003c/em\u003e \u003cstrong\u003e15\u003c/strong\u003e, 125\u0026ndash;132. https://doi.org/10.1046/j.1365-3121.2003.00473.x (2003).\u003c/li\u003e\n\u003cli\u003eRodr\u0026iacute;guez-Salgado, P. \u003cem\u003eet al.\u003c/em\u003e Mineralogical proxies of a Pliocene maar lake recording changes in precipitation at the Camp dels Ninots (Pliocene, NE Iberia). \u003cem\u003eSediment Geol\u003c/em\u003e \u003cstrong\u003e418\u003c/strong\u003e, 105910. https://doi.org/10.1016/j.sedgeo.2021.105910 (2021).\u003c/li\u003e\n\u003cli\u003eSkoczen, S. Scaptonychini Van Valen, 1967, Urotrichini and Scalopini Dobson, 1883 (Insectivora, Mammalia) in the Pliocene and Pleistocene of Poland. \u003cem\u003eActa Zoologica Cracoviensia\u003c/em\u003e \u003cstrong\u003e24\u003c/strong\u003e, 411\u0026ndash;448 (1980).\u003c/li\u003e\n\u003cli\u003eSkoczen, S. New records of Parascalops, Neurotrichus and Condylura (Talpinae, Insectivora) from the Pliocene of Poland. \u003cem\u003eActa Theriol (Warsz)\u003c/em\u003e \u003cstrong\u003e38\u003c/strong\u003e, 125\u0026ndash;137 (1993).\u003c/li\u003e\n\u003cli\u003eFuri\u0026oacute;, M. \u0026amp; Angelone, C. Insectivores (Erinaceidae, Soricidae, Talpidae; Mammalia) from the Pliocene of Capo Mannu D1 (Mandriola, central-western Sardinia, Italy). \u003cem\u003eNeues Jahrb Geol Palaontol Abh\u003c/em\u003e \u003cstrong\u003e258\u003c/strong\u003e, 229\u0026ndash;242 (2010).\u003c/li\u003e\n\u003cli\u003eLopatin, A. V. Early Miocene Small Mammals from the North Aral Region (Kazakhstan) with Special Reference to their Biostratigraphic Significance. \u003cem\u003ePaleontological journal\u003c/em\u003e \u003cstrong\u003e38\u003c/strong\u003e, 217\u0026ndash;323 (2004).\u003c/li\u003e\n\u003cli\u003eZiegler, R. Moles (Talpidae) from the late Middle Miocene of South Germany. \u003cem\u003eActa Palaeontol Pol\u003c/em\u003e \u003cstrong\u003e48\u003c/strong\u003e, 617\u0026ndash;648 (2003).\u003c/li\u003e\n\u003cli\u003eHugueney, M. \u0026amp; Maridet, O. Evolution of Oligo-Miocene talpids (Mammalia, Talpidae) in Europe: focus on the genera Myxomygale and Percymygale n. gen. \u003cem\u003eHist Biol\u003c/em\u003e \u003cstrong\u003e30\u003c/strong\u003e, 267\u0026ndash;275. https://doi.org/10.1080/08912963.2017.1282477 (2018).\u003c/li\u003e\n\u003cli\u003eLopatin, A. V. An oligocene mole (Talpidae, Insectivora, Mammalia) from Mongolia. \u003cem\u003ePaleontologicheskii Zhurnal\u003c/em\u003e \u003cstrong\u003e36\u003c/strong\u003e, 91\u0026ndash;92 (2002).\u003c/li\u003e\n\u003cli\u003eBarrow, E. \u0026amp; MacLeod, N. Shape variation in the mole dentary (Talpidae: Mammalia). \u003cem\u003eZool J Linn Soc\u003c/em\u003e \u003cstrong\u003e153\u003c/strong\u003e, 187\u0026ndash;211. https://doi.org/10.1111/j.1096-3642.2008.00376.x (2008).\u003c/li\u003e\n\u003cli\u003eMeier, P. S., Bickelmann, C., Scheyer, T. M., Koyabu, D. \u0026amp; S\u0026aacute;nchez-Villagra, M. R. Evolution of bone compactness in extant and extinct moles (Talpidae): Exploring humeral microstructure in small fossorial mammals. \u003cem\u003eBMC Evol Biol\u003c/em\u003e \u003cstrong\u003e13\u003c/strong\u003e, (2013).\u003c/li\u003e\n\u003cli\u003eSansalone, G. \u003cem\u003eet al.\u003c/em\u003e Influence of Evolutionary Allometry on Rates of Morphological Evolution and Disparity in strictly Subterranean Moles (Talpinae, Talpidae, Lipotyphla, Mammalia). \u003cem\u003eJ Mamm Evol\u003c/em\u003e \u003cstrong\u003e25\u003c/strong\u003e, 1\u0026ndash;14 (2018).\u003c/li\u003e\n\u003cli\u003eHutchison, J. H. Fossil Talpidae (lnsectivora, Mammalia) from the later Tertiary of Oregon. \u003cem\u003eBulletin of the Museum of Natural History\u003c/em\u003e 1\u0026ndash;117 (1968).\u003c/li\u003e\n\u003cli\u003eSansalone, G. \u003cem\u003eet al.\u003c/em\u003e Decoupling Functional and Morphological Convergence, the Study Case of Fossorial Mammalia. \u003cem\u003eFront Earth Sci (Lausanne)\u003c/em\u003e \u003cstrong\u003e8\u003c/strong\u003e, 112 (2020).\u003c/li\u003e\n\u003cli\u003eFurio, M. The shrew pleads \u0026lsquo;not guilty\u0026rsquo; to the mole\u0026rsquo;s murder: comment on Benn\u0026agrave;sar et al. (2015). \u003cem\u003eHist Biol\u003c/em\u003e \u003cstrong\u003e29\u003c/strong\u003e, 230-233. https://doi.org/10.1080/08912963.2016.1151016 (2017).\u003c/li\u003e\n\u003cli\u003eGambaryan, P. P., Gasc, J.-P. \u0026amp; Renous, S. Cinefluorographical study of the burrowing movements in the common mole, Talpa europaea (Lipotyphla, Talpidae). \u003cem\u003eRuss J Theriol\u003c/em\u003e \u003cstrong\u003e1\u003c/strong\u003e, 91\u0026ndash;109 (2002).\u003c/li\u003e\n\u003cli\u003eHickman, G. C. Swimming ability of talpid moles, with particular reference to the semi-aquatic Condylura cristata. \u003cem\u003eMammalia\u003c/em\u003e \u003cstrong\u003e48\u003c/strong\u003e, 505\u0026ndash;514. https://doi.org/10.1515/mamm.1984.48.4.505 (1984).\u003c/li\u003e\n\u003cli\u003eS\u0026aacute;nchez-Villagra, M. R., Horovitz, I. \u0026amp; Motokawa, M. A comprehensive morphological analysis of talpid moles (Mammalia) phylogenetic relationships. \u003cem\u003eCladistics\u003c/em\u003e \u003cstrong\u003e22\u003c/strong\u003e, 59\u0026ndash;88. https://doi.org/10.1111/j.1096-0031.2006.00087.x (2006).\u003c/li\u003e\n\u003cli\u003eM\u0026auml;hler, B., Schwermann, A. H., Wuttke, M., Schultz, J. A. \u0026amp; Martin, T. Four-dimensional virtopsy and the taphonomy of a mole from the Oligocene of Lake Enspel (Germany). \u003cem\u003ePaleobiodivers Paleoenviron\u003c/em\u003e \u003cstrong\u003e95\u003c/strong\u003e, 115\u0026ndash;131 (2015).\u003c/li\u003e\n\u003cli\u003eBehrensmeyer, A. K. Terrestrial vertebrate accumulations. In: Allison, P.A., Briggs, D.E.G. (Eds.), Taphonomy: Releasing the Data Locked in the Fossil Record. \u003cem\u003eTaphonomy: Releasing the Data Locked in the Fossil Record. Plenum Press, New York\u003c/em\u003e 291\u0026ndash;335 (1991).\u003c/li\u003e\n\u003cli\u003eSabol, M. \u003cem\u003eet al.\u003c/em\u003e Early Late Pliocene site of Hajn\u0026aacute;čka I (Southern Slovakia) - Geology, palaeovolcanic evolution, fossil assemblages and palaeoenvironment. \u003cem\u003eCFS Courier Forschungsinstitut Senckenberg\u003c/em\u003e 261\u0026ndash;274 (2006).\u003c/li\u003e\n\u003cli\u003eSyme, C.E., Salisbury, S. W. Patterns of aquatic decay and disarticulation in juvenile Indo-Pacific crocodiles (Crocodylus porosus), and implications for the taphonomic interpretation of fossil crocodyliform material. \u003cem\u003ePalaeogeogr Palaeoclimatol Palaeoecol\u003c/em\u003e \u003cstrong\u003e412\u003c/strong\u003e, 108\u0026ndash;123 (2014).\u003c/li\u003e\n\u003cli\u003eMoore, M., J., Mitchell, G. H., Rowles, T., K., Early, G. Dead Cetacean? Beach, Bloat, Float, Sink. \u003cem\u003eFront. Mar. Sci.\u003c/em\u003e \u003cstrong\u003e7\u003c/strong\u003e. https://doi.org/10.3389/fmars.2020.00333 (2020).\u003c/li\u003e\n\u003cli\u003eUhl, D., Wuttke, M., Aiglstorfer, M., Gee, C.T., Grandi, F., H\u0026ouml;ltke, O., Kaiser, T.M., Kaulfuss, U., Lee, D., Lehmann, T., Oms, O., Poschmann, M.J., Rasser, M.J., Schindler, T., Smith, K.T., Suhr, P., Wappler, T., Wedmann, S. Deep‑time maar lakes and other volcanogenic lakes as Fossil‑Lagerstatten \u0026ndash; An overview. \u003cem\u003ePaleobiodivers Paleoenviron\u003c/em\u003e \u003cstrong\u003e104\u003c/strong\u003e, 763\u0026ndash;848 (2024).\u003c/li\u003e\n\u003cli\u003eCampeny Vall-Llosera, G. \u0026amp; G\u0026oacute;mez de Soler, B. \u003cem\u003eEl Camp Dels Ninots: Rastres de l\u0026rsquo;Evoluci\u0026oacute;\u003c/em\u003e. \u003cem\u003eEl Camp dels Ninots: Rastres de l\u0026rsquo;evoluci\u0026oacute;\u003c/em\u003e (Cambridge University Press, Caldes de Malavella, 2010).\u003c/li\u003e\n\u003cli\u003eBol\u0026oacute;s, X., Oms, O., Rodr\u0026iacute;guez-Salgado, P., Mart\u0026iacute;, J., G\u0026oacute;mez De Soler, B., Campeny, G. Eruptive evolution and 3D geological modeling of Camp dels Ninots maar-diatreme (Catalonia) through continuous intra-crater drill coring. J. Volcanol. \u003cem\u003eGeotherm. Res. 419, 107369.\u003c/em\u003e \u003cstrong\u003e419\u003c/strong\u003e. https://doi.org/10.1016/j.jvolgeores.2021.107369 (2021).\u003c/li\u003e\n\u003cli\u003eGrandi, F., Del Valle, H., C\u0026aacute;ceres, I., Rodr\u0026iacute;guez-Salgado, P., Oms, O., Fern\u0026aacute;ndez-Jalvo, Y., Garc\u0026iacute;a, F., Campeny, G \u0026amp; G\u0026oacute;mez de Soler, B. Exceptional preservation of large fossil vertebrates in a volcanic setting (Camp dels Ninots, Spain). \u003cem\u003eHist. Biol\u003c/em\u003e.\u003cstrong\u003e 35\u003c/strong\u003e, 1234-1249. https://doi.org/10.1080/08912963.2022.2085570 (2023).\u003c/li\u003e\n\u003cli\u003eKlietmann, J., Nagel, D., Rummel, M. \u0026amp; van den Hoek Ostende, L. W. A gap in digging: the Talpidae of Petersbuch 28 (Germany, Early Miocene). \u003cem\u003ePalaontol Z\u003c/em\u003e \u003cstrong\u003e89\u003c/strong\u003e, 563\u0026ndash;592 (2015).\u003c/li\u003e\n\u003cli\u003eVan den Hoek Ostende, L. W. The Talpidae (Insectivora, Mammalia) of Eggingen-Mittelhart (Baden-Wurttenberg, FRG) with special reference to the Paratalpa-Desmanodan lineage. Preprint at (1988).\u003c/li\u003e\n\u003cli\u003eSwofford, D. L. PAUP*. Phylogenetic Analysis Using Parsimony (*And Other Methods), version 4. \u003cem\u003eSinauer Associates, Sunderland, Massachusetts.\u003c/em\u003e \u003cstrong\u003eVersion 4\u003c/strong\u003e, (2002).\u003c/li\u003e\n\u003cli\u003eBremer, K. R. Branch support and tree stability. \u003cem\u003eCladistics\u003c/em\u003e \u003cstrong\u003e10\u003c/strong\u003e, 295\u0026ndash;304 (1994).\u003c/li\u003e\n\u003cli\u003eGoloboff, P. A. Estimating character weights during tree search. \u003cem\u003eCladistics\u003c/em\u003e \u003cstrong\u003e9\u003c/strong\u003e, 83\u0026ndash;91 (1993).\u003c/li\u003e\n\u003cli\u003eSwofford, D. L., \u0026amp; Maddison, W. P. Reconstructing ancestral character states under Wagner parsimony. \u003cem\u003eMath Biosci\u003c/em\u003e \u003cstrong\u003e87\u003c/strong\u003e, 199\u0026ndash;229 (1987).\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"scientific-reports","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"scirep","sideBox":"Learn more about [Scientific Reports](http://www.nature.com/srep/)","snPcode":"","submissionUrl":"","title":"Scientific Reports","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Scientific Reports","inReviewEnabled":true,"inReviewRevisionsEnabled":true},"keywords":"Camp dels Ninots, Maar, Konservat-Lagerstätten, Spain, Fossorial","lastPublishedDoi":"10.21203/rs.3.rs-6215385/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6215385/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThe Pliocene Konservat-Lagerst\u0026auml;tten maar lake site of Camp dels Ninots (NE Iberian Peninsula) has recently delivered a partial skeleton of a mole (family Talpidae) with many elements in anatomical connection. At a first glance, molar and humerus size, geological time interval, and geographical location suggested that this specimen could correspond to \u003cem\u003eTalpa minor\u003c/em\u003e. However, after some mechanical preparation of the clay block (matrix removal, consolidation, and cleaning) and a micro-CT scan, this excellently preserved specimen turned out to be an unknown species to science. The resulting 3D models of this new form, \u003cem\u003eVulcanoscaptor ninoti\u003c/em\u003e gen. et sp. nov., revealed some peculiar morphological traits in teeth, mandible, and postcranial elements, which according to the phylogenetic analysis carried out, would allocate this new species within the tribe Scalopini. This is surprising, because the representatives of this tribe are nowadays restricted to North America and Asia, and only some taxa had been previously reported in the Oligocene and Miocene fossil record from Europe. The postcranial construction of this specimen reveals a highly fossorial lifestyle supported by a complex forelimb structure. How such a specialized digging animal reached the maar lake sediments where it was finally preserved is still to be solved. Some hypotheses consider swimming abilities for this extinct species. Alternatively, this specimen could be the remaining portions of a floated or scavenged carcass whose remains fell into the lake and reached the anoxic bottom.\u003c/p\u003e","manuscriptTitle":"An unexpected New World mole (Scalopini, Mammalia) from the Pliocene of Europe sheds light on the phylogeny of talpids","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-04-07 09:47:27","doi":"10.21203/rs.3.rs-6215385/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2025-04-17T12:13:12+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-04-14T15:58:02+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-04-06T09:42:58+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"317273042357040628574096689475388142996","date":"2025-03-28T11:37:20+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"277248401448356075452051282302912809951","date":"2025-03-28T10:27:09+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-03-28T09:03:29+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-03-28T08:57:11+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"","date":"2025-03-19T11:11:54+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-03-19T11:10:59+00:00","index":"","fulltext":""},{"type":"submitted","content":"Scientific Reports","date":"2025-03-13T00:52:58+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"scientific-reports","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"scirep","sideBox":"Learn more about [Scientific Reports](http://www.nature.com/srep/)","snPcode":"","submissionUrl":"","title":"Scientific Reports","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Scientific Reports","inReviewEnabled":true,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"48b5b66f-01f9-40e5-a97b-baf75cd2e210","owner":[],"postedDate":"April 7th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[{"id":46478368,"name":"Biological sciences/Evolution/Palaeontology"},{"id":46478369,"name":"Biological sciences/Evolution/Phylogenetics"},{"id":46478370,"name":"Biological sciences/Evolution/Taxonomy"}],"tags":[],"updatedAt":"2025-07-14T16:03:13+00:00","versionOfRecord":{"articleIdentity":"rs-6215385","link":"https://doi.org/10.1038/s41598-025-10396-1","journal":{"identity":"scientific-reports","isVorOnly":false,"title":"Scientific Reports"},"publishedOn":"2025-07-10 15:57:46","publishedOnDateReadable":"July 10th, 2025"},"versionCreatedAt":"2025-04-07 09:47:27","video":"","vorDoi":"10.1038/s41598-025-10396-1","vorDoiUrl":"https://doi.org/10.1038/s41598-025-10396-1","workflowStages":[]},"version":"v1","identity":"rs-6215385","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6215385","identity":"rs-6215385","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}