identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
7B23EDC9FA205A76932CD302BDAECBD3.text	7B23EDC9FA205A76932CD302BDAECBD3.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Democricetodon decipiens (Freudenthal & Daams 1988)	<div><p>Democricetodon decipiens (Freudenthal &amp; Daams, 1988)</p><p>Figs 5, 6, 7</p><p>Localities.</p><p>MCX 1, MCX 3, MTR 1, MTR 2, BC 1, FS 1, MAB 0 A, MAB 0 B, MAB 3, MAB 5, MAB 10, MAB 11, MAB 13, CBR 0 B, and CBR 1.</p><p>Material (number of remains).</p><p>Suppl. material 1; MCX 1 (2): 2 M 1; MCX 3 (1): 1 m 1; MTR 1 (2): 2 m 2; MTR 2 (11): 1 m 1, 3 m 2, 1 m 3, 2 M 1, 3 M 2, 1 M 3; BC 1 (2): 1 M 3, 1 m 1; MAB 0 A (3): 1 m 1, 1 m 2, 1 M 2; MAB 0 B (1): 1 m 3; FS 1 (3): 1 M 2, 1 M 3, 1 m 3; MAB 3 (85): 15 m 1, 10 m 2, 13 m 3, 15 M 1, 19 M 2, 13 M 3; MAB 5 (69): 15 m 1, 11 m 2, 4 m 3, 12 M 1, 14 M 2, 13 M 3; MAB 10 (1): 1 m 1; MAB 11 (31): 4 m 1, 7 m 2, 2 m 3, 6 M 1, 8 M 2, 4 M 3; MAB 13 (1): 1 M 3; CBR 0 B (4): 1 m 2, 3 M 2 (2); CBR 1: 1 m 1, 1 m 2.</p><p>Measurements.</p><p>Suppl. material 2.</p><p>Description.</p><p>(See Suppl. material 3).</p><p>m 1 (MAB 3; Fig. 5 C – F): the anteroconid is simple and it may be located close to the metaconid (8 out of 13) or slightly further away (5 out of 13), and they may be contacting (2 out of 13) or not (11 out of 13). The labial anterolophid is low and may be in contact with the base of the protoconid (11 out of 12) or not (1 out of 12). The anterolophulid may be short (10 out of 15), connected to the protoconid (3 out of 15), or absent (2 out of 15). The metalophulid may be anterior (13 out of 15) or transverse (2 out of 15). The hypolophulid is anterior. The mesolophid may be short (2 out of 15), medium (4 out of 15), long (7 out of 15), or absent (2 out of 15). The ectomesolophid is absent. The sinusid is directed anteriorly, and may be closed by a ridge (5 out of 12), not closed by a ridge (1 out of 12), or have neither ridge nor cusp (6 out of 12). The posterolophid may connect with the entoconid (5 out of 13) or not (8 out of 13).</p><p>Variability in other sites: In MCX 3 the metalophulid is absent and the hypolophulid is transverse. In MTR 2 (Fig. 5 A), the hypolophulid is transverse. In BC 1 (Fig. 5 B), the posterolophid has a posterior ridge; in MAB 5 (Fig. 5 G, H), the anteroconid is typically closer to the protoconid. In one specimen, the anterolophulid contacts the metaconid. The metalophulid is anterior or absent, and the mesolophid is invariably present. In MAB 10 (Fig. 5 I), the anterolophulid does not contact the metalophulid. In MAB 0 A, MAB 11 (Fig. 5 J), and CBR 1, there are no significant morphological differences from the m 1 of MAB 3. Biometrically, the earliest material from the Ribesalbes-Alcora Basin, categorised as belonging to the local L. florancei biozone, exhibits a slightly larger size relative to more recent material classified as the L. ellipticus biozone (Fig. 6 A, Suppl. material 4).</p><p>m 2 (MAB 3; Fig. 5 O – R): the labial anterolophid contacts the anterolabial edge of the protoconid. The lingual anterolophid may be weak and fused to the metaconid (7 out of 10) or absent (3 out of 10). The metalophulid and hypolophulid are anterior. The mesolophid may be short (7 out of 9) or medium (2 out of 9) in length. The ectomesolophid is absent. The sinusid is directed anteriorly. The labial mesocingulum may be a ridge (5 out of 7), cusp-shaped (1 out of 7), or neither cusp-shaped nor ridge (1 out of 7). The posterolophid may be either connected to the entoconid (5 out of 8) or not (3 out of 8). The posterolophid has a posterior ridge.</p><p>Variability in other sites: In MTR 2 (Fig. 5 L – N), there is one specimen with an incipient ectomesolophid. In MAB 5 (Fig. 5 S), the labial anterolophid may or may not contact the labial mesocingulid around the protoconid, in another the lingual anterolophid is long, in another the sinusid is perpendicular. Finally, two specimens have no posterior crest of the posterolophid. In MAB 11 (Fig. 5 T, U), there is one specimen in which the labial anterolophid contacts the mesocingulum. There are no significant morphological differences in MTR 1, MAB 0 A, CBR 1, and CBR 0 B (Fig. 5 V). Biometrically, the older material from the Ribesalbes-Alcora Basin, belonging to the local L. florancei biozone, is similar to the more modern material belonging to the L. ellipticus biozone (Fig. 6 B, Suppl. material 4).</p><p>m 3 (MAB 3; Fig. 5 Z – AC): the labial anterolophid may connect to the antero-labial edge of the protoconid (10 out of 11) or connect to the labial mesocingulum (1 out of 11). The lingual anterolophid may be medium (1 out of 11), short (6 out of 11), or incipient (4 out of 11). The mesolophid is mostly absent, with only one specimen showing an incipient one. The metaconid and posterolophid are connected. The mesosinusid is occluded by a ridge (6 out of 10) or opens to the labial side unobstructed (4 out of 10).</p><p>Variability in other sites: In MTR 2 (Fig. 5 W), the mesolophid is double, with the anterior one being incipient. In MAB 0 B (Fig. 5 X), the mesolophid is incipient, with the posterior part of the tooth narrower than in other sites. In FS 1 (Fig. 5 Y), there is an incipient mesolophid, and the sinusid has a cusp on the labial edge. In MAB 5 (Fig. 5 AD – AE), the labial anterolophid may be short or absent as well as long, the lingual anterolophid may be long, and the metaconid and posterolophid are not connected in two specimens. In MAB 11 (Fig. 5 AF), there is one specimen with a strong lingual anterolophid, and the mesosinusid has a cusp on the labial margin. A biometric analysis reveals a decrease in size of the specimens over time in the Ribesalbes-Alcora Basin (Fig. 6 C, Suppl. material 4).</p><p>M 1 (MAB 3; Fig. 7 C – F): The anterocone may be simple (13 out of 14) or shallowly divided (1 out of 14), with a platform on the anterior side. The labial part of the anterocone may be larger than the lingual part (12 out of 13) or they are of similar size (1 out of 13). The labial anteroloph may connect to the paracone (12 out of 13), but in one specimen, the labial anteroloph runs from the anterocone to the mesosinus. The lingual anteroloph may connect with the protocone (7 out of 13) or not (6 out of 13). The anteroloph connects the lingual part of the anterocone with the protocone. The labial ridge of the anterolophule may be incipient (4 out of 15) or absent (11 out of 15). The protoloph may be double (1 out of 14), almost double with incipient or short anterior protoloph (3 out of 14), or posterior, connecting with the longitudinal ridge near the protocone (10 out of 14). The metaloph may connect with the posteroloph (11 out of 13) or the hypocone (2 out of 13). The mesoloph may be short (3 out of 13), medium (3 out of 13), or long (7 out of 13). The mesostyle may be present (6 out of 9) or not present (3 out of 9). The ectoloph may be incipient (6 out of 13) or absent (7 out of 13). The metacone ridge is absent. The entostyle may take the form of a cusp (3 out of 11) or a cingulum (8 out of 11). The posterosinus may be small (4 out of 9), medium (1 out of 9), long (2 out of 9), or absent (2 out of 9). The posteroloph may be connected with the metacone (6 out of 9) or not (3 out of 9).</p><p>Variability in other sites: In MCX 1 (Fig. 7 A), the metalophule is absent. In MTR 2 (Fig. 7 B), a ridge may be present in front of the anterocone. In MAB 5 (Fig. 7 G), there is a decrease in the number of individuals where the contact between the lingual anteroloph and protocone is present, and there is one specimen with a short labial crest of the anteroloph. The ectoloph is less common; there is one M 1 with a metacone ridge, and the posterosinus is smaller. In MAB 11 (Fig. 7 H – J), two specimens were observed with a ridge in front of the anterocone, the lingual anteroloph consistently contacts the protocone, and the labial ridge of the anteroloph may be double (2 out 6), with the anterior one developed and the posterior one incipient (2 out 6) or absent (2 out 6). There are two specimens with a double protolophule. Two specimens exhibit a double metalophule, one of these with the formation of a double mesoloph, and the other one surrounding the mesoloph. In the latter, the surrounding mesoloph is longer. The ridge of the metacone is present in two specimens, and the posterosinus is smaller. Biometrically, there is a slight tendency for specimens to become larger over time (Fig. 6 D, Suppl. material 4).</p><p>M 2 (MAB 3; Fig. 7 M – P): the lingual anteroloph may be of three distinct lengths: it may be long and reach the antero-lingual border of the protocone (9 out of 16), of medium length (1 out of 16), or short (9 out of 16). The labial anteroloph may either be long and connect to the paracone (15 out of 17) or be disconnected (2 out of 17). The protolophule may be double (7 out of 17), and there are two protolophule, but the posterior one is incomplete (9 out of 17) or simple and connected to the antero-labial part of the protocone (1 out of 17). The metalophule may be anterior (9 out of 18), posterior (2 out of 18), or there are two metalophules, but the anterior one is incomplete (1 out of 18), double (1 out of 18), or disconnected from the metaconule (1 out of 18). The mesoloph may be long, contacting the labial border (3 out of 18), long (12 out of 18), or medium (3 out of 18). The ectoloph may be present (5 out of 16) or absent (11 out of 16). The metaconal ridge may be present (2 out of 17) or not (15 out of 17). The lingual mesocingulum may close the sinus (10 out of 15) or not (5 out of 15). The mesostyle is present in 9 of 16 specimens. The sinus is transverse. The posteroloph may either connect to the metacone (15 out of 17) or not (2 out of 17).</p><p>Variability in other sites: In MTR 2 (Fig. 7 K, L), the sinus is posterior in one specimen. In MAB 5 (Fig. 7 Q), the lingual anteroloph is shorter, and the metalophule may be transverse. MAB 11 (Fig. 7 R – T) has one specimen with a posterior metalophule. No significant morphological differences were observed in MAB 0 A, CBR 0 B, and FS 1. Biometrically, the Ribesalbes-Alcora Basin sites were found to be similar, except for MAB 11, where they were generally narrower (Fig. 6 E, Suppl. material 4).</p><p>M 3 (MAB 3; Fig. 7 W – Z): The labial anteroloph may reach the antero-labial side of the paracone (6 out of 12), or not (6 out of 12), it may be long (7 out of 11) or of medium length (4 out of 11). The lingual anteroloph reaches the protocone base; it may be long (2 out of 12), medium in length (5 out of 12), short (4 out of 12), or platform-shaped (1 out of 12). The hypocone may be either large (3 out of 12) or small (9 out of 12). The metacone’s position relative to the tooth’s ridge is another distinguishing factor: it may be incorporated within the ridge surrounding the tooth (8 out of 10) or not (2 out of 10). The metalophule is connected to the anterior ridge of the hypocone (7 out of 11), does not reach the metacone (1 out of 11), is connected to the neo-entoloph (2 out of 11), or connects to the anterior ridge of the hypocone and the axioloph (1 out of 11). The mesoloph may be long (1 out of 13), short (7 out of 13), incipient (1 out of 12), or absent (2 out of 12). The axioloph may be long (3 out of 12), long with contact to the paracone / protolophule (3 out of 12), short (3 out of 12), incipient (1 out of 12), or absent (2 out of 12). The sinus may be relatively long (3 out of 12), medium (4 out of 12), or short (5 out of 12). The mesosinus may be wide (4 out of 12) or narrow (8 out of 12).</p><p>Variability in other sites: The mesoloph contacts the paracone in MTR 2 (Fig. 7 U). In MAB 5 (Fig. 7 AA – AC), the hypocone may be absent, and the mesoloph is more often absent. In MAB 11 (Fig. 7 AD), there is one specimen without a hypocone, and the mesoloph is shorter. No significant morphological differences were observed in BC 1 (Fig. 7 V), MAB 13, and FS 1. A biometric analysis reveals that the different sites in the Ribesalbes-Alcora Basin are morphologically similar (Fig. 6 F, Suppl. material 4).</p><p>Remarks.</p><p>The taxonomic status of the genus Democricetodon has been the subject of extensive debate in the literature (Kälin 1999; van der Meulen et al. 2003; Casanovas-Vilar 2007; Jovells-Vaquè and Casanovas-Vilar 2018 a). This debate has encompassed a range of perspectives, including the proposal of the existence of multiple genera, such as Democricetodon, Fahlbuschia, Pseudofahlbuschia, and Renzimys (Freudenthal and Daams 1988; Freudenthal 2006; Ruiz-Sánchez et al. 2013). Conversely, alternative perspectives advocate for the exclusive consideration of Democricetodon and Fahlbuschia as synonyms (Aguilar 1981; Aguilar et al. 2010 a). A third viewpoint suggests the avoidance of such deliberations (Murelaga et al. 2008; de Bruijn 2010; Prieto et al. 2010, among others). However, some scholars have proposed that all the aforementioned genera be subsumed under the genus Democricetodon (van der Meulen et al. 2003; Álvarez-Sierra et al. 2006; Fejfar et al. 2011; Casanovas-Vilar et al. 2016; García-Paredes et al. 2016; among others). The latter hypothesis is the one that is more widely accepted, and, as a consequence, will be followed in this publication, as it is not a topic of discussion.</p><p>The provenance of the genus under consideration may be located in Anatolia, with the first records in Turkey immediately following the MN 1. In contrast, the earliest evidence of the genus in China and Europe is not until MN 3 / 4 (Fejfar et al. 2011). Alternatively, the genus may have originated in Central Asia (Maridet et al. 2011; Flynn and Wessels 2013) and arrived in Europe at the end of MN 3 (van der Meulen et al. 2012; among others).</p><p>This genus is clearly distinguishable from the species of muroids present in the Ribesalbes-Alcora Basin, which are assigned to Megacricetodon and Eumyarion, by virtue of its intermediate size and more rounded teeth. Furthermore, Democricetodon differs from Megacricetodon in having an undivided anterocone of M 1 and a generally more complex tooth pattern. Compared to Eumyarion, it exhibits a more simplified and advanced pattern (Fejfar 1999). A further distinction is the development of the cusps of the labial side of the upper molars and the lingual part of the lower molars, which are higher than those of the other region (Figs 2 J, 5 K). In the remaining genera, there is typically less variation in the height of the cusps.</p><p>The material under discussion is compared with that described by Freudenthal and Daams (1988), van der Meulen et al. (2003), Freudenthal (2006), and Jovells-Vaquè and Casanovas-Vilar (2021). In addition, a comparison is made with the measurements of Democricetodon franconicus Fahlbusch, 1966 by Fahlbusch (1966).</p><p>In the M 1 from the localities of the Ribesalbes-Alcora Basin, the anterocones are simple, as in the species D. hispanicus, D. decipiens, Democricetodon moralesi van der Meulen et al., 2003, Democricetodon sacedonensis Freudenthal, 2006, and D. franconicus . About the protolophid, while this character is predominantly double in D. hispanicus and the older populations of D. franconicus, it is less represented (“ morphotype double ”) in D. moralesi and D. sacedonensis, the percentages of representation or morphotypes in the Ribesalbes-Alcora Basin are very different in D. moralesi and D. sacedonensis, similar to those described for D. decipiens by van der Meulen et al. (2003) for the Calatayud-Montalbán and Buñol basins. The length of the mesoloph is very similar to that described for D. decipiens . It differs from that of D. hispanicus, D. sacedonensis, and D. franconicus, where it is usually medium-long, and also from that of D. moralesi, which is shorter. With regard to size (see Fig. 6), the material studied is generally larger than D. hispanicus from Villafeliche 2 A, smaller than D. moralesi from La Col D, slightly smaller than D. sacedonensis from Córcoles, and very similar to those of D. decipiens from Buñol and D. franconicus from Erkertshofen. About the L / W index of M 1, the studied material is within the range of variability observed in each of the aforementioned species (Table 1).</p><p>In the M 2, the mesolophs of the material from the Ribesalbes-Alcora Basin are of a medium-long length, as observed in D. franconicus, D. hispanicus, and D. decipiens . In this regard, the former exhibits a longer length than that observed in D. moralesi and D. sacedonensis . The ectoloph is absent in D. hispanicus and is very poorly represented in D. franconicus, D. decipiens, and D. moralesi, as observed in the localities studied. In the material under consideration, protolophules are predominantly double and metalophules are primarily anterior, as seen in D. decipiens, D. franconicus, and D. sacedonensis . The size of the material from the Ribesalbes-Alcora Basin is generally larger than that of D. hispanicus from Villafeliche 2 A, smaller than that of D. moralesi from La Col D, and similar to that of D. decipiens from Buñol, D. franconicus from Erkertshofen, and D. sacedonensis from Córcoles.</p><p>The M 3 from the Ribesalbes-Alcora Basin is comparable in size to D. hispanicus, D. franconicus, and D. decipiens, with a narrower width than D. sacedonensis and a smaller overall size than D. moralesi .</p><p>For the m 1, while in D. hispanicus the anteroconid-metaconid contact is invariably present, in D. decipiens it is present in approximately half of the examined remains. At the same time, in D. franconicus it is highly variable. In D. moralesi and D. sacedonensis, the contact is present in only a few specimens. In the material studied, something similar to D. decipiens occurs, tending to have a higher percentage of connections in the localities belonging to the L. florancei local biozone. However, there are only a few specimens in these sites. On the other hand, in the Ribesalbes-Alcora Basin, the metalophulid is anterior, rarely transverse or absent, whereas in D. franconicus, the metalophulid is usually transverse, in D. sacedonensis transverse or anterior, and variable in the rest. The mesolophid of the material from the Ribesalbes-Alcora Basin is typically long, as in D. hispanicus, and shorter in D. decipiens, D. franconicus, D. sacedonensis, and D. moralesi . The remaining morphologies of m 1 are analogous to those previously documented for D. hispanicus, D. franconicus, D. sacedonensis, D. decipiens, and D. moralesi . With regard to size, the material from the Ribesalbes-Alcora Basin is typically larger than D. hispanicus from Villafeliche 2 A, smaller than D. moralesi from La Col D, and of the same size as D. decipiens from Buñol, D. sacedonensis from Córcoles, and D. franconicus from Erkertshofen. The L / W index indicates that the studied material falls within the variability of one of the described species (Table 1).</p><p>The mesolophid m 2 from the Ribesalbes-Alcora Basin material is characterised by either a short or, on rare occasions, a long form, with a length comparable to that of the species D. franconicus, D. hispanicus, D. decipiens, and D. moralesi, and shorter than that of D. sacedonensis . The size of the m 2 from the Ribesalbes-Alcora Basin is typically larger than those of D. hispanicus from Villafeliche 2 A, smaller than those of D. moralesi from La Col D, and D. sacedonensis from Córcoles, and very similar to those of D. decipiens from Buñol and D. franconicus from Erkertshofen.</p><p>The m 3 from the Ribesalbes-Alcora Basin exhibits mesolophids in the local L. florancei biozone localities, although the number of identified remains of this element is limited. Such occurrences are absent in D. sacedonensis and observed only infrequently in D. hispanicus and D. decipiens . The size of the m 3 in this study is generally larger than that of D. hispanicus from Villafeliche 2 A, smaller than that of D. moralesi from La Col D, not as wide as that of D. sacedonensis from Córcoles, and very similar in size to that of D. decipiens from Buñol and D. franconicus from Erkertshofen.</p><p>The material under study exhibits notable disparities compared with other European species of the genus Democricetodon of the same age. It is distinguished from Democricetodon anatolicus Theocharopoulos, 2000 by its larger size and more robust anteroconid (Theocharopoulos 2000); and from Democricetodon doukasi Theocharopoulos, 2000 by its larger size, less developed M 1 -2 ectoloph of the M 1 -2, and longer mesolophids (Theocharopoulos 2000); from Democricetodon affinis (Schaub, 1925) is distinguished for being smaller, having shorter mesolophs, longer mesolophids and the anterocone exhibiting a slight division (Maridet 2003); from Democricetodon brevis (Schaub, 1925) from MN 6, of which there is a citation in MN 4 from Port la Nouvelle (Aguilar et al. 1999) for having a less divided anterocone and shorter mesoloph / phids (Maridet 2003); from Democricetodon gracilis (Fahlbusch 1964) for having shorter mesolophs and larger size (Maridet 2003); and from Democricetodon mutilus (Fahlbusch, 1964) because the protolophule of M 1 is usually double or nearly double, the mesolophule is shorter, the lingual valleys of m 1 and 2 are not as anterior, and m 3 usually has an entoconid and is smaller (Maridet 2003); from Democricetodon gaillardi (Schaub, 1925) due to the smaller size, the shorter mesoloph / ids, the less elongated m 1, and because the anteroconid is weaker (Maridet, 2003); from Democricetodon freudenthali (Antunes and Mein 1981) because of the smaller size and the longer mesolophs (Antunes and Mein 1981); from Democricetodon romieviensis (Freudenthal, 1963) for being larger, having a less developed double protolophule of M 1 and metalophule of M 2, and for having shorter mesoloph / ids (Daams and Freudenthal 1974).</p><p>Compared with all previously documented Democricetodon species, it can be concluded that the Democricetodon material from the Ribesalbes-Alcora Basin localities can be assigned to D. decipiens . Its morphology and size fall within the range of intrapopulation variability, and it exhibits novel morphological features not observed in any other species of the genus, including the mesolophs and the double labial ridges of the anteroloph of MAB 11.</p></div>	https://treatment.plazi.org/id/7B23EDC9FA205A76932CD302BDAECBD3	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Crespo, Vicente D.;Ríos, María;Marquina-Blasco, Rafael;Montoya, Plini	Crespo, Vicente D., Ríos, María, Marquina-Blasco, Rafael, Montoya, Plini (2025): The Early Miocene muroids (Muroidea, Rodentia) of the Ribesalbes-Alcora Basin (Spain): A thriving haven during a time of migration. Fossil Record 28 (1): 187-218, DOI: 10.3897/fr.28.138478
54385458995854BE8DB1539FA71E8762.text	54385458995854BE8DB1539FA71E8762.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Democricetodon Fahlbusch 1964	<div><p>Genus Democricetodon Fahlbusch, 1964</p><p>Type species.</p><p>Democricetodon crassus Freudenthal, 1969 . Sansan, Middle Miocene.</p></div>	https://treatment.plazi.org/id/54385458995854BE8DB1539FA71E8762	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Crespo, Vicente D.;Ríos, María;Marquina-Blasco, Rafael;Montoya, Plini	Crespo, Vicente D., Ríos, María, Marquina-Blasco, Rafael, Montoya, Plini (2025): The Early Miocene muroids (Muroidea, Rodentia) of the Ribesalbes-Alcora Basin (Spain): A thriving haven during a time of migration. Fossil Record 28 (1): 187-218, DOI: 10.3897/fr.28.138478
7E46BB1091C65B429709B21C5A5003B6.text	7E46BB1091C65B429709B21C5A5003B6.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Eumyarion Thaler 1966	<div><p>Genus Eumyarion Thaler, 1966</p><p>Type species.</p><p>Cricetodon medium Lartet, 1851 (= Cricetodon helveticum Schaub, 1925). Sansan, Middle Miocene.</p></div>	https://treatment.plazi.org/id/7E46BB1091C65B429709B21C5A5003B6	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Crespo, Vicente D.;Ríos, María;Marquina-Blasco, Rafael;Montoya, Plini	Crespo, Vicente D., Ríos, María, Marquina-Blasco, Rafael, Montoya, Plini (2025): The Early Miocene muroids (Muroidea, Rodentia) of the Ribesalbes-Alcora Basin (Spain): A thriving haven during a time of migration. Fossil Record 28 (1): 187-218, DOI: 10.3897/fr.28.138478
604F7AFCF92852BB85D8BE950A8B8191.text	604F7AFCF92852BB85D8BE950A8B8191.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Eumyarion weinfurteri (Schaub & Zapfe 1953)	<div><p>Eumyarion weinfurteri (Schaub &amp; Zapfe, 1953)</p><p>Figs 8 A – T, 9</p><p>Localities.</p><p>FS 1, MAB 3, and MAB 5.</p><p>Material (number of remains).</p><p>Suppl. material 1; FS 1 (1): 1 m 2; MAB 3 (21): 3 M 1, 1 M 2, 3 M 3, 3 m 1, 6 m 2, 5 m 3; MAB 5 (8): 1 M 1, 1 M 3, 2 m 1, 2 m 2, 2 m 3.</p><p>Measurements.</p><p>Suppl. material 2.</p><p>Description.</p><p>m 1 (MAB 3; Fig. 8 A, B): the anteroconid is simple. The anterolophulid may be complete (1 out of 3), low (1 out of 3), or incomplete (1 out of 3). In the latter case, the anterolophulid is a ridge that starts from the protoconid. A ridge begins from the metaconid and may contact the anteroconid (2 out of 3) or not (1 out of 3). The anterior metalophulid is directed slightly forward, and it may either connect with the anterolophulid (2 out of 3) or not (1 out of 3). The posterior metalophulid may contact with the posterior ridge of the protoconid (1 out of 3), there is only a spur (1 out of 3), or absent (1 out of 3). The protosinusid is almost closed by a low ridge coming from the anteroconid. The anterosinusid may be well developed (1 out of 3) or practically disappear by the anterior spur of the metaconid (2 out of 3). The posterior ridge of the protoconid may be short, and does not connect with the mesolophid (1 out of 3) or is of medium size and contacts the mesolophid (2 out of 3). The mesolophid is of medium length. The ectomesolophid may reach the labial margin (2 out of 3) or be absent (1 out of 3). The hypolophulid is transverse and connects with the ectolophid. The posterior ridge of the hypoconid may be present (2 out of 3) or not (1 out of 3), resulting in a high posterolophid connected to the entoconid, which encloses the posterosinusid.</p><p>Variability in other sites: in MAB 5 (Fig. 8 C, D), the anterior ridge of the metaconid does not contact the anteroconid, the anterior metalophulid is a spur, while the posterior one is directed towards the mesolophid in one specimen. In one, the mesolophid is long, while the ectomesolophid is either a spur or absent. The m 1 from MAB 5 is comparatively larger than that of MAB 3 (Fig. 9 A).</p><p>m 2 (MAB 3; Fig. 8 F, G): the lingual anterolophid is of medium length, and it may not reach the lingual margin (4 out of 5) or be absent (1 out of 5). The anterosinusid is closed by the anterolophid (4 out of 5) or absent (1 out of 5). The labial anterolophid connects to the base of the protoconid. The metalophulid is simple and connects to the anterolophid. On the lingual side, the posterior ridge of the metaconid connects with the lingual part of the posterior ridge of the protoconid (4 out of 5), or not (1 out of 5), almost completely enclosing the mesosinusid. The posterior ridge of the protoconid is of medium length, and two specimens contact the mesolophid. The mesolophid is medium length (1 out of 6), short (4 out of 6), or absent (1 out of 6). The ectomesolophid may be short (1 out of 5), medium (1 out of 5), or absent (3 out of 5). The posterior crest of the hypoconid is long (2 out of 6) or only a spur (4 out of 6). The posterolophid is high and connects with the entoconid, enclosing the posterosinusid (4 out of 6) or not contacting the entoconid (2 out of 6).</p><p>Variability at other sites: FS 1 (Fig. 8 E) exhibits no notable differences. However, in MAB 5, the lingual anterolophid is shorter (Fig. 8 H, I). Additionally, the m 2 of the study material appears to lengthen with time in terms of biometry (Fig. 9 B).</p><p>m 3 (MAB 3; Fig. 8 J, K): the lingual anterolophid is short, and the labial one connects with the base of the protoconid. The metalophulid contacts obliquely forward. On the lingual side, the posterior ridge of the metaconid connects with the anterior ridge of the protoconid and with the entoconid, enclosing the mesosinusid. The posterior ridge of the protoconid is of medium-long length. The ectomesolophid may be short (1 out of 5) or absent (4 out of 5). The hypolophulid is transverse and complete. The posterolophid is high and connects with the entoconid, enclosing the posterosinusid.</p><p>Variability in other sites: in MAB 5 (Fig. 8 L, M), the posterior ridge of the protoconid is more delayed. However, no difference in size is apparent (Fig. 9 C).</p><p>M 1 (MAB 3; Fig. 8 N, O): The tooth surface is rough. The anterocone is simple and extends transversely. One specimen has an anterior ridge on the labial side of the anterocone. The postero-labial ridge of the anterocone is attached to the labial side of the tooth in one individual, and in another it is isolated. The anterocone has a central posterior ridge: in one specimen it is directed towards the labial side, in another towards the protocone, and in the last one it is straight. The anteroloph may be present (2 out of 3) or absent (1 out of 3). The labial ridge of the anterolophule may be incipient (1 out of 3) or absent (2 out of 3). The anterior ridge of the protocone is directed forward and, in one specimen, connects with the median ridge of the anterocone. The posterior protoloph is transverse and connects labially to the entoloph, posterior to the protocone. The ectoloph may be present (2 out of 3) or absent (1 out of 3). In one individual, there is a double mesoloph. In the latter case, the anterior mesoloph is incipient and is directed towards the posterior mesoloph. The posterior or main mesoloph is long. The ectomesoloph may be absent (2 out of 3) or only a spur (1 out of 3). The sinus is relatively narrow and directed forward. The metaloph is connected to the entoloph. The posteroloph is long, and it may connect to the base of the metacone (1 out of 3) or not (2 out of 3).</p><p>Variability is present in other sites: in MAB 5 (Fig. 8 P), the absence of a posterior ridge of the anterocone is observed, and the metaloph is connected to the anterior part of the hypocone. No significant disparities in size are detected (Fig. 9 D).</p><p>M 2 (MAB 3; Fig. 8 Q): The surface is rough. The lingual anteroloph is short. The protoloph is transverse and connects with the protocone. The mesoloph is of medium length. The ectomesoloph is absent. The sinus is narrow and directed forward. The metaloph is transverse and connects with the hypocone. The posteroloph is long and isolated labially.</p><p>M 3 (MAB 3; Fig. 8 R – T): the labial anteroloph is characterised as either long (2 out of 3) or medium (1 out of 3) in length, with the possibility of either connecting to the protocone (1 out of 3) or the protoloph (2 out of 3). The lingual anteroloph is either small (2 out of 3) or absent (1 out of 3). The protoloph is simple, transverse, or shifted slightly forward, connecting to the anterior part of the protocone. The posterior ridge of the paracone connects to the base of the metacone, forming a labial cingulum that closes the mesosinus. The neo-entoloph may be complete and high (1 out of 3), short and weak (1 out of 3), or incomplete (1 out of 3). The axioloph may be complete and located in the central part of the tooth (2 out of 3), or long and incomplete (1 out of 3). The mesoloph may be medium (1 out of 3) or long and connects to the labial cingulum (2 out of 3). The metaloph is long and directed forward. The metaloph may connect to the metacone and the neo-entoloph through the axioloph (1 out of 3), the metacone and the posterior part of the protocone (1 out of 3), or the anterior part of the hypocone (1 out of 3). The hypocone may be well developed (1 out of 3) or absent (2 out of 3). The posteroloph may be high and short, connecting the neo-entoloph and the metacone, enclosing a small posterosinus (1 out of 3), which is small and isolated (1 out of 3), or high, short, and not connected to the metacone (1 out of 3).</p><p>Variability is present in other sites: the tooth from MAB 5 is similar to the one described before.</p><p>Remarks.</p><p>The genus Eumyarion is a small to medium-sized muroid characterized by primitive dental features (Thaler 1966). The origins of this genus are complex and likely related to Eucricetodon from the Oligocene and Early Miocene (Kälin 1999). A comprehensive revision of both genera could potentially result in their synonymy, encompassing some basal forms of Eumyarion (Kälin 1999; de Bruijn et al. 2013).</p><p>The provenance of this genus has been the subject of considerable debate among scholars, with various authors proposing a range of potential locations. These include the upper Oligocene in Turkey (Ünay et al. 2003), the MN 1 in Asia (Lopatin 2004 a; Maridet et al. 2011), and the upper Oligocene in Europe, more specifically in Bosnia and Herzegovina, have all been proposed as potential locations for the origin of this genus. (de Bruijn et al. 2013). The earliest known fossil of this species from Central and Western Europe was discovered at Dolnice 1 in the Cheb Basin in the Czech Republic and dates back to the first half of the lower Aragonian (MN 4; Fejfar 1989). During the Early-Middle Miocene in Europe, a variety of species of Eumyarion were present, namely Eumyarion bifidus (Fahlbusch, 1964), Eumyarion latior (Schaub &amp; Zapfe, 1953), Eumyarion medius (Lartet, 1851), and E. weinfurteri (which may be a synonym of E. latior, according to de Bruijn (2009)). In addition, within the geographical confines of Turkey, the following species have been identified: Eumyarion carbonicus de Bruijn &amp; Saraç, 1991, Eumyarion orhani de Bruijn et al., 2006, Eumyarion montanus de Bruijn &amp; Saraç, 1991, Eumyarion beyderensis Bilgin et al., 2023, Eumyarion aegeaniensis Bilgin et al., 2023, and Eumyarion gordesensis Peláez-Campomanes et al., 2019 were also discovered (de Bruijn 2009; Peláez-Campomanes et al. 2019; Bilgin et al. 2023).</p><p>In the Iberian Peninsula, the earliest fossil records date from the lower Aragonian (upper part of the MN 4), specifically at the localities of Buñol (Cerro de la Cruz, Magro River Basin, last part of the local biozone C; Daams and Freudenthal (1974); local biozone according to Crespo et al. 2019), Montalvos 2 (Teruel Basin, local biozone Ca, Hordijk et al. 2015), Vargas 4 A (Calatayud-Montalbán Basin, local biozone Cb, van der Meulen et al. 2012), and Les Cases de la Valenciana, Can Martí Vell, Els Casots or San Mamet (Vallès-Penedès Basin, local biozone C, Casanovas-Vilar et al. 2016, Jovells-Vaquè et al. 2017). During the Early Miocene appear in the Iberian Peninsula, the species E. medius, Eumyarion valencianum Daams &amp; Freudenthal, 1974 (nomem dubium in de Bruijn and Saraç (1991), probably E. weinfurteri or E. latior according to de Bruijn 2009) and E. weinfurteri (Daams and Freudenthal 1974; Ruiz-Sánchez et al. 2003; Casanovas-Vilar et al. 2016), and subsequently, Eumyarion leemani (Hartenberger, 1965) (Hartenberger 1965; Casanovas-Vilar 2007).</p><p>The classification of medium-sized Eumyarion poses a considerable challenge, primarily due to the extensive intraspecific variation observed in their morphology and size. This observation has prompted several researchers to synonymise species and question the homogeneity of populations (Engesser 1972; Vasileiadou and Zouros 2012). A particularly illustrative example of this phenomenon is E. weinfurteri, a species initially synonymised with E. latior by de Bruijn and Saraç (1991), and subsequently by Schötz (1993). However, de Bruijn (2009) subsequently raised questions regarding its status, albeit identifying differences between the two in M 1 and M 2 of Sandelzhausen.</p><p>The material of the genus Eumyarion recovered from the Ribesalbes-Alcora Basin exhibits a larger size compared to the smaller species of the genus (namely Eumyarion intercentralis de Bruijn &amp; Saraç, 1991, Eumyarion microps de Bruijn &amp; Saraç, 1991), E. gordensensis, and E. orhani . It is slightly larger than the type population of E. weinfurteri (although resembling other populations of this species), E. bifidus, and E. carbonicus, and exhibits similarities with E. aegeaniensis, E. beyderensis, E. montanus, and E. medius . However, in comparison to E. latior, it is similar or slightly smaller, depending on the specific element (Fig. 9).</p><p>The MAB 3 sample exhibits a combination of primitive and derived features in the m 1 morphology, with one specimen displaying an anteroconid that is strikingly similar to that of E. orhani or Eumyarion tremulus Lopatin, 1996, and lacking protoconid-anteroconid contact. In the remaining material, the anteroconid has a decreasing crescent shape and it is connected to the metaconid and protoconid by a well-developed anterolophulid, a feature typical of E. carbonicus or E. latior / weinfurteri (Daams and Freudenthal 1974; Bulot 1979; de Bruijn et al. 2006). It differs from E. beyderensis because in this species the anteroconid is near to be isolated, and E. aegeaniensis by this more developed anteroconid (Bilgin et al. 2023). One specimen displays the distinctive characteristics of E. montanus from Keseköy, i. e., a short mesolophid and a posterior ridge of the protoconid (de Bruijn and Saraç 1991). Conversely, the remaining specimens exhibit a convergence of both ridges, a morphology consistent with that of E. latior / weinfurteri as observed in Buñol, Aliveri, and Dolnice (Daams and Freudenthal 1974; Hofmeijer and de Bruijn 1988; Fejfar 1989), or E. orhani (de Bruijn et al. 2006) . The occurrence of the posterior crest of the hypoconid varies significantly within the genus Eumyarion . It is present in our material in a comparable percentage to E. montanus, E. tremulus, or E. latior / weinfurteri (de Bruijn and Saraç 1991; Lopatin 2004 a; de Bruijn 2009). The morphological similarity of the m 1 of MAB 5 to the E. latior / weinfurteri populations from Buñol or Aliveri (Daams and Freudenthal 1974; Hofmeijer and de Bruijn 1988) is remarkable. This element is biometrically smaller than the type population of E. latior, and only MAB 5 is larger than the type population of E. weinfurteri, while the remains of MAB 3 are smaller than the type population of E. medius (Fig. 9 A).</p><p>The morphology of m 2 is distinctive. In 33 % of the specimens, the posterior ridge of the protoconid is present, a feature that is more commonly seen in more recent species. Conversely, the short or absent mesolophid is a characteristic of more primitive species. This combination of morphologies is a characteristic of E. latior / weinfurteri from Aliveri and Eumyarion margueritae from de Bruijn et al. 2013, from Banovići (Hofmeijer and de Bruijn 1988; de Bruijn et al. 2013). In terms of size, the specimens from the Ribesalbes-Alcora Basin are smaller than the type population of E. latior, somewhat larger than E. weinfurteri, and fall within the type variability shown for each species (Fig. 9 B).</p><p>The m 3 has no mesolophid, the posterior crest of the hypoconid is absent, and the posterior crest of the protoconid is medium to long. These characteristics are comparable to those found in E. latior / weinfurteri de Aliveri and E. margueritae, yet still shorter than those of the Buñol population (Daams and Freudenthal 1974; Hofmeijer and de Bruijn 1988; de Bruijn et al. 2013). With regard to dimensions, they are marginally larger than the E. weinfurteri type population (Fig. 9 C).</p><p>The M 1 is distinguished by the variability and complexity of the posterior spurs of the anterocone, exhibiting a morphology analogous to that observed in E. latior / weinfurteri populations from Aliveri or Dolnice 1 (Hofmeijer and de Bruijn 1988; Fejfar 1989). The number of specimens with the paracone spur not attached to the mesoloph is comparable to that observed in E. montanus and E. tremulus, exhibiting a medium mesoloph consistent with that seen in E. montanus (de Bruijn and Saraç 1991; Lopatin 2004 a). De Bruijn (2009) regards this as one of the distinguishing elements between E. latior and E. weinfurteri . According to the classification system proposed by de Brujin (2009), the material from the Ribesalbes-Alcora Basin’s “ BBB ” morphotype exhibits the characteristic morphologies of E. weinfurteri, including a lingual spur on the anterocone, no connection between mesoloph and metacone, and no paracone spur and mesoloph. The MAB 3 specimen is slightly larger than the E. weinfurteri type population and longer than E. bifidus (Fig. 9 D).</p><p>A single M 2 has been identified at the MAB 3 locality, and it is characterised by a small posterior paracone spur. This morphology is similar to that described in E. montanus from Keseköy and E. weinfurteri from Sandelzhausen or Bézian (Bulot 1979; de Bruijn and Saraç 1991; de Bruijn 2009), while in E. latior from Sandelzhausen or E. latior / weinfurteri from Buñol, this crest is more developed (Daams and Freudenthal 1974; de Bruijn 2009). The protoloph is single, in contrast to the double protoloph observed in E. orhani or E. bifidus (de Bruijn et al. 2006; de Bruijn 2009). The mesoloph is of medium length, a feature that is analogous to that of E. montanus (de Bruijn and Saraç 1991), yet it is comparatively shorter than in E. latior / weinfurteri (Daams and Freudenthal 1974; de Bruijn 2009). This morphology is another distinguishing factor between E. latior and E. weinfurteri (de Bruijn 2009; Vasileiadou and Zouros 2012). According to the classification proposed by Bruijn (2009), the material under study falls into the category “ BBB ”: characterised by a simple protoloph, with no connection between the metacone and mesoloph, nor between the paracone spur and the mesoloph. This category is typical of E. weinfurteri . The specimen of MAB 3 is of a size that falls between the type populations of E. weinfurteri and E. latior, coinciding with the variability of the rest of the species shown (Fig. 9 E).</p><p>The morphology of the third molar exhibits considerable variation, with the axioloph potentially being either complete with a long mesoloph or incomplete with a short mesoloph. The former combination has been observed in E. montanus or early populations of E. latior / weinfurteri (Daams and Freudenthal 1974; de Bruijn and Saraç 1991), while the latter is exclusively found in the MAB 3 specimens. With regard to biometrics, the M 3 of MAB 3 is larger than the type population of E. weinfurteri and smaller than that of E. medius, which is in agreement with the other species shown (Fig. 9 F).</p><p>As posited by de Bruijn (2009), in his seminal work on the subject, the material of the genus Eumyarion from the Ribesalbes-Alcora Basin must be ascribed to E. weinfurteri, albeit with a slightly larger size than that of the type population.</p></div>	https://treatment.plazi.org/id/604F7AFCF92852BB85D8BE950A8B8191	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Crespo, Vicente D.;Ríos, María;Marquina-Blasco, Rafael;Montoya, Plini	Crespo, Vicente D., Ríos, María, Marquina-Blasco, Rafael, Montoya, Plini (2025): The Early Miocene muroids (Muroidea, Rodentia) of the Ribesalbes-Alcora Basin (Spain): A thriving haven during a time of migration. Fossil Record 28 (1): 187-218, DOI: 10.3897/fr.28.138478
92F8A1CE7FE1509893323EDEBD1463B7.text	92F8A1CE7FE1509893323EDEBD1463B7.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Megacricetodon Fahlbusch 1964	<div><p>Genus Megacricetodon Fahlbusch, 1964</p><p>Type species.</p><p>Megacricetodon gregarious (Schaub, 1925), La Grive M, Upper Miocene.</p></div>	https://treatment.plazi.org/id/92F8A1CE7FE1509893323EDEBD1463B7	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Crespo, Vicente D.;Ríos, María;Marquina-Blasco, Rafael;Montoya, Plini	Crespo, Vicente D., Ríos, María, Marquina-Blasco, Rafael, Montoya, Plini (2025): The Early Miocene muroids (Muroidea, Rodentia) of the Ribesalbes-Alcora Basin (Spain): A thriving haven during a time of migration. Fossil Record 28 (1): 187-218, DOI: 10.3897/fr.28.138478
CA97BFB8994B5D1F8B931FF183CCAF9F.text	CA97BFB8994B5D1F8B931FF183CCAF9F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Megacricetodon primitivus (Freudenthal 1963)	<div><p>Megacricetodon primitivus (Freudenthal, 1963)</p><p>Figs 2, 3, 4</p><p>Localities.</p><p>MCX 2, MCX 3, MCX 4, MTR 1, MTR 2, MTR 3, BC 1, BC 2, FS 1, MAB 0 A, MAB 0 B, MAB 2, MAB 3, MAB 4, MAB 5, MAB 11, MAB 11 B, MAB 12, CBR 0 B, CBR 0 E, CBR 0 F, CBR 1, CBR 2, and CBR 4.</p><p>Material (number of remains).</p><p>Suppl. material 1; MCX 2 (1): 1 mandible, 1 m 1; MCX 3 (62): 1 mandible, 12 m 1, 9 m 2, 9 m 3, 10 M 1, 12 M 2, 9 M 3; MCX 4: 1 M 1; MTR 1 (3): 1 m 1, 2 M 2; MTR 2 (47): 13 m 1, 9 m 2, 3 m 3, 12 M 1, 5 M 2, 5 M 3; MTR 3 (2): 1 M 1, 1 M 2; BC 1 (51): 10 m 1, 8 m 2, 7 m 3, 15 M 1, 7 M 2, 4 M 3; BC 2 (1): 1 M 1; MAB 0 A (22): 3 m 1, 4 m 2, 1 m 3, 7 M 1, 4 M 2, 3 M 3; MAB 0 B (9): 2 m 1, 3 m 2, 2 m 3, 2 M 1; FS 1 (15): 3 m 1, 1 m 3, 3 M 1, 7 M 2, 1 M 3; MAB 2 (2): 1 m 1, 1 m 3; MAB 3 (92): 14 m 1, 20 m 2, 19 m 3, 17 M 1, 12 M 2, 10 M 3; MAB 3 A (1): 1 m 2; MAB 4 (1): 1 M 2; MAB 5 (85): 18 m 1, 13 m 2, 14 m 3, 14 M 1, 15 M 2, 11 M 3; MAB 11 (10): 1 m 1, 3 m 2, 1 m 3, 3 M 1, 2 M 2; MAB 11 B (1): 1 M 3; MAB 12 (1): 1 m 3; CBR 0 B (1): 1 m 2; CBR 0 E (1): 1 M 1; CBR 0 F (1): 1 m 2; CBR 1 (19): 1 m 1, 4 m 2, 2 m 3, 5 M 1, 4 M 2, 2 M 3; CBR 2 (1): 1 m 3; CBR 4 (1): 1 M 1.</p><p>Measurements.</p><p>Suppl. material 2.</p><p>Description.</p><p>(See Suppl. material 3).</p><p>Mandible (MCX 3; Fig. 2 A): In occlusal view, the mandible exhibits a slight inclination, which does not obscure the mental foramen. This foramen is located between the incisor and the first molar on the labial side. The masseteric process is deep and originates at the level of the anterior part of the m 1 with a ridge.</p><p>Concerning other sites, no significant disparities were observed in the material of MCX 2.</p><p>m 1 (MAB 5; Fig. 2 F – I): the anteroconid is simple and rounded, and slightly lower than the rest of the cusps (between morphology B and C as described by Oliver and Peláez-Campomanes (2016) (Fig. 2 H)). The labial spur of the anterolophulid may be developed (1 out of 16), incipient (1 out of 16), or absent (14 out of 16). The metalophulid is positioned anteriorly and is directed forward, not connected to the protoconid. The mesolophid may be categorised as follows: long and developed to the lingual side (1 out of 16), long without reaching the lingual side (4 out of 16), medium (8 out of 16), or short (3 out of 16). The ectomesolophid is absent. The hypolophulid is directed forward. The posterolophid descends towards the base of the entoconid but does not connect to it.</p><p>Variability in other sites: In the remains of MCX 2 and MCX 3 (Fig. 2 B), the anterolophulid is consistently absent, and the mesolophid is either short or medium. In MTR 1, the anterolophulid is absent. In the remains from MTR 2 (Fig. 2 C), two specimens have a double metalophulid, one has no mesolophid, one has an ectomesolophid and one has the posterolophid attached to the entoconid. In BC 1, the mesolophid is shorter (Fig. 2 D). In one individual, the posterolophid is joined to the entoconid, and there is greater development of the labial ridge of the anterolophulid. In MAB 0 B (Fig. 2 E), one individual is observed to possess two ridges at the lingual-anterior and labial side of the anteroconid; the anterolophulid is more developed, and in another, the metalophulid is isolated. In FS 1, an individual is noted with the metalophulid isolated from the protoconid. In MAB 3, a tooth with an incipient ectomesolophid is observed. The material from the other sites (MAB 2, MAB 0 A, MAB 11, and CBR 1) shows no significant differences. Generally, there seems to be a slight tendency for the mesolophid to elongate over time. As illustrated in Fig. 3 A, Suppl. material 4, the biometric values of M. primitivus from localities MCX 2 and MCX 3 are the smallest observed in this basin. In addition, there is a tendency for specimens to lengthen in time.</p><p>m 2 (MAB 3; Fig. 2 M – P): the lingual anterolophid may terminate at the antero-lingual border of the metaconid (2 out of 16), fail to reach the corner (10 out of 16), or be absent (4 out of 16). The anterosinusid may be small and narrow (7 out of 16) or absent (9 out of 16). The labial anterolophid is long and descends towards the base of the protoconid. The mesolophid can be categorised into several distinct forms, including long and reaching the lingual margin (2 out of 18), long without reaching the lingual margin (3 out of 18), medium (6 out of 18), short (5 out of 18), or absent (3 out of 18). The ectolophid is continuous. The ectomesolophid is absent, and the posterolophid is not connected to the entoconid.</p><p>Variability in other sites: In MCX 3 (Fig. 2 K), the lingual anterolophid is always present and longer, and the mesolophid is short or medium in length. In BC 1 (Fig. 2 L), the lingual anterolophid is always present; in one specimen, the labial anterolophid joins the labial mesocingulum, and the mesolophid is usually longer. In MAB 0 A, the mesolophid is usually longer. In MAB 0 B, the posterolophid may be connected to the entoconid. In MAB 5 (Fig. 2 Q), the anterolophid is always present. In CBR 1, the posterolophid is connected to the entoconid; in another specimen, the labial anterolophid is connected to the labial mesocingulum. The m 2 morphology of MTR 2, FS 1, MAB 3 A, MAB 11, CBR 0 B (Fig. 2 R), and CBR 0 F does not differ significantly from that of MAB 3. About the biometry (Fig. 3 B, Suppl. material 4) between the material from the different sites of this species in the Ribesalbes-Alcora Basin, the material from MCX 3 is slightly smaller, while that from MAB 11 is slightly longer, and that from BC 1 is slightly narrower. In addition, there is a slight tendency for specimens to become larger over time.</p><p>m 3 (MAB 3; Fig. 2 V – Z): The lingual anterolophid may be of medium size (7 out of 19), short (6 out of 19), or absent (6 out of 19). A small anterosinusid is present in six specimens. The labial anterolophid may be long (9 out of 18) or short (9 out of 18), descending towards the base of the protoconid (9 out of 18). The metalophulid is short and complete, while the mesolophid is absent. The mesosinusid is narrow and transverse. The posterosinusid is medium to large in size. The protoconid and the hypoconid may be separate (6 out of 18) or not (12 out of 18). The metaconid, entoconid, and posterolophid form a continuous wall along the lingual margin (10 out of 17), or the metaconid is independent (7 out of 17).</p><p>Variability in other sites: In MCX 3 (Fig. 2 S), the lingual anterolophid is shorter and the labial is longer, with the anterosinusid occurring more frequently. In BC 1 (Fig. 2 T), there is a tooth with an anterior cingulum in front of the anterolophid. In MAB 2 (Fig. 2 U), an individual is observed with a developed mesolophid. In MAB 5 (Fig. 2 AA – AB), the lingual anterolophid is long in two individuals, while the metalophulid is absent in another and the mesolophid is present in a third specimen. In CBR 1, in addition to the mesolophid, the metaconid has a ridge that contacts the lingual part of the ectolophid in one specimen. In the more modern sites of the basin, such as MAB 5 and CBR 1, the presence of mesolophids is more prevalent. The morphology of m 3 from localities MTR 2, MAB 0 A, MAB 0 B, FS 1, MAB 11, MAB 12, and CBR 2 does not differ significantly from that described in MAB 3. The comparison of the biometry (Fig. 3 C, Suppl. material 4) of the m 3 of M. primitivus from the various localities of the Ribesalbes-Alcora Basin shows that the specimen from MAB 0 B is the largest, while that from CBR 2 is the smallest (MAB 0 B). In addition, there is a slight tendency for specimens to become shorter and wider over time.</p><p>M 1 (MAB 3; Fig. 4 E – I): the anterocone may be not subdivided (1 out of 15), be slightly subdivided and the sulcus shallow (1 out of 15), or deeply subdivided and have a deeply subdivided with a platform in front of the anterocone (10 out of 15) and with even a cingulum (3 out of 15). The labial part of the anterocone may be larger (13 out of 15) or equal (2 out of 15) to the lingual one. The protolophule may be posterior to the protocone (5 out of 15), posterior almost double, formed by the labial ridge of the anteroloph or by the front ridge of the paracone (10 out of 15). The anterolophule may connect over the lingual cusp (8 out of 15) or between the two cusps of the anterocone (7 out of 15). The labial spur of the anterolophule may be short (4 of 15), incipient (7 of 15), or absent (4 of 15). The lingual mesocingulum is present but poorly developed (8 of 16) or absent (8 out of 16). The ectoloph (or paracone ridge) may be long (8 out of 16) or short (8 out of 16). The length of the mesoloph is recorded as either long and reaching the labial margin (2 out of 17), long (11 out of 17), medium (2 out of 17), or short (2 out of 17). The connection between the mesoloph and ectoloph of the paracone is categorised as either present (4 out of 18) or absent (14 out of 18). The metalophule is directed backwards, thereby reducing the posterosinus. The posterosinus may be narrow (10 out of 17) or of medium width (7 out of 17), being either deep (8 out of 15) or deep (8 out of 15) or shallow (7 out of 15).</p><p>Variability in other sites: In the M 1 of MCX 3, there is a higher prevalence of uniform size anterocone cusps, the absence of a double protolophule, the anterolophule connection with the lingual cusp, the ectoloph is absent in one specimen, and the mesoloph is long and lack of contact with the posterior crest of the paracone. In M 1 from MTR 2 (Fig. 4 A), the protolophule is posterior, the anterolophule connects to the lingual cusp, the labial crest of the anterolophule is absent, the lingual mesocingulum is rare, the mesoloph and ectoloph of the paracone never seem to be connected, and finally the metalophule usually connects to the most anterior part of the posteroloph. It may even be directed more posteriorly, even transversely. In BC 1 (Fig. 4 B, C), there are two specimens with a deeply divided anterocone without platform or cingulum, the protolophule is simple and posterior, the anterolophule connects with the lingual cusp, its labial crest is always absent, the lingual mesocingulum is less frequent, the mesoloph never connects with the ectoloph of the paracone and finally the metalophule connects with the posteroloph posterior to the hypocone in six out of fifteen cases. In MAB 0 A (Fig. 4 D), most individuals have an anterior cingulum of the anterocone, the protolophule is usually posterior, the ectoloph is long in three out of six individuals, and in another, the metalophule does not reduce the posterosinus. In FS 1, the anterocone is deeply divided and has no anterior platform or cingulum; in one individual, the protolophule is double; in another, the protolophule is double; in another, the ectoloph is absent. In MAB 5 (Fig. 4 J), the anterocone is less divided and there are no specimens with a cingulum, the proportion of material with anterocone cusps of equal size is higher, the lingual mesocingulum is less common, the protolophule is mostly posterior, the ectoloph is absent in one tooth, and there are four specimens with a metalophule that connects posteriorly but without reducing the posterosinus. In MAB 11 (Fig. 4 K), the anterocone is less divided; in another, the metalophule does not reduce the posterosinus. In CBR 1 (Fig. 4 L), the protolophule is posterior, the anterolophule touches the lingual cusp, and the metalophule does not reduce the posterosinus in three individuals and is transverse in another. In the remaining sites (MCX 4, MTR 3, BC 2, MAB 0 B, CBR 0 E, and CBR 4), no significant morphological differences were observed. About biometrics (Fig. 3 D, Suppl. material 4), the material from the study sites belongs to the L. florancei local biozone, which is generally slightly smaller in size. In addition, there is a slight tendency for specimens to become larger over time.</p><p>M 2 (MAB 5; Fig. 4 Q – T): the protolophule may be single and transverse, connected to the protocone (11 out of 14), or double with an incomplete posterior protolophule (3 out of 14). The ectoloph may be long and contact the labial side (4 out of 13), long (2 out of 13), short (4 out of 13), or absent (3 out of 13). The mesoloph is long and reaches the labial edge (4 out of 13), long without reaching the labial edge (6 out of 13), or medium length (3 out of 13). The ectoloph and the mesoloph may be connected (5 out of 13) or not (6 out of 13), with two specimens in which the ectoloph is connected to the labial border. The sinus may be directed slightly anteriorly (2 out of 16) or perpendicular (14 out of 16). The metalophule may be directed posteriorly, connecting with the posteroloph posterior to the hypocone (3 out of 13), or directed forward and connecting with the entoloph (10 out of 13). The posteroloph may be connected to the posterior part of the metaconid (8 out of 13) or not (5 out of 13). The posterosinus may be either long (8 out of 13) or short (5 out of 13).</p><p>Variability in other sites: In MCX 3 (Fig. 4 M), there is one individual with a double protolophule, another with a lingual mesocingulum, and the ectoloph is usually longer. In BC 1 (Fig. 4 N), the morphology of the protolophule is more variable, even with a double protolophule, a double protolophule with both protolophules complete in one individual, the mesoloph is short in another. It does not appear to be connected to the ectoloph. In MAB 0 A, the protolophule may be directed either anteriorly or posteriorly. In FS 1 (Fig. 4 O), there is one individual with a double protolophule and another with a metalophule attached to the posteroloph. In MAB 3 (Fig. 4 P) there are three individuals with the protolophule attached to the anteroloph anterior to the protocone another with a double protolophule, the ectoloph always present, another with a short mesoloph, and three others with a transverse metalophule. In MAB 11 (Fig. 4 U) has one specimen with an anterior protolophule and another with a transverse metalophule. In CBR 1 (Fig. 4 V), there is one individual with an incomplete anterior protolophule, an oblique sinus in three out of four specimens, another with a transverse metalophule, and another with a reduced posterosinus. No significant differences exist in MTR 1, MTR 2, MTR 3, and MAB 4. Regarding biometry (Fig. 3 E, Suppl. material 4), the examined material belonging to local biozone L. florancei tends to be smaller than that belonging to local biozone L. ellipticus . In addition, there is a slight tendency for specimens to become larger over time.</p><p>M 3 (MAB 5; Fig. 4 AC – AF): the lingual anteroloph and the protosinus may be poorly developed (2 out of 11) or absent (9 out of 11). The labial anteroloph may be short (5 out of 11) or long (6 out of 11), connecting with the paracone (7 out of 11) or not (4 out of 11). This morphology of the labial anteroloph influences the presence of a short but relatively wide anterosinus (2 out of 11), a long anterosinus (6 out of 11), or the absence of an anterosinus (3 out of 11). The metalophule (centroloph) may be connected to the neo-entoloph (3 out of 11), to the anterior ridge of the hypocone (4 out of 11), to the anterior ridge of the hypocone and protolophule (2 out of 11), to the posterior ridge of the protocone (1 out of 11) or the axioloph (1 out of 11). The sinus may be shallow (4 out of 11) or deep (7 out of 11). The mesoloph may be long (2 out of 11), short (4 out of 11), incipient (2 out of 11), or absent (3 out of 11). The neo-entoloph may be short and connected to the protocone and hypocone (8 out of 11) or absent (3 out of 11). The axioloph may be incipient (3 out of 11) or absent (8 out of 11). The posteroloph may be long and curved (10 out of 11) or straight (1 out of 11), forming a labial wall together with the metacone and the posterior wall of the paracone (7 out of 11), and may not connect with the metacone (4 out of 11).</p><p>Variability in other sites: In MCX 3 (Fig. 4 W), the labial anteroloph is shorter; in two out of eight individuals, the neo-entoloph is absent, and the protocone and hypocone are directly connected. In MTR 2 (Fig. 4 X), there is one individual without a labial anteroloph, and the metaloph seems more often connected to the hypocone. In BC 1 (Fig. 4 Y), there is one individual without a metalophule; the sinus is always deep, and the mesoloph is absent. In MAB 3 (Fig. 4 Z – AA), the metalophule disappears in some individuals, while the mesoloph is less common. There are no significant differences in FS 1, MAB 0 A, MAB 11 B, and CBR 1. Regarding biometry (Fig. 3 F, Suppl. material 4), no significant differences can be observed between the different sites in the Ribesalbes-Alcora basin. In addition, there is a slight tendency for specimens to become larger over time.</p><p>Remarks.</p><p>The genus Megacricetodon has been the focus of significant research interest due to its abundance and wide distribution during the Early and Middle Miocene (Daams and Freudenthal 1988; Lazzari and Aguilar 2007; Wessels and Reumer 2009; Maridet et al. 2011; Oliver and Peláez-Campomanes 2013, 2014, 2016; Bonilla-Salomón et al. 2021; Jovells-Vaqué and Casanovas-Vilar 2021; Čermák et al. 2023, among others). The probable origin of this genus is in Asia, with the possibility of Megacricetodon beijiangensis Maridet et al., 2011 in China (Maridet et al. 2011) or Megacricetodon dzhungaricus (Kordikova and de Bruijn, 2001) in Kazakhstan (Kordikova and de Bruijn 2001), being possibly the oldest species described so far. An alternative hypothesis proposes an origin in Turkey, with an indeterminate form found in Keseköy (Wessels et al. 2001). These localities correlate with the European MN 3 (Wessels et al. 2001; Wessels 2009; Maridet et al. 2011; Flynn and Wessels 2013). The temporal range of this genus extends to the island site of Gargano (MN 13). However, the karstic origin of the latter site means that the record is a mixture of materials of different ages, according to Freudenthal and Martín Suárez (2010). Therefore, it can be concluded that the most recent record is that cited from the MN 9 by Kälin (1999).</p><p>The species M. primitivus, in conjunction with Megacricetodon bezianensis Bulot, 1980, represents the most primitive species identified in south-western Europe (Oliver and Peláez-Campomanes 2014; Čermák et al. 2023). Furthermore, the existence of multiple species of this genus in the Iberian Peninsula dates back to the Middle Miocene, as evidenced by the coexistence of M. primitivus with other species of the same genus during this period (Sesé 2006; van der Meulen et al. 2012; Casanovas-Vilar et al. 2016). In the Early Miocene of France, M. primitivus undoubtedly coexists with other species of the same genus, including M. bezianensis and Megacricetodon aunayi Lazzari &amp; Aguilar, 2007 (Antoine et al. 2000; Ginsburg and Bulot 2000; Bulot et al. 2009).</p><p>The populations of the Calatayud-Montalbán Basin, as meticulously documented by Oliver and Peláez-Campomanes (2014), provide a detailed understanding of the morphology of M. primitivus . This understanding is defined by specific characteristics: the m 1 with a rounded and generally simple anteroconid, a short anterolophulid, and an ectomesolophid that is typically absent; the m 3 mesolophids are usually present at low frequency; the m 1 has double anteroconid, which are typically accompanied by a small anterior platform, a strong lingual mesocingulum that is present in 20 % of specimens, and a mesoloph that is usually short to medium length. This detailed morphology is comparable to that observed in the material studied here, although with some minor variations, which will be discussed in further detail below.</p><p>The material under study is distinguished from other small and medium-sized species of this genus from Europe and Turkey by its relatively smaller size, Megacricetodon andrewsi Peláez-Campomanes &amp; Daams, 2002, Megacricetodon similis (Fahlbusch, 1964), and Megacricetodon grueneri Čermák et al., 2022 (Peláez-Campomanes and Daams 2002; Čermák et al. 2023). The material under study exhibits significant distinguishing characteristics in comparison with Megacricetodon collongensis (Mein, 1958), including a higher frequency of double anteroconids, longer anterolophs / lophids, more symmetrical anterocones of M 1, and a greater prevalence of protolophs and double metalophs in M 1 and M 2. It differs from Megacricetodon debruijni Freudenthal, 1968, in that it exhibits split anteroconids on M 1 and short mesolophids. It is also distinguished from Megacricetodon freudenthali García Moreno (in Álvarez-Sierra and García Moreno, 1986), Megacricetodon lopezae García Moreno (in Álvarez-Sierra and García Moreno, 1986), Megacricetodon minor (Lartet, 1851), Megacricetodon minutus Daxner, 1967, and Megacricetodon rafaeli Daams &amp; Freudenthal, 1988 . The material under study differs from these in possessing an undivided anterocone. Furthermore, it differs from Megacricetodon tautavelensis Lazzari &amp; Aguilar, 2007 in terms of its smaller size, a higher percentage of divided anterocones, shorter mesolophs / ids, and a higher percentage of representation of crescent-shaped anteroconids. Finally, Megacricetodon hellenicus Oliver and Peláez-Campomanes, 2016 differs from the aforementioned material by having the M 1 in that it has a less deeply divided anterocone, longer mesolophs and entolophs; M 2 has a higher percentage of ectolophs; M 3 has longer entolophs; the anteroconid of m 1 is lower, and the mesolophids and lingual anterolophid of m 2 are the longest (Oliver and Peláez-Campomanes 2014, 2016).</p><p>Given the substantial number of localities in which this species is present, a comparison was made between the present material and that from the localities of the Calatayud-Montalbán Basin. The most numerous population from each of the localities, Ca (Artesilla), Cb (Vargas 1 a), and the type locality (Valtorres, MN 5, local area Db), was selected for analysis (Oliver and Peláez-Campomanes 2014).</p><p>In the M 1 s of the Calatayud-Montalbán Basin, the anterocone generally exhibits a small anterior platform; however, in a smaller proportion of specimens, this feature is absent, and in a lesser degree, the anterocones display a small cingulum or the anterocone is barely divided. In the material studied, this form is dominant only in MTR 2. In the remaining localities, the presence of this anterior platform or the cingulum is exceedingly rare. It is noteworthy that MAB 5 contains multiple specimens exhibiting a slight division of the anterocone. About the symmetry of the anterocone of M 1, the prevailing morphology in the populations used for comparisons is that in which the labial cusp is larger than the lingual cusp, except Artesilla, where the two cusps are almost of the same size. A similar phenomenon is observed in the localities of the Ribesalbes-Alcora Basin, where something similar occurs, where three sites (MCX 3, MAB 5, and CBR 1) exhibit comparable sizes between the anterocone cusps. In the type locality, the connection of the anteroloph of M 1 to the anterocone occurs at similar frequencies along the middle part of the cusps of the anterocone or on the lingual cusp, with a slightly higher frequency in the latter case.</p><p>In the Ribesalbes-Alcora Basin, except MAB 3 and MAB 5, where both morphologies are represented in equal proportions, there is a striking prevalence of the anteroloph attached to the lingual cusp in the remaining materials. The presence of the partially developed or incipient labial ridge of the anteroloph of M 1 is an infrequent occurrence in the localities with which it is compared. In contrast, it is similar in our localities, with the incipient stage slightly more common. The protoloph of M 1 in the localities used for comparisons of the Calatayud-Montalbán Basin is predominantly single and posterior, with a lesser prevalence of complete or interrupted double forms. These also occur in our material, although in MAB 3 the dominant form is that with the small unconnected ridge.</p><p>Concerning the lingual mesocingulum of M 1, the number of individuals exhibiting a robust lingual mesocingulum was documented in the populations utilised for comparisons. This character is not particularly prevalent in the material under consideration. However, when only the presence or absence of the mesocingulum-anteroloph connection is considered, the proportions are similar or even higher. The ectoloph is typically longer in the localities of the Ribesalbes-Alcora Basin than in the localities used for comparisons of the Calatayud-Montalbán Basin, and a comparable development is observed solely at the Artesilla site. In contrast, the number of connections with the mesoloph is very similar in both groups of localities. The mesoloph of M 1 demonstrates a comparable development, except for MCX 3, where they are longer. At the same time, the metalophule of M 1 is predominantly connected to the base of the posteroloph in the localities used for comparisons. In the localities of the Ribesalbes-Alcora Basin, there is material where a posteriorly directed metalophule dominates, thereby reducing the size of the posterosinus.</p><p>Regarding the development of the M 2 protolophule, in most of the populations utilised for comparisons, it is anterior, or an incomplete posterior protolophule manifests with significantly reduced frequency, and the transverse, double, or posterior forms are residual. Conversely, within the material studied, the proportions observed in the sites belonging to the L. florancei biozone are consistent with those seen in other populations, while in the sites belonging to the L. ellipticus biozone, the transverse forms predominate. The ectoloph’s length and percentage of connections with the mesoloph are comparable. Notably, the ectoloph-mesoloph junction is generally less frequent in the oldest deposits belonging to the local L. florancei biozone of the Ribesalbes-Alcora Basin. The development of the M 2 metalophule is very similar in both populations, with the anterior form dominating, although the transverse form predominates in MCX 3 and the posterior metalophule in MAB 0 A.</p><p>The morphology of the metalophule of M 3 is similar in both basins, predominantly connected to the anterior ridge of the hypocone, except for the exception of BC 1, where the majority of specimens are devoid of a metalophule.</p><p>In m 1, the anteroconid is consistently simple, although at very low frequencies, the populations employed for comparisons from the Calatayud-Montalbán Basin may be slightly divided. The labial ridge of the m 1 anterolophulid is predominantly absent in both basins, although incipient ridges are well represented in BC 1 and well-developed ridges are present in Vargas 1 A. The anterior metalophulid of m 1 is present in the majority of specimens from both basins, more frequently separated from the protoconid in the Ribesalbes-Alcora Basin than in the populations used for comparisons of the Calatayud-Montalbán Basin. The mesolophid lengths of m 1, m 2, and m 3 are similar in the localities of both basins, although in m 2 of the localities studied, there are proportionally more specimens that reach the labial edge. On the other hand, the mesolophids of m 3 occur only in the more modern deposits belonging to local biozone L. ellipticus in the Ribesalbes-Alcora Basin. The absence of ectomesolophids in m 1 and the length of the anterolophid of m 2 (short) are similar in both basins. Although in the latter case, the length of the m 2 anterolophid is shorter, the well-developed form is more abundant in the MCX 3, BC 1, and MAB 5 sites.</p><p>The specimens from the Calatayud-Montalbán and the Ribesalbes-Alcora basins studied here are similar in size (see Fig. 3). However, minor discrepancies do emerge: the m 1 is similar to those of the populations utilised for comparisons of local biozone C (Ca and Cb), though with some shorter specimens in MCX 2, MCX 3, and MTR 2; the m 2 are also more similar to the local biozone C populations, with maximum length values slightly higher than in the material under study; the m 3 the sizes are similar to those of the deposits of the same local biozone, except for one specimen found in MAB 0 B; the M 1, although similar, is generally closer to the range of the locality used for comparisons in local biozone Cb; the M 2 are identical to those of this locality, only the material from MAB 0 A is longer; and finally, regarding the M 3, there is one specimen from BC 1 and one from MAB 3 that are narrower than those from this locality, while the rest are of similar size.</p><p>In relation to the Buñol zone, the Barranco de Candel site is distinguished by the following features in comparison with our material: one specimen with a slightly bilobed anteroconid of the M 1; the anterolophid of the M 2 is typically more developed; the M 2 mesolophid is longer; the M 1 ectolophs are more prevalent; and the M 1 mesolophs are longer. Regarding biometrics, the specimens from the Barranco del Candel site are broadly similar to those studied from the Ribesalbes-Alcora Basin, although the M 1 s are slightly smaller (Adrover et al. 1987). Furthermore, the morphology of the material from the Buñol site is comparable to that of the Ribesalbes-Alcora Basin sites, with the exception of the mesolophid of m 1, which is slightly longer, and the anterolophid of m 1, which is somewhat shorter. Also, the lingual anterolophid of m 2 is more developed, the protolophule is almost double in half of the specimens, which is the same as in MAB 3, and the mesoloph of M 1 is shorter. The measurements are similar, except M 2 is slightly smaller and M 3 is slightly broader (Daams and Freudenthal 1974).</p><p>In the Morteral section (1–20 A), within the Magro Basin (including M. primitivus and some taxa in open nomenclature ( Megacricetodon sp. 1 and 2), the morphology of these taxa is similar. However, there are some differences when compared to our sites. In the more modern sites, there are representatives with a slightly doubled anteroconid of m 1 and shorter mesolophids. The m 2 exhibits a slightly shorter mesolophid, and the labial ridge of the anterolophid of M 1 is less frequently observed. The ectoloph of the M 1 is longer. In M 2, there is a reduction in the number of partially double and transverse protolophids. Furthermore, in the more modern deposits, the ectoloph is more frequently attached to the mesoloph. The metalophule of M 2 exhibits a comparable morphology, with the exception of MCX 3, where it is transverse. Biometrically, they are similar, except for Megacricetodon sp. 2, which has larger lower molars, and the upper molars are of a similar size (Ruiz-Sánchez 1999; Ruiz-Sánchez et al. 2003).</p><p>In the Vallès-Penedès Basin, the remains analysed in this study demonstrate notable similarities to those exhibited by the material found in the following sites: Can Julià 6 and Can Martí Vell I and II, Els Casots 73 and 76. However, a distinguishing feature emerges in the analysis of Can Martí Vell I, where the protolophid is consistently located posteriorly, and the mesolophid of m 1 is typically shorter. Biometrically, the material from Catalonia corresponds with the material studied here, including the Megacricetodon sp. described by Agustí (1983), for which he emphasized its larger size. While the most recent publications from this basin indicate an increase in size in the more recent deposits, similar to what occurred in the present basin, the measurements are, on average, slightly larger than those of the material studied (Jovells-Vaqué and Casanovas-Vilar 2021).</p><p>In the Loranca Basin, particularly at the La Retama site, the mesoloph of M 1 is characterised by a reduction in length, the lingual mesocingulum exhibits increased prevalence, the protolophule is posteriorly positioned, and the metalophule is anteriorly situated. In the M 2, the mesoloph is shorter; in the M 1, there is a higher prevalence of divided anteroconids. In the m 2, the lingual anterolophid is less prevalent, and the mesolophids are shorter in length. Biometrically, M 2, m 2, and m 3 are typically smaller in size (Álvarez-Sierra et al. 2006).</p><p>Therefore, the morphological and biometric study of the material of M. primitivus is in accordance with the findings of Oliver and Peláez-Campomanes (2014), which indicate that the species is highly stable over time, exhibiting intraspecific variability. Additionally, there is no evident evolutionary trend in either size or morphology. Furthermore, the distribution of dental characters varies considerably across different sites, yet they always retain a resemblance to the type of material.</p></div>	https://treatment.plazi.org/id/CA97BFB8994B5D1F8B931FF183CCAF9F	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Crespo, Vicente D.;Ríos, María;Marquina-Blasco, Rafael;Montoya, Plini	Crespo, Vicente D., Ríos, María, Marquina-Blasco, Rafael, Montoya, Plini (2025): The Early Miocene muroids (Muroidea, Rodentia) of the Ribesalbes-Alcora Basin (Spain): A thriving haven during a time of migration. Fossil Record 28 (1): 187-218, DOI: 10.3897/fr.28.138478
B8CDB4FA63D05A668306D1A62FEC49DD.text	B8CDB4FA63D05A668306D1A62FEC49DD.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Melissiodon Schaub 1925	<div><p>Genus Melissiodon Schaub, 1925</p><p>Type species.</p><p>Melissiodon dominans Schaub, 1925 . Wintershof-West. Early Miocene.</p></div>	https://treatment.plazi.org/id/B8CDB4FA63D05A668306D1A62FEC49DD	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Crespo, Vicente D.;Ríos, María;Marquina-Blasco, Rafael;Montoya, Plini	Crespo, Vicente D., Ríos, María, Marquina-Blasco, Rafael, Montoya, Plini (2025): The Early Miocene muroids (Muroidea, Rodentia) of the Ribesalbes-Alcora Basin (Spain): A thriving haven during a time of migration. Fossil Record 28 (1): 187-218, DOI: 10.3897/fr.28.138478
E62062C7CCE00F06244D9C1AE4B76A03.text	E62062C7CCE00F06244D9C1AE4B76A03.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Muroidea Illiger 1811	<div><p>Muroidea indet.</p><p>Fig. 8 Z</p><p>Localities. MAB 0 A.</p><p>Material (number of remains). MAB 0 A (1): 1 M 2.</p><p>Description. M 2 (MAB 0 A; Fig. 8 Z): small broken tooth. The protoloph is double, the anterior one being transverse and connected to the protocone, and the posterior one incomplete, without contacting the paracone. The mesoloph is double: the anterior is short and the posterior is medium, both parallel. The crest of the protocone is short and does not contact the mesoloph. The ectomesoloph is absent. The sinus is narrow and directed forward. The metaloph is anterior and connects with the entoloph. The posteroloph is long and isolated labially. There is a small posterior ridge of the hypocone, in the posterosinus.</p><p>Remarks. The tooth fragment displays characteristics consistent with those observed in Eumyarion, Pseudocricetodon, or Eucricetodon, including the presence of double mesolophs and the posterior crest of the hypocone. Given its diminutive dimensions and the presence of a double protoloph, it seems not plausible that this specimen can be attributed to the Eumyarion species identified at other localities. The Turkish fossil record is replete with small Eumyarion species, yet one m 1 has been described from the Blanquàtere 1 location, which belongs to one of these species (Aguilar et al. 2010 b). It is imprudent to draw comparisons between the two, as they represent distinct elements. However, this combination of characteristics is absent in E. intercentralis or E. microps from Keseköy (de Bruijn and Saraç 1991). A specimen from Sabuncubeli exhibits some resemblance but displays a longer mesoloph, and the two crests are fused. Additionally, it is larger in E. orhani (de Bruijn et al. 2006) . The presence of double mesolophs or the posterior crest of the hypocone has been documented in Eucricetodon aquitanicus Baudelot &amp; Bonis, 1968 from the Early Miocene (Hugueney 1999 b), and in addition, the presence of double mesolophs has been documented in Eucricetodon atavoides Freudenthal, 1996 from the early Oligocene (Freudenthal 1996; Gomes-Rodrigues et al. 2013), within the Eucricetodon genus. The species Sindemys aguilari (Lindsay, 1988) from Pakistan and certain Asian Eucricetodon forms (see plate 7.3, fig. 12; Wessels 2009; Gomes-Rodrigues et al. 2012) exhibit this or a similar configuration, despite originating from outside the European and Turkish regions.</p><p>The complicated primitive morphology of the taxon precludes its categorisation within any specific genus. If the specimen were determined to belong to Eumyarion, it would represent the earliest known appearance of this genus in the Levantine basins from the Iberian Peninsula. Conversely, if the specimen belonged to Eucricetodon, it would represent the surviving member of the genus following the absence of representatives of the European MN 3 following the Cricetid Vacuum (Hugueney 1999 b). Alternatively, it could be an exotic immigrant species from Asia or another continent during a period of widespread intercontinental migration.</p></div>	https://treatment.plazi.org/id/E62062C7CCE00F06244D9C1AE4B76A03	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Crespo, Vicente D.;Ríos, María;Marquina-Blasco, Rafael;Montoya, Plini	Crespo, Vicente D., Ríos, María, Marquina-Blasco, Rafael, Montoya, Plini (2025): The Early Miocene muroids (Muroidea, Rodentia) of the Ribesalbes-Alcora Basin (Spain): A thriving haven during a time of migration. Fossil Record 28 (1): 187-218, DOI: 10.3897/fr.28.138478
